Anatomy and Physiology! https://pressbooks.bccampus.ca/apdouglas2017 Open Textbook Wed, 30 Aug 2017 18:17:37 +0000 en-US 1.2 https://pressbooks.bccampus.ca/ https://pressbooks.bccampus.ca/apdouglas2017 10 1 https://wordpress.org/?v=4.8.1 Preface https://pressbooks.bccampus.ca/apdouglas2017/preface/ Thu, 13 Jul 2017 20:38:05 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/Preface.png 19 0 0 0 Preface2 https://pressbooks.bccampus.ca/apdouglas2017/preface2/ Thu, 13 Jul 2017 20:38:06 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/Preface2.png 20 0 0 0 01_01ab_Gross_and_Microscopic_Anatomy https://pressbooks.bccampus.ca/apdouglas2017/01_01ab_gross_and_microscopic_anatomy/ Thu, 13 Jul 2017 20:38:07 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/01_01ab_Gross_and_Microscopic_Anatomy.jpg 23 0 0 0 101_Levels_of_Org_in_Body 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20:42:40 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/810_Femur_and_Patella.jpg 288 0 0 0 818_Femur_Q_Angle https://pressbooks.bccampus.ca/apdouglas2017/818_femur_q_angle/ Thu, 13 Jul 2017 20:42:41 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/818_Femur_Q_Angle.jpg 289 0 0 0 811_Tibia_and_fibula https://pressbooks.bccampus.ca/apdouglas2017/811_tibia_and_fibula/ Thu, 13 Jul 2017 20:42:42 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/811_Tibia_and_fibula.jpg 290 0 0 0 812_Bones_of_the_Foot https://pressbooks.bccampus.ca/apdouglas2017/812_bones_of_the_foot/ Thu, 13 Jul 2017 20:42:43 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/812_Bones_of_the_Foot.jpg 291 0 0 0 footbones https://pressbooks.bccampus.ca/apdouglas2017/footbones/ Thu, 13 Jul 2017 20:42:44 +0000 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2021 0 0 9.1 https://pressbooks.bccampus.ca/apdouglas2017/chapter/9-1-classification-of-joints/9-1/ Fri, 14 Jul 2017 22:46:07 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/9.1.png 2967 2114 0 0 10.2 https://pressbooks.bccampus.ca/apdouglas2017/chapter/10-2-skeletal-muscle/10-2/ Fri, 14 Jul 2017 22:47:47 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/10.2.png 2969 2147 0 0 18.1 https://pressbooks.bccampus.ca/apdouglas2017/chapter/18-1-an-overview-of-blood/18-1/ Fri, 14 Jul 2017 22:55:40 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/18.1.png 2971 2200 0 0 19.1 https://pressbooks.bccampus.ca/apdouglas2017/chapter/19-1-heart-anatomy/19-1/ Fri, 14 Jul 2017 22:56:58 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/19.1.png 2973 2237 0 0 20.5 https://pressbooks.bccampus.ca/apdouglas2017/chapter/20-5-circulatory-pathways/20-5/ Fri, 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https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-1-organs-and-structures-of-the-respiratory-system/22-1/ Fri, 14 Jul 2017 23:04:07 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/22.1.png 2984 2373 0 0 22.5 https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-5-transport-of-gases/22-5/ Fri, 14 Jul 2017 23:05:34 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/22.5.png 2986 2393 0 0 1.5 https://pressbooks.bccampus.ca/apdouglas2017/chapter/1-5-homeostasis-2/1-5-2/ Mon, 17 Jul 2017 18:51:49 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/1.5-1.png 2988 2428 0 0 2.2 https://pressbooks.bccampus.ca/apdouglas2017/chapter/2-2-chemical-bonds-2/2-2/ Mon, 17 Jul 2017 18:53:33 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/2.2.png 2990 2442 0 0 3.5 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https://pressbooks.bccampus.ca/apdouglas2017/chapter/13-2-the-central-nervous-system/13-2/ Mon, 17 Jul 2017 19:03:24 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/13.2.png 3000 2522 0 0 14.1 hearing and balance https://pressbooks.bccampus.ca/apdouglas2017/chapter/14-1-sensory-perception/14-1-hearing-and-balance/ Mon, 17 Jul 2017 19:05:40 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/14.1-hearing-and-balance.png 3002 2553 0 0 14.1 taste and smell https://pressbooks.bccampus.ca/apdouglas2017/chapter/14-1-sensory-perception/14-1-taste-and-smell/ Mon, 17 Jul 2017 19:06:12 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/14.1-taste-and-smell.png 3003 2553 0 0 14.1 vision https://pressbooks.bccampus.ca/apdouglas2017/chapter/14-1-sensory-perception/14-1-vision/ Mon, 17 Jul 2017 19:08:32 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/14.1-vision.png 3004 2553 0 0 14.3 https://pressbooks.bccampus.ca/apdouglas2017/chapter/14-3-motor-responses/14-3/ Mon, 17 Jul 2017 19:10:13 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/14.3.png 3007 2568 0 0 15.1 https://pressbooks.bccampus.ca/apdouglas2017/chapter/15-1-divisions-of-the-autonomic-nervous-system-1203/15-1/ Mon, 17 Jul 2017 19:13:12 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/15.1.png 3009 2574 0 0 17.1 https://pressbooks.bccampus.ca/apdouglas2017/chapter/17-1-an-overview-of-the-endocrine-system/17-1/ Mon, 17 Jul 2017 19:14:29 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/17.1.png 3011 2588 0 0 22.5 https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-5-transport-of-gases-2/22-5-2/ Mon, 17 Jul 2017 19:16:17 +0000 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https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/24.1.png 3021 2637 0 0 25.1 https://pressbooks.bccampus.ca/apdouglas2017/chapter/25-1-physical-characteristics-of-urine/25-1/ Mon, 17 Jul 2017 19:37:36 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/25.1.png 3023 2713 0 0 25.5 https://pressbooks.bccampus.ca/apdouglas2017/chapter/25-5-physiology-of-urine-formation/25-5/ Mon, 17 Jul 2017 19:38:49 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/25.5.png 3025 2732 0 0 27.1 https://pressbooks.bccampus.ca/apdouglas2017/chapter/27-1-anatomy-and-physiology-of-the-male-reproductive-system/27-1/ Mon, 17 Jul 2017 19:42:44 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/27.1.png 3027 2773 0 0 27.2 https://pressbooks.bccampus.ca/apdouglas2017/chapter/27-2-anatomy-and-physiology-of-the-female-reproductive-system/27-2/ Mon, 17 Jul 2017 19:43:44 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/27.2.png 3029 2783 0 0 28.1 https://pressbooks.bccampus.ca/apdouglas2017/chapter/28-1-fertilization/28-1/ Mon, 17 Jul 2017 19:44:43 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/28.1.png 3031 2789 0 0 28.2 https://pressbooks.bccampus.ca/apdouglas2017/chapter/28-2-embryonic-development/28-2/ Mon, 17 Jul 2017 19:45:31 +0000 https://pressbooks.bccampus.ca/apdouglas2017/wp-content/uploads/sites/221/2017/07/28.2.png 3033 2803 0 0 Appendix https://pressbooks.bccampus.ca/apdouglas2017/back-matter/appendix/ Mon, 10 Jul 2017 16:28:25 +0000 https://pressbooks.bccampus.ca/apdouglas2017/back-matter/appendix/ 7 0 1 0 1.2 Structural Organization of the Human Body https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1840 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1840

Learning Objectives

[caption id="" align="alignright" width="550"]This illustration shows biological organization as a pyramid. The chemical level is at the apex of the pyramid where atoms bond to form molecules with three dimensional structures. An example is shown with two white hydrogen atoms bonding to a red oxygen atom to create water. The next level down on the pyramid is the cellular level, as illustrated with a long, tapered, smooth muscle cell. At this level, a variety of molecules combine to form the interior fluid and organelles of a body cell. The next level down is the tissue level. A community of similar cells forms body tissue. The example given here is a section of smooth muscle tissue, which contains many smooth muscle cells closely bound side by side. The next level down is the organ level, as illustrated with the bladder and urethra. The bladder contains smooth muscle while the urethra contains skeletal muscle. These are both examples of muscle tissues. The next level down is the organ system level, as illustrated by the entire urinary system containing the kidney, ureters, bladder and urethra. At this level, two or more organs work closely together to perform the functions of a body system. At the base of the pyramid is the organismal level illustrated with a woman drinking water. At this level, many organ systems work harmoniously together to perform the functions of an independent organism. Figure 1. Levels of Structural Organization of the Human Body. The organization of the body often is discussed in terms of six distinct levels of increasing complexity, from the smallest chemical building blocks to a unique human organism.[/caption] By the end of this section, you will be able to:
  • Describe the structure of the human body in terms of six levels of organization
  • List the eleven organ systems of the human body and identify at least one organ and one major function of each
Before you begin to study the different structures and functions of the human body, it is helpful to consider its basic architecture; that is, how its smallest parts are assembled into larger structures. It is convenient to consider the structures of the body in terms of fundamental levels of organization that increase in complexity: subatomic particles, atoms, molecules, organelles, cells, tissues, organs, organ systems, organisms and biosphere (Figure 1).

The Levels of Organization

To study the chemical level of organization, scientists consider the simplest building blocks of matter: subatomic particles, atoms and molecules. All matter in the universe is composed of one or more unique pure substances called elements, familiar examples of which are hydrogen, oxygen, carbon, nitrogen, calcium, and iron. The smallest unit of any of these pure substances (elements) is an atom. Atoms are made up of subatomic particles such as the proton, electron and neutron. Two or more atoms combine to form a molecule, such as the water molecules, proteins, and sugars found in living things. Molecules are the chemical building blocks of all body structures. A cell is the smallest independently functioning unit of a living organism. Even bacteria, which are extremely small, independently-living organisms, have a cellular structure. Each bacterium is a single cell. All living structures of human anatomy contain cells, and almost all functions of human physiology are performed in cells or are initiated by cells. A human cell typically consists of flexible membranes that enclose cytoplasm, a water-based cellular fluid together with a variety of tiny functioning units called organelles. In humans, as in all organisms, cells perform all functions of life. A tissue is a group of many similar cells (though sometimes composed of a few related types) that work together to perform a specific function. An organ is an anatomically distinct structure of the body composed of two or more tissue types. Each organ performs one or more specific physiological functions. An organ system is a group of organs that work together to perform major functions or meet physiological needs of the body. [caption id="" align="alignright" width="500"]The lymphatic system returns fluid to the blood and defends against pathogens. The lymphatic system includes the thymus in the chest, the spleen in the abdomen, the lymphatic vessels that spread throughout the body, and the lymph nodes distributed along the lymphatic vessels. The respiratory system removes carbon dioxide from the body and delivers oxygen to the blood. The respiratory system includes the nasal passages, the trachea, and the lungs. The digestive system processes food for use by the body and removes wastes from undigested food. The digestive system includes the stomach, the liver, the gall bladder (connected to the liver), the large intestine, and the small intestine. The urinary system controls water balance in the body and removes and excretes waste from the blood. The urinary system includes the kidneys and the urinary bladder. The reproductive system of males and females produce sex hormones and gametes. The male reproductive system is specialized to deliver gametes to the female while the female reproductive system is specialized to support the embryo and fetus until birth and produce milk for the infant after birth. The male reproductive system includes the two testes within the scrotum as well as the epididymis which wraps around each testis. The female reproductive system includes the mammary glands within the breasts and the ovaries and uterus within the pelvic cavity. Figure 3. Organ Systems of the Human Body (continued). Organs that work together are grouped into organ systems.[/caption] [caption id="" align="alignright" width="500"]This illustration shows eight silhouettes of a human female, each showing the components of a different organ system. The integumentary system encloses internal body structures and is the site of many sensory receptors. The integumentary system includes the hair, skin, and nails. The skeletal system supports the body and, along with the muscular system, enables movement. The skeletal system includes cartilage, such as that at the tip of the nose, as well as the bones and joints. The muscular system enables movement, along with the skeletal system, but also helps to maintain body temperature. The muscular system includes skeletal muscles, as well as tendons that connect skeletal muscles to bones. The nervous system detects and processes sensory information and activates bodily responses. The nervous system includes the brain, spinal cord, and peripheral nerves, such as those located in the limbs. The endocrine system secretes hormones and regulates bodily processes. The endocrine system includes the pituitary gland in the brain, the thyroid gland in the throat, the pancreas in the abdomen, the adrenal glands on top of the kidneys, and the testes in the scrotum of males as well as the ovaries in the pelvic region of females. The cardiovascular system delivers oxygen and nutrients to the tissues as well as equalizes temperature in the body. The cardiovascular system includes the heart and blood vessels. Figure 2. Organ Systems of the Human Body. Organs that work together are grouped into organ systems.[/caption] This book covers eleven distinct organ systems in the human body (Figure 2 and Figure 3). Assigning organs to organ systems can be imprecise since organs that “belong” to one system can also have functions integral to another system. In fact, most organs contribute to more than one system.
The organism level is the highest level of organization. An organism is a living being that has a cellular structure and that can independently perform all physiologic functions necessary for life. In multicellular organisms, including humans, all cells, tissues, organs, and organ systems of the body work together to maintain the life and health of the organism.
]]> 1840 1834 2 0 1.4 Requirements for Human Life https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1847 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1847

Learning Objectives

By the end of this section, you will be able to:
  • Discuss the role of oxygen and nutrients in maintaining human survival
  • Explain why extreme heat and extreme cold threaten human survival
  • Explain how the pressure exerted by gases and fluids influences human survival

Humans have been adapting to life on Earth for at least the past 200,000 years. Earth and its atmosphere have provided us with air to breathe, water to drink, and food to eat, but these are not the only requirements for survival. Although you may rarely think about it, you also cannot live outside of a certain range of temperature and pressure that the surface of our planet and its atmosphere provides. The next sections explore these four requirements of life.

Oxygen

Atmospheric air is only about 20 percent oxygen, but that oxygen is a key component of the chemical reactions that keep the body alive, including the reactions that produce ATP. Brain cells are especially sensitive to lack of oxygen because of their requirement for a high-and-steady production of ATP. Brain damage is likely within five minutes without oxygen, and death is likely within ten minutes.

Nutrients

A nutrient is a substance in foods and beverages that is essential to human survival. The three basic classes of nutrients are water, the energy-yielding and body-building nutrients, and the micronutrients (vitamins and minerals).

The most critical nutrient is water. Depending on the environmental temperature and our state of health, we may be able to survive for only a few days without water. The body’s functional chemicals are dissolved and transported in water, and the chemical reactions of life take place in water. Moreover, water is the largest component of cells, blood, and the fluid between cells, and water makes up about 70 percent of an adult’s body mass. Water also helps regulate our internal temperature and cushions, protects, and lubricates joints and many other body structures.

The energy-yielding nutrients are primarily carbohydrates and lipids, while proteins mainly supply the amino acids that are the building blocks of the body itself. You ingest these in plant and animal foods and beverages, and the digestive system breaks them down into molecules small enough to be absorbed. The breakdown products of carbohydrates and lipids can then be used in the metabolic processes that convert them to ATP. Although you might feel as if you are starving after missing a single meal, you can survive without consuming the energy-yielding nutrients for at least several weeks.

Water and the energy-yielding nutrients are also referred to as macronutrients because the body needs them in large amounts. In contrast, micronutrients are vitamins and minerals. These elements and compounds participate in many essential chemical reactions and processes, such as nerve impulses, and some, such as calcium, also contribute to the body’s structure. Your body can store some of the micronutrients in its tissues, and draw on those reserves if you fail to consume them in your diet for a few days or weeks. Some others micronutrients, such as vitamin C and most of the B vitamins, are water-soluble and cannot be stored, so you need to consume them every day or two.

Narrow Range of Temperature

You have probably seen news stories about athletes who died of heat stroke, or hikers who died of exposure to cold. Such deaths occur because the chemical reactions upon which the body depends can only take place within a narrow range of body temperature, from just below to just above 37°C (98.6°F). When body temperature rises well above or drops well below normal, certain proteins (enzymes) that facilitate chemical reactions lose their normal structure and their ability to function and the chemical reactions of metabolism cannot proceed.

That said, the body can respond effectively to short-term exposure to heat (Figure 1) or cold. One of the body’s responses to heat is, of course, sweating. As sweat evaporates from skin, it removes some thermal energy from the body, cooling it. Adequate water (from the extracellular fluid in the body) is necessary to produce sweat, so adequate fluid intake is essential to balance that loss during the sweat response. Not surprisingly, the sweat response is much less effective in a humid environment because the air is already saturated with water. Thus, the sweat on the skin’s surface is not able to evaporate, and internal body temperature can get dangerously high.
[caption id="" align="aligncenter" width="420"]This photo shows two white-clad men riding camels through a sparse desert. Two canvas tents are visible in the background. Figure 1. Extreme Heat. Humans adapt to some degree to repeated exposure to high temperatures. (credit: McKay Savage/flickr)[/caption]

The body can also respond effectively to short-term exposure to cold. One response to cold is shivering, which is random muscle movement that generates heat. Another response is increased breakdown of stored energy to generate heat. When that energy reserve is depleted, however, and the core temperature begins to drop significantly, red blood cells will lose their ability to give up oxygen, denying the brain of this critical component of ATP production. This lack of oxygen can cause confusion, lethargy, and eventually loss of consciousness and death. The body responds to cold by reducing blood circulation to the extremities, the hands and feet, in order to prevent blood from cooling there and so that the body’s core can stay warm. Even when core body temperature remains stable, however, tissues exposed to severe cold, especially the fingers and toes, can develop frostbite when blood flow to the extremities has been much reduced. This form of tissue damage can be permanent and lead to gangrene, requiring amputation of the affected region.

Narrow Range of Atmospheric Pressure

Pressure is a force exerted by a substance that is in contact with another substance. Atmospheric pressure is pressure exerted by the mixture of gases (primarily nitrogen and oxygen) in the Earth’s atmosphere. Although you may not perceive it, atmospheric pressure is constantly pressing down on your body. This pressure keeps gases within your body, such as the gaseous nitrogen in body fluids, dissolved. If you were suddenly ejected from a space ship above Earth’s atmosphere, you would go from a situation of normal pressure to one of very low pressure. The pressure of the nitrogen gas in your blood would be much higher than the pressure of nitrogen in the space surrounding your body. As a result, the nitrogen gas in your blood would expand, forming bubbles that could block blood vessels and even cause cells to break apart.

Atmospheric pressure does more than just keep blood gases dissolved. Your ability to breathe—that is, to take in oxygen and release carbon dioxide—also depends upon a precise atmospheric pressure. Altitude sickness occurs in part because the atmosphere at high altitudes exerts less pressure, reducing the exchange of these gases, and causing shortness of breath, confusion, headache, lethargy, and nausea. Mountain climbers carry oxygen to reduce the effects of both low oxygen levels and low barometric pressure at higher altitudes (Figure 2).

[caption id="" align="aligncenter" width="380"]This photo shows Mount Everest as seen from a distance. It is a large, pyramid-shaped, craggy peak with many smaller snow-covered peaks in the foreground. The peak of Mount Everest is partially occluded by clouds. Figure 2. Harsh Conditions. Climbers on Mount Everest must accommodate extreme cold, low oxygen levels, and low barometric pressure in an environment hostile to human life. (credit: Melanie Ko/flickr)[/caption]
]]> 1847 1834 4 0 1.6 Anatomical Terminology https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1858 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1858

Learning Objectives

By the end of this section, you will be able to:
  • Demonstrate the anatomical position
  • Describe the human body using directional and regional terms
  • Identify three planes most commonly used in the study of anatomy
  • Distinguish between the posterior (dorsal) and the anterior (ventral) body cavities, identifying their subdivisions and representative organs found in each
  • Describe serous membrane and explain its function

Anatomists and health care providers use terminology that can be bewildering to the uninitiated. However, the purpose of this language is not to confuse, but rather to increase precision and reduce medical errors. For example, is a scar “above the wrist” located on the forearm two or three inches away from the hand? Or is it at the base of the hand? Is it on the palm-side or back-side? By using precise anatomical terminology, we eliminate ambiguity. Anatomical terms derive from ancient Greek and Latin words. Because these languages are no longer used in everyday conversation, the meaning of their words does not change.

Anatomical terms are made up of roots, prefixes, and suffixes. The root of a term often refers to an organ, tissue, or condition, whereas the prefix or suffix often describes the root. For example, in the disorder hypertension, the prefix “hyper-” means “high” or “over,” and the root word “tension” refers to pressure, so the word “hypertension” refers to abnormally high blood pressure.

[caption id="" align="alignright" width="550"]This illustration shows an anterior and posterior view of the human body. The cranial region encompasses the upper part of the head while the facial region encompasses the lower half of the head beginning below the ears. The eyes are referred to as the ocular region. The cheeks are referred to as the buccal region. The ears are referred to as the auricle or otic region. The nose is referred to as the nasal region. The chin is referred to as the mental region. The neck is referred to as the cervical region. The trunk of the body contains, from superior to inferior, the thoracic region encompassing the chest, the mammary region encompassing each breast, the abdominal region encompassing the stomach area, the coxal region encompassing the belt line, and the pubic region encompassing the area above the genitals. The umbilicus, or naval, is located at the center of the abdomen. The pelvis and legs contain, from superior to inferior, the inguinal or groin region between the legs and the genitals, the pubic region surrounding the genitals, the femoral region encompassing the thighs, the patellar region encompassing the knee, the crural region encompassing the lower leg, the tarsal region encompassing the ankle, the pedal region encompassing the foot and the digital/phalangeal region encompassing the toes. The great toe is referred to as the hallux. The regions of the upper limbs, from superior to inferior, are the axillary region encompassing the armpit, the brachial region encompassing the upper arm, the antecubital region encompassing the front of the elbow, the antebrachial region encompassing the forearm, the carpal region encompassing the wrist, the palmar region encompassing the palm, and the digital/phalangeal region encompassing the fingers. The thumb is referred to as the pollux. The posterior view contains, from superior to inferior, the cervical region encompassing the neck, the dorsal region encompassing the upper back and the lumbar region encompassing the lower back. The regions of the back of the arms, from superior to inferior, include the cervical region encompassing the neck, acromial region encompassing the shoulder, the brachial region encompassing the upper arm, the olecranal region encompassing the back of the elbow, the antebrachial region encompasses the back of the arm, and the manual region encompassing the palm of the hand. The posterior regions of the legs, from superior to inferior, include the gluteal region encompassing the buttocks, the femoral region encompassing the thigh, the popliteus region encompassing the back of the knee, the sural region encompassing the back of the lower leg, and the plantar region encompassing the sole of the foot. Some regions are combined into larger regions. These include the trunk, which is a combination of the thoracic, mammary, abdominal, naval, and coxal regions. The cephalic region is a combination of all of the head regions. The upper limb region is a combination of all of the arm regions. The lower limb region is a combination of all of the leg regions. Figure 1. Regions of the Human Body. The human body is shown in anatomical position in an (a) anterior view and a (b) posterior view. The regions of the body are labeled in boldface.[/caption]

Anatomical Position

To further increase precision, anatomists standardize the way in which they view the body. Just as maps are normally oriented with north at the top, the standard body “map,” or anatomical position, is that of the body standing upright, with the feet at shoulder width and parallel, toes forward. The upper limbs are held out to each side, and the palms of the hands face forward as illustrated in Figure 1. Using this standard position reduces confusion. It does not matter how the body being described is oriented, the terms are used as if it is in anatomical position. For example, a scar in the “anterior (front) carpal (wrist) region” would be present on the palm side of the wrist. The term “anterior” would be used even if the hand were palm down on a table.
A body that is lying down is described as either prone or supine. Prone describes a face-down orientation, and supine describes a face up orientation. These terms are sometimes used in describing the position of the body during specific physical examinations or surgical procedures.

Regional Terms

The human body’s numerous regions have specific terms to help increase precision (see Figure 1). Notice that the term “brachium” or “arm” is reserved for the “upper arm” and “antebrachium” or “forearm” is used rather than “lower arm.” Similarly, “femur” or “thigh” is correct, and “leg” or “crus” is reserved for the portion of the lower limb between the knee and the ankle. You will be able to describe the body’s regions using the terms from the figure.

 

Directional Terms

[caption id="" align="alignright" width="450"]This illustration shows two diagrams: one of a side view of a female and the other of an anterior view of a female. Each diagram shows directional terms using double-sided arrows. The cranial-distal arrow runs vertically behind the torso and lower abdomen. The cranial arrow is pointing toward the head while the caudal arrow is pointing toward the tail bone. The posterior/anterior arrow is running horizontally through the back and chest. The posterior or dorsal arrow is pointing toward the back while the anterior, or ventral arrow, is pointing toward the abdomen. On the anterior view, the proximal/distal arrow is on the right arm. The proximal arrow is pointing up toward the shoulder while the distal arrow is pointing down toward the hand. The lateral-medial arrow is a horizontal arrow on the abdomen. The medial arrow is pointing toward the navel while the lateral arrow is pointing away from the body to the right. Right refers to the right side of the woman’s body from her perspective while left refers to the left side of the woman’s body from her perspective. Figure 2. Directional Terms Applied to the Human Body. Paired directional terms are shown as applied to the human body.[/caption] Certain directional anatomical terms appear throughout this and any other anatomy textbook (Figure 2). These terms are essential for describing the relative locations of different body structures. For instance, an anatomist might describe one band of tissue as “inferior to” another or a physician might describe a tumor as “superficial to” a deeper body structure. Commit these terms to memory to avoid confusion when you are studying or describing the locations of particular body parts.
  • Anterior (or ventral) Describes the front or direction toward the front of the body. The toes are anterior to the foot.
  • Posterior (or dorsal) Describes the back or direction toward the back of the body. The popliteus is posterior to the patella.
  • Superior (or cranial) describes a position above or higher than another part of the body proper. The orbits are superior to the oris.
  • Inferior (or caudal) describes a position below or lower than another part of the body proper; near or toward the tail (in humans, the coccyx, or lowest part of the spinal column). The pelvis is inferior to the abdomen.
  • Lateral describes the side or direction toward the side of the body. The thumb (pollex) is lateral to the digits.
  • Medial describes the middle or direction toward the middle of the body. The hallux is the medial toe.
  • Proximal describes a position in a limb that is nearer to the point of attachment or the trunk of the body. The brachium is proximal to the antebrachium.
  • Distal describes a position in a limb that is farther from the point of attachment or the trunk of the body. The crus is distal to the femur.
  • Superficial describes a position closer to the surface of the body. The skin is superficial to the bones.
  • Deep describes a position farther from the surface of the body. The brain is deep to the skull.

Body Planes

[caption id="" align="alignleft" width="350"]This illustration shows a female viewed from her right, front side. The anatomical planes are depicted as blue rectangles passing through the woman’s body. The frontal or coronal plane enters through the right side of the body, passes through the body, and exits from the left side. It divides the body into front (anterior) and back (posterior) halves. The sagittal plane enters through the back and emerges through the front of the body. It divides the body into right and left halves. The transverse plane passes through the body perpendicular to the frontal and sagittal planes. This plane is a cross section which divides the body into upper and lower halves. Figure 3. Planes of the Body. The three planes most commonly used in anatomical and medical imaging are the sagittal, frontal (or coronal), and transverse plane.[/caption]

A section is a two-dimensional surface of a three-dimensional structure that has been cut. Modern medical imaging devices enable clinicians to obtain “virtual sections” of living bodies. We call these scans. Body sections and scans can be correctly interpreted, however, only if the viewer understands the plane along which the section was made. A plane is an imaginary two-dimensional surface that passes through the body. There are three planes commonly referred to in anatomy and medicine, as illustrated in Figure 3.

  • The sagittal plane is the plane that divides the body or an organ vertically into right and left sides. If this vertical plane runs directly down the middle of the body, it is called the midsagittal or median plane. If it divides the body into unequal right and left sides, it is called a parasagittal plane or less commonly a longitudinal section.
  • The frontal plane is the plane that divides the body or an organ into an anterior (front) portion and a posterior (rear) portion. The frontal plane is often referred to as a coronal plane. (“Corona” is Latin for “crown.”)
  • The transverse plane is the plane that divides the body or organ horizontally into upper and lower portions. Transverse planes produce images referred to as cross sections.

Body Cavities and Serous Membranes

The body maintains its internal organization by means of membranes, sheaths, and other structures that separate compartments. The dorsal (posterior) cavity and the ventral (anterior) cavity are the largest body compartments (Figure 4). These cavities contain and protect delicate internal organs, and the ventral cavity allows for significant changes in the size and shape of the organs as they perform their functions. The lungs, heart, stomach, and intestines, for example, can expand and contract without distorting other tissues or disrupting the activity of nearby organs.

[caption id="" align="alignright" width="550"]This illustration shows a lateral and anterior view of the body and highlights the body cavities with different colors. The cranial cavity is a large, bean-shaped cavity filling most of the upper skull where the brain is located. The vertebral cavity is a very narrow, thread-like cavity running from the cranial cavity down the entire length of the spinal cord. Together the cranial cavity and vertebral cavity can be referred to as the dorsal body cavity. The thoracic cavity consists of three cavities that fill the interior area of the chest. The two pleural cavities are situated on both sides of the body, anterior to the spine and lateral to the breastbone. The superior mediastinum is a wedge-shaped cavity located between the superior regions of the two thoracic cavities. The pericardial cavity within the mediastinum is located at the center of the chest below the superior mediastinum. The pericardial cavity roughly outlines the shape of the heart. The diaphragm divides the thoracic and the abdominal cavities. The abdominal cavity occupies the entire lower half of the trunk, anterior to the spine. Just under the abdominal cavity, anterior to the buttocks, is the pelvic cavity. The pelvic cavity is funnel shaped and is located inferior and anterior to the abdominal cavity. Together the abdominal and pelvic cavity can be referred to as the abdominopelvic cavity while the thoracic, abdominal, and pelvic cavities together can be referred to as the ventral body cavity. Figure 4. Dorsal and Ventral Body Cavities. The ventral cavity includes the thoracic and abdominopelvic cavities and their subdivisions. The dorsal cavity includes the cranial and spinal cavities.[/caption]

Subdivisions of the Posterior (Dorsal) and Anterior (Ventral) Cavities

The posterior (dorsal) and anterior (ventral) cavities are each subdivided into smaller cavities. In the posterior (dorsal) cavity, the cranial cavity houses the brain, and the spinal cavity (or vertebral cavity) encloses the spinal cord. Just as the brain and spinal cord make up a continuous, uninterrupted structure, the cranial and spinal cavities that house them are also continuous. The brain and spinal cord are protected by the bones of the skull and vertebral column and by cerebrospinal fluid, a colorless fluid produced by the brain, which cushions the brain and spinal cord within the posterior (dorsal) cavity.

 

The anterior (ventral) cavity has two main subdivisions: the thoracic cavity and the abdominopelvic cavity (see Figure 4). The thoracic cavity is the more superior subdivision of the anterior cavity, and it is enclosed by the rib cage. The thoracic cavity contains the lungs and the heart, which is located in the mediastinum. The diaphragm forms the floor of the thoracic cavity and separates it from the more inferior abdominopelvic cavity. The abdominopelvic cavity is the largest cavity in the body. Although no membrane physically divides the abdominopelvic cavity, it can be useful to distinguish between the abdominal cavity, the division that houses the digestive organs, and the pelvic cavity, the division that houses the organs of reproduction.

[caption id="" align="alignleft" width="565"]This illustration has two parts. Part A shows the abdominopelvic regions. These regions divide the abdomen into nine squares. The upper right square is the right hypochondriac region and contains the base of the right ribs. The upper left square is the left hypochondriac region and contains the base of the left ribs. The epigastric region is the upper central square and contains the bottom edge of the liver as well as the upper areas of the stomach. The diaphragm curves like an upside down U over these three regions. The central right region is called the right lumbar region and contains the ascending colon and the right edge of the small intestines. The central square contains the transverse colon and the upper regions of the small intestines. The left lumbar region contains the left edge of the transverse colon and the left edge of the small intestine. The lower right square is the right iliac region and contains the right pelvic bones and the ascending colon. The lower left square is the left iliac region and contains the left pelvic bone and the lower left regions of the small intestine. The lower central square contains the bottom of the pubic bones, upper regions of the bladder and the lower region of the small intestine. Part B shows four abdominopelvic quadrants. The right upper quadrant (RUQ) includes the lower right ribs, right side of the liver, and right side of the transverse colon. The left upper quadrant (LUQ) includes the lower left ribs, stomach, and upper left area of the transverse colon. The right lower quadrant (RLQ) includes the right half of the small intestines, ascending colon, right pelvic bone and upper right area of the bladder. The left lower quadrant (LLQ) contains the left half of the small intestine and left pelvic bone. Figure 5. Regions and Quadrants of the Peritoneal Cavity. There are (a) nine abdominal regions and (b) four abdominal quadrants in the peritoneal cavity.[/caption]

Abdominal Regions and Quadrants

To promote clear communication, for instance about the location of a patient’s abdominal pain or a suspicious mass, health care providers typically divide up the cavity into either nine regions or four quadrants (Figure 5).

The more detailed regional approach subdivides the cavity with one horizontal line immediately inferior to the ribs and one immediately superior to the pelvis, and two vertical lines drawn as if dropped from the midpoint of each clavicle (collarbone). There are nine resulting regions. The simpler quadrants approach, which is more commonly used in medicine, subdivides the cavity with one horizontal and one vertical line that intersect at the patient’s umbilicus (navel).

Membranes of the Anterior (Ventral) Body Cavity

A serous membrane (also referred to a serosa) is one of the thin membranes that cover the walls and organs in the thoracic and abdominopelvic cavities. The parietal layers of the membranes line the walls of the body cavity (pariet- refers to a cavity wall). The visceral layer of the membrane covers the organs (the viscera). Between the parietal and visceral layers is a very thin, fluid-filled serous space, or cavity (Figure 6).

[caption id="" align="alignright" width="420"]This diagram shows the pericardium on the left next to an analogy of a hand punching a balloon on the right. The pericardium is a two-layered sac that surrounds the entire heart except where the blood vessels emerge on the heart’s superior side. The pericardium has two layers because it folds over itself in the shape of the letter U. The inner layer that borders the heart is the visceral pericardium while the outer layer is the parietal pericardium. The space between the two layers is called the pericardial cavity. The heart sits in the cavity much like a fist punching into a balloon. The balloon surrounds the lower part of the fist with a two-layered sac, with the top of the balloon, where it contacts the fist, being analogous to the visceral pericardium. The bottom of the balloon, where it is tied off, is analogous to the parietal pericardium. The air within the balloon is analogous to the pericardial cavity. Figure 6. Serous Membrane. Serous membrane lines the pericardial cavity and reflects back to cover the heart—much the same way that an underinflated balloon would form two layers surrounding a fist.[/caption] There are three serous cavities and their associated membranes. The pleura is the serous membrane that surrounds the lungs in the pleural cavity; the pericardium is the serous membrane that surrounds the heart in the pericardial cavity; and the peritoneum is the serous membrane that surrounds several organs in the abdominopelvic cavity.The serous membranes form fluid-filled sacs, or cavities, that are meant to cushion and reduce friction on internal organs when they move, such as when the lungs inflate or the heart beats. Both the parietal and visceral serosa secrete the thin, slippery serous fluid located within the serous cavities. The pleural cavity reduces friction between the lungs and the body wall. Likewise, the pericardial cavity reduces friction between the heart and the wall of the pericardium. The peritoneal cavity reduces friction between the abdominal and pelvic organs and the body wall. Therefore, serous membranes provide additional protection to the viscera they enclose by reducing friction that could lead to inflammation of the organs.

]]> 1858 1834 6 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1903 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1903 [caption id="" align="aligncenter" width="500"]This figure shows a double helix. Figure 1. Human DNA. Human DNA is described as a double helix that resembles a molecular spiral staircase. In humans the DNA is organized into 46 chromosomes.[/caption]

Chapter Objectives

After studying this chapter, you will be able to:
  • Describe the fundamental composition of matter
  • Identify the three subatomic particles
  • Identify the four most abundant elements in the body
  • Explain the relationship between an atom’s number of electrons and its relative stability
  • Distinguish between ionic bonds, covalent bonds, and hydrogen bonds
  • Explain how energy is invested, stored, and released via chemical reactions, particularly those reactions that are critical to life
  • Explain the importance of the inorganic compounds that contribute to life, such as water, salts, acids, and bases
  • Compare and contrast the four important classes of organic (carbon-based) compounds—proteins, carbohydrates, lipids and nucleic acids—according to their composition and functional importance to human life
The smallest, most fundamental material components of the human body are basic chemical elements. In fact, chemicals called nucleotide bases are the foundation of the genetic code with the instructions on how to build and maintain the human body from conception through old age. There are about three billion of these base pairs in human DNA. Human chemistry includes organic molecules (carbon-based) and biochemicals (those produced by the body). Human chemistry also includes elements. In fact, life cannot exist without many of the elements that are part of the earth. All of the elements that contribute to chemical reactions, to the transformation of energy, and to electrical activity and muscle contraction—elements that include phosphorus, carbon, sodium, and calcium, to name a few—originated in stars. These elements, in turn, can form both the inorganic and organic chemical compounds important to life, including, for example, water, glucose, and proteins. This chapter begins by examining elements and how the structures of atoms, the basic units of matter, determine the characteristics of elements by the number of protons, neutrons, and electrons in the atoms. The chapter then builds the framework of life from there.]]> 1903 1901 1 0 2.1 Elements and Atoms: the Building Blocks of Matter https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1910 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1910

Learning Objectives

By the end of this section, you will be able to:
  • Discuss the relationships between matter, mass, elements, compounds, atoms, and subatomic particles
  • Distinguish between atomic number and mass number
  • Identify the key distinction between isotopes of the same element
  • Explain how electrons occupy electron shells and their contribution to an atom’s relative stability

The substance of the universe—from a grain of sand to a star—is called matter. Scientists define matter as anything that occupies space and has mass. An object’s mass and its weight are related concepts, but not quite the same. An object’s mass is the amount of matter contained in the object, and the object’s mass is the same whether that object is on Earth or in the zero-gravity environment of outer space. An object’s weight, on the other hand, is its mass as affected by the pull of gravity. Where gravity strongly pulls on an object’s mass its weight is greater than it is where gravity is less strong. An object of a certain mass weighs less on the moon, for example, than it does on Earth because the gravity of the moon is less than that of Earth. In other words, weight is variable, and is influenced by gravity. A piece of cheese that weighs a pound on Earth weighs only a few ounces on the moon.

Elements and Compounds

All matter in the natural world is composed of one or more of the 92 fundamental substances called elements. An element is a pure substance that is distinguished from all other matter by the fact that it cannot be created or broken down by ordinary chemical means. While your body can assemble many of the chemical compounds needed for life from their constituent elements, it cannot make elements. They must come from the environment.

A familiar example of an element that you must take in is calcium (Ca++). Calcium is essential to the human body; it is absorbed and used for a number of processes, including strengthening bones. When you consume dairy products your digestive system breaks down the food into components small enough to cross into the bloodstream. Among these is calcium, which, because it is an element, cannot be broken down further. The elemental calcium in cheese, therefore, is the same as the calcium that forms your bones. Some other elements you might be familiar with are oxygen, sodium, and iron. The elements in the human body are shown in Figure 1, beginning with the most abundant: oxygen (O), carbon (C), hydrogen (H), and nitrogen (N). Each element’s name can be replaced by a one- or two-letter symbol; you will become familiar with some of these during this course. All the elements in your body are derived from the foods you eat and the air you breathe.
[caption id="" align="alignleft" width="585"]This figure shows a human body with the percentage of the main elements in the body, in the left panel. In the right panel, a table lists the elements and the percentages in the body. Figure 1. Elements of the Human Body. The main elements that compose the human body are shown from most abundant to least abundant.[/caption]

In nature, elements rarely occur alone. Instead, they combine to form compounds. A compound is a substance composed of two or more elements joined by chemical bonds. For example, the compound glucose is an important body fuel. It is always composed of the same three elements: carbon, hydrogen, and oxygen. Moreover, the elements that make up any given compound always occur in the same relative amounts. In glucose, there are always six carbon and six oxygen units for every twelve hydrogen units. But what, exactly, are these “units” of elements?

Atoms and Subatomic Particles

An atom is the smallest quantity of an element that retains the unique properties of that element. In other words, an atom of hydrogen is a unit of hydrogen—the smallest amount of hydrogen that can exist. As you might guess, atoms are almost unfathomably small. The period at the end of this sentence is millions of atoms wide.

Atomic Structure and Energy

Atoms are made up of even smaller subatomic particles, three types of which are important: the proton, neutron, and electron. The number of positively-charged protons and non-charged (“neutral”) neutrons, gives mass to the atom, and the number of each in the nucleus of the atom determine the element. The number of negatively-charged electrons that “spin” around the nucleus at close to the speed of light equals the number of protons. An electron has about 1/2000th the mass of a proton or neutron.

Figure 2 shows two models that can help you imagine the structure of an atom—in this case, helium (He). In the planetary model, helium’s two electrons are shown circling the nucleus in a fixed orbit depicted as a ring. Although this model is helpful in visualizing atomic structure, in reality, electrons do not travel in fixed orbits, but whiz around the nucleus erratically in a so-called electron cloud.

[caption id="" align="alignleft" width="285"]The top panel of this figure shows two electrons orbiting around the nucleus of a Helium atom. The bottom panel of this figure shows a cloud of electrons surrounding the nucleus of a Helium atom. Figure 2. Two Models of Atomic Structure. (a) In the planetary model, the electrons of helium are shown in fixed orbits, depicted as rings, at a precise distance from the nucleus, somewhat like planets orbiting the sun. (b) In the electron cloud model, the electrons of carbon are shown in the variety of locations they would have at different distances from the nucleus over time.[/caption]

An atom’s protons and electrons carry electrical charges. Protons, with their positive charge, are designated p+. Electrons, which have a negative charge, are designated e–. An atom’s neutrons have no charge: they are electrically neutral. Just as a magnet sticks to a steel refrigerator because their opposite charges attract, the positively charged protons attract the negatively charged electrons. This mutual attraction gives the atom some structural stability. The attraction by the positively charged nucleus helps keep electrons from straying far. The number of protons and electrons within a neutral atom are equal, thus, the atom’s overall charge is balanced.

Atomic Number and Mass Number

An atom of carbon is unique to carbon, but a proton of carbon is not. One proton is the same as another, whether it is found in an atom of carbon, sodium (Na), or iron (Fe). The same is true for neutrons and electrons. So, what gives an element its distinctive properties—what makes carbon so different from sodium or iron? The answer is the unique quantity of protons each contains. Carbon by definition is an element whose atoms contain six protons. No other element has exactly six protons in its atoms. Moreover, all atoms of carbon, whether found in your liver or in a lump of coal, contain six protons. Thus, the atomic number, which is the number of protons in the nucleus of the atom, identifies the element. Because an atom usually has the same number of electrons as protons, the atomic number identifies the usual number of electrons as well.

In their most common form, many elements also contain the same number of neutrons as protons. The most common form of carbon, for example, has six neutrons as well as six protons, for a total of 12 subatomic particles in its nucleus. An element’s mass number is the sum of the number of protons and neutrons in its nucleus. So the most common form of carbon’s mass number is 12. (Electrons have so little mass that they do not appreciably contribute to the mass of an atom.) Carbon is a relatively light element. Uranium (U), in contrast, has a mass number of 238 and is referred to as a heavy metal. Its atomic number is 92 (it has 92 protons) but it contains 146 neutrons; it has the most mass of all the naturally occurring elements.

The periodic table of the elements, shown below in Figure 3, is a chart identifying the 92 elements found in nature, as well as several larger, unstable elements discovered experimentally. The elements are arranged in order of their atomic number, with hydrogen and helium at the top of the table, and the more massive elements below. The periodic table is a useful device because for each element, it identifies the chemical symbol, the atomic number, and the mass number, while organizing elements according to their propensity to react with other elements. The number of protons and electrons in an element are equal. The number of protons and neutrons may be equal for some elements, but are not equal for all.

[caption id="" align="alignleft" width="650"]This figure shows the periodic table. Figure 3. The Periodic Table of the Elements. (credit: R.A. Dragoset, A. Musgrove, C.W. Clark, W.C. Martin)[/caption]
[caption id="" align="alignright" width="120"]QR Code representing a URL Visit this website to view the periodic table.[/caption]   In the periodic table of the elements, elements in a single column have the same number of electrons that can participate in a chemical reaction. These electrons are known as “valence electrons.” For example, the elements in the first column all have a single valence electron, an electron that can be “donated” in a chemical reaction with another atom. What is the meaning of a mass number shown in parentheses?

Isotopes

Although each element has a unique number of protons, it can exist as different isotopes. An isotope is one of the different forms of an element, distinguished from one another by different numbers of neutrons. The standard isotope of carbon is 12C, commonly called carbon twelve. 12C has six protons and six neutrons, for a mass number of twelve. All of the isotopes of carbon have the same number of protons; therefore, 13C has seven neutrons, and 14C has eight neutrons. The different isotopes of an element can also be indicated with the mass number hyphenated (for example, C-12 instead of 12C). Hydrogen has three common isotopes, shown in Figure 4.

[caption id="" align="alignright" width="380"]This figure shows the three isotopes of hydrogen: hydrogen, deuterium, and tritium. Figure 4. Isotopes of Hydrogen. Protium, designated 1H, has one proton and no neutrons. It is by far the most abundant isotope of hydrogen in nature. Deuterium, designated 2H, has one proton and one neutron. Tritium, designated 3H, has two neutrons.[/caption]

An isotope that contains more than the usual number of neutrons is referred to as a heavy isotope. An example is 14C. Heavy isotopes tend to be unstable, and unstable isotopes are radioactive. A radioactive isotope is an isotope whose nucleus readily decays, giving off subatomic particles and electromagnetic energy. Different radioactive isotopes (also called radioisotopes) differ in their half-life, the time it takes for half of any size sample of an isotope to decay. For example, the half-life of tritium—a radioisotope of hydrogen—is about 12 years, indicating it takes 12 years for half of the tritium nuclei in a sample to decay. Excessive exposure to radioactive isotopes can damage human cells and even cause cancer and birth defects, but when exposure is controlled, some radioactive isotopes can be useful in medicine. For more information, see the Career Connections.

Radioisotopes emit subatomic particles that can be detected and tracked by imaging technologies. One of the most advanced uses of radioisotopes in medicine is the positron emission tomography (PET) scanner, which detects the activity in the body of a very small injection of radioactive glucose, the simple sugar that cells use for energy. The PET camera reveals to the medical team which of the patient’s tissues are taking up the most glucose. Thus, the most metabolically active tissues show up as bright “hot spots” on the images (Figure 5). PET can reveal some cancerous masses because cancer cells consume glucose at a high rate to fuel their rapid reproduction.

The Behavior of Electrons

In the human body, atoms do not exist as independent entities. Rather, they are constantly reacting with other atoms to form and to break down more complex substances. To fully understand anatomy and physiology you must grasp how atoms participate in such reactions. The key is understanding the behavior of electrons.

Although electrons do not follow rigid orbits a set distance away from the atom’s nucleus, they do tend to stay within certain regions of space called electron shells. An electron shell is a layer of electrons that encircle the nucleus at a distinct energy level.

The atoms of the elements found in the human body have from one to five electron shells, and all electron shells hold eight electrons except the first shell, which can only hold two. This configuration of electron shells is the same for all atoms. The precise number of shells depends on the number of electrons in the atom. Hydrogen and helium have just one and two electrons, respectively. If you take a look at the periodic table of the elements, you will notice that hydrogen and helium are placed alone on either sides of the top row; they are the only elements that have just one electron shell (Figure 6). A second shell is necessary to hold the electrons in all elements larger than hydrogen and helium.

Lithium (Li), whose atomic number is 3, has three electrons. Two of these fill the first electron shell, and the third spills over into a second shell. The second electron shell can accommodate as many as eight electrons. Carbon, with its six electrons, entirely fills its first shell, and half-fills its second. With ten electrons, neon (Ne) entirely fills its two electron shells. Again, a look at the periodic table reveals that all of the elements in the second row, from lithium to neon, have just two electron shells. Atoms with more than ten electrons require more than two shells. These elements occupy the third and subsequent rows of the periodic table.
[caption id="" align="alignright" width="420"]This four panel figure shows four different atoms with the electrons in orbit around the nucleus. Figure 6. Electron Shells. Electrons orbit the atomic nucleus at distinct levels of energy called electron shells. (a) With one electron, hydrogen only half-fills its electron shell. Helium also has a single shell, but its two electrons completely fill it. (b) The electrons of carbon completely fill its first electron shell, but only half-fills its second. (c) Neon, an element that does not occur in the body, has 10 electrons, filling both of its electron shells.[/caption]

The factor that most strongly governs the tendency of an atom to participate in chemical reactions is the number of electrons in its valence shell. A valence shell is an atom’s outermost electron shell. If the valence shell is full, the atom is stable; meaning its electrons are unlikely to be pulled away from the nucleus by the electrical charge of other atoms. If the valence shell is not full, the atom is reactive; meaning it will tend to react with other atoms in ways that make the valence shell full. Consider hydrogen, with its one electron only half-filling its valence shell. This single electron is likely to be drawn into relationships with the atoms of other elements, so that hydrogen’s single valence shell can be stabilized.

All atoms (except hydrogen and helium with their single electron shells) are most stable when there are exactly eight electrons in their valence shell. This principle is referred to as the octet rule, and it states that an atom will give up, gain, or share electrons with another atom so that it ends up with eight electrons in its own valence shell. For example, oxygen, with six electrons in its valence shell, is likely to react with other atoms in a way that results in the addition of two electrons to oxygen’s valence shell, bringing the number to eight. When two hydrogen atoms each share their single electron with oxygen, covalent bonds are formed, resulting in a molecule of water, H2O.

In nature, atoms of one element tend to join with atoms of other elements in characteristic ways. For example, carbon commonly fills its valence shell by linking up with four atoms of hydrogen. In so doing, the two elements form the simplest of organic molecules, methane, which also is one of the most abundant and stable carbon-containing compounds on Earth. As stated above, another example is water; oxygen needs two electrons to fill its valence shell. It commonly interacts with two atoms of hydrogen, forming H2O. Incidentally, the name “hydrogen” reflects its contribution to water (hydro- = “water”; -gen = “maker”). Thus, hydrogen is the “water maker.”

]]> 1910 1901 2 0 3.3 The Nucleus and DNA Replication https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1961 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1961

By the end of this section, you will be able to:
  • Describe the structure and features of the nuclear membrane
  • List the contents of the nucleus
  • Explain the organization of the DNA molecule within the nucleus
  • Describe the process of DNA replication
[caption id="" align="alignright" width="382"]This figure shows the structure of the nucleus. The nucleolus is inside the nucleus, surrounded by the chromatin and covered by the nuclear envelope. Figure 1. The Nucleus. The nucleus is the control center of the cell. The nucleus of living cells contains the genetic material that determines the entire structure and function of that cell.[/caption] The nucleus is the largest and most prominent of a cell’s organelles (Figure 1). The nucleus is generally considered the control center of the cell because it stores all of the genetic instructions for manufacturing proteins. Interestingly, some cells in the body, such as muscle cells, contain more than one nucleus (Figure 2), which is known as multinucleated. Other cells, such as mammalian red blood cells (RBCs), do not contain nuclei at all. RBCs eject their nuclei as they mature, making space for the large numbers of hemoglobin molecules that carry oxygen throughout the body (Figure 3). Without nuclei, the life span of RBCs is short, and so the body must produce new ones constantly.    
[caption id="" align="alignleft" width="380"]This micrograph shows a muscle cell with multiple nuclei. Figure 2. Multinucleate Muscle Cell. Unlike cardiac muscle cells and smooth muscle cells, which have a single nucleus, a skeletal muscle cell contains many nuclei, and is referred to as “multinucleated.” These muscle cells are long and fibrous (often referred to as muscle fibers). During development, many smaller cells fuse to form a mature muscle fiber. The nuclei of the fused cells are conserved in the mature cell, thus imparting a multinucleate characteristic to mature muscle cells. LM × 104.3. Visit the University of Michigan MedScope to view the sample further. http://141.214.65.171/Histology/Basic%20Tissues/Muscle/058thin_HISTO_83X.svs/view.apml (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]
[caption id="" align="alignright" width="317"]This set of micrographs shows a red blood cell extruding its nucleus. In the left panel, the nucleus is partially extruded from the red blood cell and in the right panel, the nucleus is completely extruded from the cell. Figure 3. Red Blood Cell Extruding Its Nucleus. Mature red blood cells lack a nucleus. As they mature, erythroblasts extrude their nucleus, making room for more hemoglobin. The two panels here show an erythroblast before and after ejecting its nucleus, respectively.[/caption]  
 
           

Inside the nucleus lies the blueprint that dictates everything a cell will do and all of the products it will make. This information is stored within DNA. The nucleus sends “commands” to the cell via molecular messengers that translate the information from DNA. Each cell in your body (with the exception of germ cells) contains the complete set of your DNA. When a cell divides, the DNA must be duplicated so that the each new cell receives a full complement of DNA. The following section will explore the structure of the nucleus and its contents, as well as the process of DNA replication.

Organization of the Nucleus and Its DNA

Like most other cellular organelles, the nucleus is surrounded by a membrane called the nuclear envelope. This membranous covering consists of two adjacent lipid bilayers with a thin fluid space in between them. Spanning these two bilayers are nuclear pores. A nuclear pore is a tiny passageway for the passage of proteins, RNA, and solutes between the nucleus and the cytoplasm. Proteins called pore complexes lining the nuclear pores regulate the passage of materials into and out of the nucleus.

[caption id="" align="alignleft" width="420"]This diagram shows the macrostructure of DNA. A chromosome and its component chromatin are shown to expand into nucleosomes with histones, which further unravel into a DNA helix and finally into a DNA ladder. Figure 4. DNA Macrostructure. Strands of DNA are wrapped around supporting histones. These proteins are increasingly bundled and condensed into chromatin, which is packed tightly into chromosomes when the cell is ready to divide.[/caption] Inside the nuclear envelope is a gel-like nucleoplasm with solutes that include the building blocks of nucleic acids. There also can be a dark-staining mass often visible under a simple light microscope, called a nucleolus (plural = nucleoli). The nucleolus is a region of the nucleus that is responsible for manufacturing the RNA necessary for construction of ribosomes. Once synthesized, newly made ribosomal subunits exit the cell’s nucleus through the nuclear pores.

The genetic instructions that are used to build and maintain an organism are arranged in an orderly manner in strands of DNA. Within the nucleus are threads of chromatin composed of DNA and associated proteins (Figure 4). Along the chromatin threads, the DNA is wrapped around a set of histone proteins. A nucleosome is a single, wrapped DNA-histone complex. Multiple nucleosomes along the entire molecule of DNA appear like a beaded necklace, in which the string is the DNA and the beads are the associated histones. When a cell is in the process of division, the chromatin condenses into chromosomes, so that the DNA can be safely transported to the “daughter cells.” The chromosome is composed of DNA and proteins; it is the condensed form of chromatin. It is estimated that humans have almost 22,000 genes distributed on 46 chromosomes.

[caption id="" align="alignleft" width="450"]This figure shows the DNA double helix on the top left panel. The different nucleotides are color-coded. In the top right panel, the interaction between the nucleotides through the hydrogen bonds and the location of the sugar-phosphate backbone is shown. In the bottom panel, the structure of a nucleotide is described in detail. Figure 5. Molecular Structure of DNA. The DNA double helix is composed of two complementary strands. The strands are bonded together via their nitrogenous base pairs using hydrogen bonds.[/caption]

]]> 1961 1936 4 0 3.4 Protein Synthesis https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1967 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1967

Learning Objectives

By the end of this section, you will be able to:
  • Explain how the genetic code stored within DNA determines the protein that will form
  • Describe the process of transcription
  • Describe the process of translation
  • Discuss the function of ribosomes

It was mentioned earlier that DNA provides a “blueprint” for the cell structure and physiology. This refers to the fact that DNA contains the information necessary for the cell to build one very important type of molecule: the protein. Most structural components of the cell are made up, at least in part, by proteins and virtually all the functions that a cell carries out are completed with the help of proteins. One of the most important classes of proteins is enzymes, which help speed up necessary biochemical reactions that take place inside the cell. Some of these critical biochemical reactions include building larger molecules from smaller components (such as occurs during DNA replication or synthesis of microtubules) and breaking down larger molecules into smaller components (such as when harvesting chemical energy from nutrient molecules). Whatever the cellular process may be, it is almost sure to involve proteins. Just as the cell’s genome describes its full complement of DNA, a cell’s proteome is its full complement of proteins. Protein synthesis begins with genes. A gene is a functional segment of DNA that provides the genetic information necessary to build a protein. Each particular gene provides the code necessary to construct a particular protein. Gene expression, which transforms the information coded in a gene to a final gene product, ultimately dictates the structure and function of a cell by determining which proteins are made.

The interpretation of genes works in the following way. Recall that proteins are polymers, or chains, of many amino acid building blocks. The sequence of bases in a gene (that is, its sequence of A, T, C, G nucleotides) translates to an amino acid sequence. A triplet is a section of three DNA bases in a row that codes for a specific amino acid. Similar to the way in which the three-letter code d-o-g signals the image of a dog, the three-letter DNA base code signals the use of a particular amino acid. For example, the DNA triplet CAC (cytosine, adenine, and cytosine) specifies the amino acid valine. Therefore, a gene, which is composed of multiple triplets in a unique sequence, provides the code to build an entire protein, with multiple amino acids in the proper sequence (Figure 1). The mechanism by which cells turn the DNA code into a protein product is a two-step process, with an RNA molecule as the intermediate.

[caption id="" align="aligncenter" width="400"]This diagram shows the translation of RNA into proteins. A DNA template strand is shown to become an RNA strand through transcription. Then the RNA strand undergoes translation and becomes proteins. Figure 1. The Genetic Code. DNA holds all of the genetic information necessary to build a cell’s proteins. The nucleotide sequence of a gene is ultimately translated into an amino acid sequence of the gene’s corresponding protein.[/caption]

From DNA to RNA: Transcription

DNA is housed within the nucleus, and protein synthesis takes place in the cytoplasm, thus there must be some sort of intermediate messenger that leaves the nucleus and manages protein synthesis. This intermediate messenger is messenger RNA (mRNA), a single-stranded nucleic acid that carries a copy of the genetic code for a single gene out of the nucleus and into the cytoplasm where it is used to produce proteins.

There are several different types of RNA, each having different functions in the cell. The structure of RNA is similar to DNA with a few small exceptions. For one thing, unlike DNA, most types of RNA, including mRNA, are single-stranded and contain no complementary strand. Second, the ribose sugar in RNA contains an additional oxygen atom compared with DNA. Finally, instead of the base thymine, RNA contains the base uracil. This means that adenine will always pair up with uracil during the protein synthesis process.

Gene expression begins with the process called transcription, which is the synthesis of a strand of mRNA that is complementary to the gene of interest. This process is called transcription because the mRNA is like a transcript, or copy, of the gene’s DNA code. Transcription begins in a fashion somewhat like DNA replication, in that a region of DNA unwinds and the two strands separate, however, only that small portion of the DNA will be split apart. The triplets within the gene on this section of the DNA molecule are used as the template to transcribe the complementary strand of RNA (Figure 2). A codon is a three-base sequence of mRNA, so-called because they directly encode amino acids. Like DNA replication, there are three stages to transcription: initiation, elongation, and termination.

[caption id="" align="aligncenter" width="550"]In this diagram, RNA polymerase is shown transcribing a DNA template strand into its corresponding RNA transcript. Figure 2. Transcription: from DNA to mRNA. In the first of the two stages of making protein from DNA, a gene on the DNA molecule is transcribed into a complementary mRNA molecule.[/caption]

Stage 1: Initiation. A region at the beginning of the gene called a promoter—a particular sequence of nucleotides—triggers the start of transcription.

Stage 2: Elongation. Transcription starts when RNA polymerase unwinds the DNA segment. One strand, referred to as the coding strand, becomes the template with the genes to be coded. The polymerase then aligns the correct nucleic acid (A, C, G, or U) with its complementary base on the coding strand of DNA. RNA polymerase is an enzyme that adds new nucleotides to a growing strand of RNA. This process builds a strand of mRNA.

Stage 3: Termination. When the polymerase has reached the end of the gene, one of three specific triplets (UAA, UAG, or UGA) codes a “stop” signal, which triggers the enzymes to terminate transcription and release the mRNA transcript.

Before the mRNA molecule leaves the nucleus and proceeds to protein synthesis, it is modified in a number of ways. For this reason, it is often called a pre-mRNA at this stage. For example, your DNA, and thus complementary mRNA, contains long regions called non-coding regions that do not code for amino acids. Their function is still a mystery, but the process called splicing removes these non-coding regions from the pre-mRNA transcript (Figure 3). A spliceosome—a structure composed of various proteins and other molecules—attaches to the mRNA and “splices” or cuts out the non-coding regions. The removed segment of the transcript is called an intron. The remaining exons are pasted together. An exon is a segment of RNA that remains after splicing. Interestingly, some introns that are removed from mRNA are not always non-coding. When different coding regions of mRNA are spliced out, different variations of the protein will eventually result, with differences in structure and function. This process results in a much larger variety of possible proteins and protein functions. When the mRNA transcript is ready, it travels out of the nucleus and into the cytoplasm.

[caption id="" align="aligncenter" width="320"]In this diagram, a pre-mRNA transcript is shown in the top of a flowchart. This pre-mRNA transcript contains introns and exons. In the next step, the intron is in a structure called the spliceosome. In the last step, the intron is shown separated from the spliced RNA. Figure 3. Splicing DNA. In the nucleus, a structure called a spliceosome cuts out introns (noncoding regions) within a pre-mRNA transcript and reconnects the exons.[/caption]

From RNA to Protein: Translation

Like translating a book from one language into another, the codons on a strand of mRNA must be translated into the amino acid alphabet of proteins. Translation is the process of synthesizing a chain of amino acids called a polypeptide. Translation requires two major aids: first, a “translator,” the molecule that will conduct the translation, and second, a substrate on which the mRNA strand is translated into a new protein, like the translator’s “desk.” Both of these requirements are fulfilled by other types of RNA. The substrate on which translation takes place is the ribosome.

Remember that many of a cell’s ribosomes are found associated with the rough ER, and carry out the synthesis of proteins destined for the Golgi apparatus. Ribosomal RNA (rRNA) is a type of RNA that, together with proteins, composes the structure of the ribosome. Ribosomes exist in the cytoplasm as two distinct components, a small and a large subunit. When an mRNA molecule is ready to be translated, the two subunits come together and attach to the mRNA. The ribosome provides a substrate for translation, bringing together and aligning the mRNA molecule with the molecular “translators” that must decipher its code.

The other major requirement for protein synthesis is the translator molecules that physically “read” the mRNA codons. Transfer RNA (tRNA) is a type of RNA that ferries the appropriate corresponding amino acids to the ribosome, and attaches each new amino acid to the last, building the polypeptide chain one-by-one. Thus tRNA transfers specific amino acids from the cytoplasm to a growing polypeptide. The tRNA molecules must be able to recognize the codons on mRNA and match them with the correct amino acid. The tRNA is modified for this function. On one end of its structure is a binding site for a specific amino acid. On the other end is a base sequence that matches the codon specifying its particular amino acid. This sequence of three bases on the tRNA molecule is called an anticodon. For example, a tRNA responsible for shuttling the amino acid glycine contains a binding site for glycine on one end. On the other end it contains an anticodon that complements the glycine codon (GGA is a codon for glycine, and so the tRNAs anticodon would read CCU). Equipped with its particular cargo and matching anticodon, a tRNA molecule can read its recognized mRNA codon and bring the corresponding amino acid to the growing chain (Figure 4).

[caption id="" align="aligncenter" width="280"]The top part of this figure shows a large ribosomal subunit coming into contact with the mRNA that already has the small ribosomal subunit attached. A tRNA and an anticodon are in proximity. In the second panel, the tRNA also binds to the same site as the ribosomal subunits. In the bottom panel, a polypeptide chain is shown emerging from the complex. Figure 4. Translation from RNA to Protein. During translation, the mRNA transcript is “read” by a functional complex consisting of the ribosome and tRNA molecules. tRNAs bring the appropriate amino acids in sequence to the growing polypeptide chain by matching their anti-codons with codons on the mRNA strand.[/caption]

Much like the processes of DNA replication and transcription, translation consists of three main stages: initiation, elongation, and termination. Initiation takes place with the binding of a ribosome to an mRNA transcript. The elongation stage involves the recognition of a tRNA anticodon with the next mRNA codon in the sequence. Once the anticodon and codon sequences are bound (remember, they are complementary base pairs), the tRNA presents its amino acid cargo and the growing polypeptide strand is attached to this next amino acid. This attachment takes place with the assistance of various enzymes and requires energy. The tRNA molecule then releases the mRNA strand, the mRNA strand shifts one codon over in the ribosome, and the next appropriate tRNA arrives with its matching anticodon. This process continues until the final codon on the mRNA is reached which provides a “stop” message that signals termination of translation and triggers the release of the complete, newly synthesized protein. Thus, a gene within the DNA molecule is transcribed into mRNA, which is then translated into a protein product (Figure 5).

[caption id="" align="aligncenter" width="280"]This figure shows a schematic of a cell where transcription from DNA to mRNA takes place inside the nucleus and translation from mRNA to protein takes place in the cytoplasm. Figure 5. From DNA to Protein: Transcription through Translation. Transcription within the cell nucleus produces an mRNA molecule, which is modified and then sent into the cytoplasm for translation. The transcript is decoded into a protein with the help of a ribosome and tRNA molecules.[/caption]

Commonly, an mRNA transcription will be translated simultaneously by several adjacent ribosomes. This increases the efficiency of protein synthesis. A single ribosome might translate an mRNA molecule in approximately one minute; so multiple ribosomes aboard a single transcript could produce multiple times the number of the same protein in the same minute. A polyribosome is a string of ribosomes translating a single mRNA strand.

Check out the Khan Academy nucleus and ribosomes section to find out more!

]]> 1967 1936 5 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1979 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1979 [caption id="" align="alignright" width="600"]This micrograph shows tissue surrounding several empty spaces. The epithelial tissue occurs at the border between the rest of the tissue and the empty spaces. The normal epithelium is composed of rectangular-shaped cells neatly organized side by side. Dark purple nuclei are clear at the bottom of the epithelial cells, where they attach to the rest of the tissue. The abnormal epithelium appears as a tangled area of purple nuclei, much thicker than the normal epithelium although no distinct cells are discernible. Figure 1. Micrograph of Cervical Tissue. This figure is a view of the regular architecture of normal tissue contrasted with the irregular arrangement of cancerous cells. (credit: “Haymanj”/Wikimedia Commons)[/caption]

Chapter Objectives

After studying this chapter, you will be able to:
  • Identify the main tissue types and discuss their roles in the human body
  • Identify the four types of tissue membranes and the characteristics of each that make them functional
  • Explain the functions of various epithelial tissues and how their forms enable their functions
  • Explain the functions of various connective tissues and how their forms enable their functions
  • Describe the characteristics of muscle tissue and how these enable function
  • Discuss the characteristics of nervous tissue and how these enable information processing and control of muscular and glandular activities
The body contains at least 200 distinct cell types. These cells contain essentially the same internal structures yet they vary enormously in shape and function. The different types of cells are not randomly distributed throughout the body; rather they occur in organized layers, a level of organization referred to as tissue. The micrograph that opens this chapter shows the high degree of organization among different types of cells in the tissue of the cervix. You can also see how that organization breaks down when cancer takes over the regular mitotic functioning of a cell. The variety in shape reflects the many different roles that cells fulfill in your body. The human body starts as a single cell at fertilization. As this fertilized egg divides, it gives rise to trillions of cells, each built from the same blueprint, but organizing into tissues and becoming irreversibly committed to a developmental pathway.]]> 1979 1977 1 0 Appendix https://pressbooks.bccampus.ca/apdouglas2017/?post_type=back-matter&p=2826 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=back-matter&p=2826 This is where you can add appendices or other back matter.

]]> 2826 0 2 0 Glossary https://pressbooks.bccampus.ca/apdouglas2017/back-matter/glossary/ Wed, 02 Aug 2017 23:05:17 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=back-matter&p=2827

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broad depression located on the posterio scapula, inferior to the spine
abdaominopelvic cavity
division of the anterior (ventral) cavity that houses the abdominal and pelvic viscera
abduct
move away from midline in the sagittal plane
abduction
movement in the coronal plane that moves a limb laterally away from the body; spreading of the fingers
abductor
moves the bone away from the midline
abductor digiti minimi
muscle that abducts the little finger
adductor pollicis
muscle that adducts the thumb
abductor pollicis brevis
muscle that abducts the thumb
abductor pollicis longus
muscle that inserts into the first metacarpal
ABO blood group
blood-type classification based on the presence or absence of A and B glycoproteins on the erythrocyte membrane surface
acclimatization
process of adjustment that the respiratory system makes due to chronic exposure to high altitudes
acetabular labrum
lip of fibrocartilage that surrounds outer margin of the acetabulum on the hip bone
acetabulum
large, cup-shaped cavity located on the lateral side of the hip bone; formed by the junction of the ilium, pubis, and ischium portions of the hip bone
acetylcholine (ACh)
neurotransmitter that binds at a motor end-plate to trigger depolarization
acetylcholine (ACh)
neurotransmitter that binds at a motor end-plate to trigger depolarization
acid
building block of proteins; characterized by an amino and carboxyl functional groups and a variable side-chain
acromial process
acromion of the scapula
acromioclavicular joint
articulation between the acromion of the scapula and the acromial end of the clavicle
acromion
flattened bony process that extends laterally from the scapular spine to form the bony tip of the shoulder
acrosome
cap-like vesicle located at the anterior-most region of a sperm that is rich with lysosomal enzymes capable of digesting the protective layers surrounding the oocyte
acrosomal reaction
release of digestive enzymes by sperm that enables them to burrow through the corona radiata and penetrate the zona pellucida of an oocyte prior to fertilization
actin
protein that makes up most of the thin myofilaments in a sarcomere muscle fiber
actin
protein that makes up most of the thin myofilaments in a sarcomere muscle fiber
action potential
change in voltage of a cell membrane in response to a stimulus that results in transmission of an electrical signal; unique to neurons and muscle fibers
action potential
change in voltage of a cell membrane in response to a stimulus that results in transmission of an electrical signal; unique to neurons and muscle fibers
activation energy
amount of energy greater than the energy contained in the reactants, which must be overcome for a reaction to proceed
active immunity
immunity developed from an individual’s own immune system
active transport
form of transport across the cell membrane that requires the input of cellular energy
acute inflammation
inflammation occurring for a limited time period; rapidly developing
acute mountain sickness (AMS)
condition that occurs a result of acute exposure to high altitude due to a low partial pressure of oxygen
adaptive immune response
relatively slow but very specific and effective immune response controlled by lymphocytes
adductor
moves the bone toward the midline
adductor brevis
muscle that adducts and medially rotates the thigh
adductor longus
muscle that adducts, medially rotates, and flexes the thigh
adductor magnus
muscle with an anterior fascicle that adducts, medially rotates and flexes the thigh, and a posterior fascicle that assists in thigh extension
adenosine triphosphate (ATP)
nucleotide containing ribose and an adenine base that is essential in energy transfer
adipocytes
lipid storage cells
adipose tissue
specialized areolar tissue rich in stored fat
adductor tubercle
small, bony bump located on the superior aspect of the medial epicondyle of the femur
adduction
movement in the coronal plane that moves a limb medially toward or across the midline of the body; bringing fingers together
aerobic respiration
production of ATP in the presence of oxygen
afterload
force the ventricles must develop to effectively pump blood against the resistance in the vessels
agglutination
clustering of cells into masses linked by antibodies
agonist
(also, prime mover) muscle whose contraction is responsible for producing a particular motion
agranular leukocytes
leukocytes with few granules in their cytoplasm; specifically, monocytes, lymphocytes, and NK cells
ala
(plural = alae) small, flaring structure of a nostril that forms the lateral side of the nares
alar cartilage
cartilage that supports the apex of the nose and helps shape the nares; it is connected to the septal cartilage and connective tissue of the alae
albinism
genetic disorder that affects the skin, in which there is no melanin production
albumin
most abundant plasma protein, accounting for most of the osmotic pressure of plasma
alveolar dead space
air space within alveoli that are unable to participate in gas exchange
alveolar duct
small tube that leads from the terminal bronchiole to the respiratory bronchiole and is the point of attachment for alveoli
alveolar macrophage
immune system cell of the alveolus that removes debris and pathogens
alveolar pore
opening that allows airflow between neighboring alveoli
alveolar process of the mandible
upper border of mandibular body that contains the lower teeth
alveolar process of the maxilla
curved, inferior margin of the maxilla that supports and anchors the upper teeth
alveolar sac
cluster of alveoli
alveolus
small, grape-like sac that performs gas exchange in the lungs
amino acid
building block of proteins; characterized by an amino and carboxyl functional groups and a variable side-chain
amphipathic
describes a molecule that exhibits a difference in polarity between its two ends, resulting in a difference in water solubility
amphiarthrosis
slightly mobile joint
anal triangle
posterior triangle of the perineum that includes the anus
anaphase
third stage of mitosis (and meiosis), during which sister chromatids separate into two new nuclear regions of a dividing cell
anaphylactic shock
type of shock that follows a severe allergic reaction and results from massive vasodilation
anastomosis
(plural = anastomoses) area where vessels unite to allow blood to circulate even if there may be partial blockage in another branch
anatomical dead space
air space present in the airway that never reaches the alveoli and therefore never participates in gas exchange
anatomical neck
line on the humerus located around the outside margin of the humeral head
anatomical position
standard reference position used for describing locations and directions on the human body
anatomy
science that studies the form and composition of the body’s structures
anagen
active phase of the hair growth cycle
anconeus
small muscle on the lateral posterior elbow that extends the forearm
anemia
deficiency of red blood cells or hemoglobin
angiogenesis
formation of blood capillary networks
angle of the mandible
rounded corner located at outside margin of the body and ramus junction
angle of the rib
portion of rib with greatest curvature; together, the rib angles form the most posterior extent of the thoracic cage
anion
atom with a negative charge
ankle joint
joint that separates the leg and foot portions of the lower limb; formed by the articulations between the talus bone of the foot inferiorly, and the distal end of the tibia, medial malleolus of the tibia, and lateral malleolus of the fibula superiorly
annular ligament
intrinsic ligament of the elbow articular capsule that surrounds and supports the head of the radius at the proximal radioulnar joint
antagonist
muscle that opposes the action of an agonist
anterior
describes the front or direction toward the front of the body; also referred to as ventral
anterior arch
anterior portion of the ring-like C1 (atlas) vertebra
anterior border of the tibia
narrow, anterior margin of the tibia that extends inferiorly from the tibial tuberosity
anterior cardiac veins
vessels that parallel the small cardiac arteries and drain the anterior surface of the right ventricle; bypass the coronary sinus and drain directly into the right atrium
anterior cavity
larger body cavity located anterior to the posterior (dorsal) body cavity; includes the serous membrane-lined pleural cavities for the lungs, pericardial cavity for the heart, and peritoneal cavity for the abdominal and pelvic organs; also referred to as ventral cavity
anterior compartment of the arm
(anterior flexor compartment of the arm) the biceps brachii, brachialis, brachioradialis, and their associated blood vessels and nerves
anterior compartment of the forearm
(anterior flexor compartment of the forearm) deep and superficial muscles that originate on the humerus and insert into the hand
anterior compartment of the leg
region that includes muscles that dorsiflex the foot
anterior compartment of the thigh
region that includes muscles that flex the thigh and extend the leg
anterior cranial fossa
shallowest and most anterior cranial fossa of the cranial base that extends from the frontal bone to the lesser wing of the sphenoid bone
anterior cruciate ligament
intracapsular ligament of the knee; extends from anterior, superior surface of the tibia to the inner aspect of the lateral condyle of the femur; resists hyperextension of knee
anterior inferior iliac spine
small, bony projection located on the anterior margin of the ilium, below the anterior superior iliac spine
anterior interventricular artery
(also, left anterior descending artery or LAD) major branch of the left coronary artery that follows the anterior interventricular sulcus
anterior interventricular sulcus
sulcus located between the left and right ventricles on the anterior surface of the heart
anterior longitudinal ligament
ligament that runs the length of the vertebral column, uniting the anterior aspects of the vertebral bodies
anterior (ventral) sacral foramen
one of the series of paired openings located on the anterior (ventral) side of the sacrum
anterior sacroiliac ligament
strong ligament between the sacrum and the ilium portions of the hip bone that supports the anterior side of the sacroiliac joint
anterior scalene
a muscle anterior to the middle scalene
anterior superior iliac spine
rounded, anterior end of the iliac crest
anterior talofibular ligament
intrinsic ligament located on the lateral side of the ankle joint, between talus bone and lateral malleolus of fibula; supports talus at the talocrural joint and resists excess inversion of the foot
antibodies
(also, immunoglobulins or gamma globulins) antigen-specific proteins produced by specialized B lymphocytes that protect the body by binding to foreign objects such as bacteria and viruses
anticoagulant
substance such as heparin that opposes coagulation
antigenic determinant
(also, epitope) one of the chemical groups recognized by a single type of lymphocyte antigen receptor
antigen presentation
binding of processed antigen to the protein-binding cleft of a major histocompatibility complex molecule
antigen processing
internalization and digestion of antigen in an antigen-presenting cell
antigen receptor
two-chain receptor by which lymphocytes recognize antigen
antithrombin
anticoagulant that inactivates factor X and opposes the conversion of prothrombin (factor II) into thrombin in the common pathway
aortic valve
(also, aortic semilunar valve) valve located at the base of the aorta
anticodon
consecutive sequence of three nucleotides on a tRNA molecule that is complementary to a specific codon on an mRNA molecule
anulus fibrosus
tough, fibrous outer portion of an intervertebral disc, which is strongly anchored to the bodies of the adjacent vertebrae
aortic sinuses
small pockets in the ascending aorta near the aortic valve that are the locations of the baroreceptors (stretch receptors) and chemoreceptors that trigger a reflex that aids in the regulation of vascular homeostasis
apex
tip of the external nose
appendicular skeleton
all bones of the upper and lower limbs, plus the girdle bones that attach each limb to the axial skeleton
apneustic center
network of neurons within the pons that stimulate the neurons in the dorsal respiratory group; controls the depth of inspiration
apocrine sweat gland
type of sweat gland that is associated with hair follicles in the armpits and genital regions
aponeurosis
broad, tendon-like sheet of connective tissue that attaches a skeletal muscle to another skeletal muscle or to a bone
aponeurosis
broad, tendon-like sheet of connective tissue that attaches a skeletal muscle to another skeletal muscle or to a bone
appendicular
of the arms and legs
arcuate line of the ilium
smooth ridge located at the inferior margin of the iliac fossa; forms the lateral portion of the pelvic brim
arrector pili
smooth muscle that is activated in response to external stimuli that pull on hair follicles and make the hair “stand up”
areolar tissue
(also, loose connective tissue) a type of connective tissue proper that shows little specialization with cells dispersed in the matrix
arm
region of the upper limb located between the shoulder and elbow joints; contains the humerus bone
arteriole
(also, resistance vessel) very small artery that leads to a capillary
arteriovenous anastomosis
short vessel connecting an arteriole directly to a venule and bypassing the capillary beds
artery
blood vessel that conducts blood away from the heart; may be a conducting or distributing vessel
articular cartilage
thin layer of cartilage covering an epiphysis; reduces friction and acts as a shock absorber
articular tubercle
smooth ridge located on the inferior skull, immediately anterior to the mandibular fossa
articulation
joint of the body/where two bone surfaces meet
artificial pacemaker
medical device that transmits electrical signals to the heart to ensure that it contracts and pumps blood to the body
astrocyte
star-shaped cell in the central nervous system that regulates ions and uptake and/or breakdown of some neurotransmitters and contributes to the formation of the blood-brain barrier
atlantoaxial joint
series of three articulations between the atlas (C1) vertebra and the axis (C2) vertebra, consisting of the joints between the inferior articular processes of C1 and the superior articular processes of C2, and the articulation between the dens of C2 and the anterior arch of C1
atlanto-occipital joint
articulation between the occipital condyles of the skull and the superior articular processes of the atlas (C1 vertebra)
atlas
first cervical (C1) vertebrae
atmospheric pressure
amount of force that is exerted by gases in the air surrounding any given surface
atom
smallest unit of an element that retains the unique properties of that element
atomic number
number of protons in the nucleus of an atom
ATPase
enzyme that hydrolyzes ATP to ADP
atrial reflex
(also, called Bainbridge reflex) autonomic reflex that responds to stretch receptors in the atria that send impulses to the cardioaccelerator area to increase HR when venous flow into the atria increases
atrial reflex
mechanism for maintaining vascular homeostasis involving atrial baroreceptors: if blood is returning to the right atrium more rapidly than it is being ejected from the left ventricle, the atrial receptors will stimulate the cardiovascular centers to increase sympathetic firing and increase cardiac output until the situation is reversed; the opposite is also true
atrioventricular bundle
(also, bundle of His) group of specialized myocardial conductile cells that transmit the impulse from the AV node through the interventricular septum; form the left and right atrioventricular bundle branches
atrioventricular bundle branches
(also, left or right bundle branches) specialized myocardial conductile cells that arise from the bifurcation of the atrioventricular bundle and pass through the interventricular septum; lead to the Purkinje fibers and also to the right papillary muscle via the moderator band
atrioventricular (AV) node
clump of myocardial cells located in the inferior portion of the right atrium within the atrioventricular septum; receives the impulse from the SA node, pauses, and then transmits it into specialized conducting cells within the interventricular septum
atrioventricular septum
cardiac septum located between the atria and ventricles; atrioventricular valves are located here
atrioventricular valves
one-way valves located between the atria and ventricles; the valve on the right is called the tricuspid valve, and the one on the left is the mitral or bicuspid valve
atrium
(plural = atria) upper or receiving chamber of the heart that pumps blood into the lower chambers just prior to their contraction; the right atrium receives blood from the systemic circuit that flows into the right ventricle; the left atrium receives blood from the pulmonary circuit that flows into the left ventricle
atrophy
loss of structural proteins from muscle fibers
auricle
extension of an atrium visible on the superior surface of the heart
autolysis
breakdown of cells by their own enzymatic action
autonomic tone
contractile state during resting cardiac activity produced by mild sympathetic and parasympathetic stimulation
autorhythmicity
ability of cardiac muscle to initiate its own electrical impulse that triggers the mechanical contraction that pumps blood at a fixed pace without nervous or endocrine control
auricular surface of the ilium
roughened area located on the posterior, medial side of the ilium of the hip bone; articulates with the auricular surface of the sacrum to form the sacroiliac joint
autophagy
lysosomal breakdown of a cell’s own components
axial
of the trunk and head
axial skeleton
central, vertical axis of the body, including the skull, vertebral column, and thoracic cage
axis
second cervical (C2) vertebra
B lymphocytes
(also, B cells) lymphocytes that defend the body against specific pathogens and thereby provide specific immunity
Bachmann’s bundle
(also, interatrial band) group of specialized conducting cells that transmit the impulse directly from the SA node in the right atrium to the left atrium
Bainbridge reflex
(also, called atrial reflex) autonomic reflex that responds to stretch receptors in the atria that send impulses to the cardioaccelerator area to increase HR when venous flow into the atria increases
baroreceptor reflex
autonomic reflex in which the cardiac centers monitor signals from the baroreceptor stretch receptors and regulate heart function based on blood flow
basal cell
type of stem cell found in the stratum basale and in the hair matrix that continually undergoes cell division, producing the keratinocytes of the epidermis
basal metabolic rate (BMR)
amount of energy expended by the body at rest
base
compound that accepts hydrogen ions (H+) in solution
base of the metatarsal bone
expanded, proximal end of each metatarsal bone
basophils
granulocytes that stain with a basic (alkaline) stain and store histamine and heparin
belly
bulky central body of a muscle
bi
two
biaxial joint
type of diarthrosis; a joint that allows for movements within two planes (two axes)
biceps brachii
two-headed muscle that crosses the shoulder and elbow joints to flex the forearm while assisting in supinating it and flexing the arm at the shoulder
biceps femoris
hamstring muscle
bicipital groove
intertubercular groove; narrow groove located between the greater and lesser tubercles of the humerus
bicuspid valve
(also, mitral valve or left atrioventricular valve) valve located between the left atrium and ventricle; consists of two flaps of tissue
bilirubin
yellowish bile pigment produced when iron is removed from heme and is further broken down into waste products
biliverdin
green bile pigment produced when the non-iron portion of heme is degraded into a waste product; converted to bilirubin in the liver
bipennate
pennate muscle that has fascicles that are located on both sides of the tendon
blood
liquid connective tissue composed of formed elements—erythrocytes, leukocytes, and platelets—and a fluid extracellular matrix called plasma; component of the cardiovascular system
blood colloidal osmotic pressure (BCOP)
pressure exerted by colloids suspended in blood within a vessel; a primary determinant is the presence of plasma proteins
blood flow
movement of blood through a vessel, tissue, or organ that is usually expressed in terms of volume per unit of time
blood hydrostatic pressure
force blood exerts against the walls of a blood vessel or heart chamber
blood pressure
force exerted by the blood against the wall of a vessel or heart chamber; can be described with the more generic term hydrostatic pressure
body of the rib
shaft portion of a rib
Bohr effect
relationship between blood pH and oxygen dissociation from hemoglobin
bond
electrical force linking atoms
bone
hard, dense connective tissue that forms the structural elements of the skeleton
bone marrow biopsy
diagnostic test of a sample of red bone marrow
bone marrow transplant
treatment in which a donor’s healthy bone marrow with its stem cells replaces diseased or damaged bone marrow of a patient
Boyle’s law
relationship between volume and pressure as described by the formula: P1V1 = P2V2
brachialis
muscle deep to the biceps brachii that provides power in flexing the forearm.
brachioradialis
muscle that can flex the forearm quickly or help lift a load slowly
brain case
portion of the skull that contains and protects the brain, consisting of the eight bones that form the cranial base and rounded upper skull
brevis
short
bronchial tree
collective name for the multiple branches of the bronchi and bronchioles of the respiratory system
bridge
portion of the external nose that lies in the area of the nasal bones
bronchiole
branch of bronchi that are 1 mm or less in diameter and terminate at alveolar sacs
bronchus
tube connected to the trachea that branches into many subsidiaries and provides a passageway for air to enter and leave the lungs
buccinator
muscle that compresses the cheek
buffer
solution containing a weak acid or a weak base that opposes wide fluctuations in the pH of body fluids
buffy coat
thin, pale layer of leukocytes and platelets that separates the erythrocytes from the plasma in a sample of centrifuged blood
bundle of His
(also, atrioventricular bundle) group of specialized myocardial conductile cells that transmit the impulse from the AV node through the interventricular septum; form the left and right atrioventricular bundle branches
calcaneal tendon
(also, Achilles tendon) strong tendon that inserts into the calcaneal bone of the ankle
calcaneofibular ligament
intrinsic ligament located on the lateral side of the ankle joint, between the calcaneus bone and lateral malleolus of the fibula; supports the talus bone at the ankle joint and resists excess inversion of the foot
calcaneus
heel bone; posterior, inferior tarsal bone that forms the heel of the foot
calvaria
(also, skullcap) rounded top of the skull
canaliculi
(singular = canaliculus) channels within the bone matrix that house one of an osteocyte’s many cytoplasmic extensions that it uses to communicate and receive nutrients
capacitation
process that occurs in the female reproductive tract in which sperm are prepared for fertilization; leads to increased motility and changes in their outer membrane that improve their ability to release enzymes capable of digesting an oocyte’s outer layers
capacitance
ability of a vein to distend and store blood
capacitance vessels
veins
capillary
smallest of blood vessels where physical exchange occurs between the blood and tissue cells surrounded by interstitial fluid
capillary bed
network of 10–100 capillaries connecting arterioles to venules
capillary hydrostatic pressure (CHP)
force blood exerts against a capillary
capitate
from the lateral side, the third of the four distal carpal bones; articulates with the scaphoid and lunate proximally, the trapezoid laterally, the hamate medially, and primarily with the third metacarpal distally
capitulum
knob-like bony structure located anteriorly on the lateral, distal end of the humerus
carbaminohemoglobin
compound of carbon dioxide and hemoglobin, and one of the ways in which carbon dioxide is carried in the blood
carbaminohemoglobin
bound form of hemoglobin and carbon dioxide
carbohydrate
class of organic compounds built from sugars, molecules containing carbon, hydrogen, and oxygen in a 1-2-1 ratio
carbonic anhydrase (CA)
enzyme that catalyzes the reaction that causes carbon dioxide and water to form carbonic acid
cardiac cycle
period of time between the onset of atrial contraction (atrial systole) and ventricular relaxation (ventricular diastole)
cardiac muscle
heart muscle, under involuntary control, composed of striated cells that attach to form fibers, each cell contains a single nucleus, contracts autonomously
cardiac muscle
striated muscle found in the heart; joined to one another at intercalated discs and under the regulation of pacemaker cells, which contract as one unit to pump blood through the circulatory system. Cardiac muscle is under involuntary control
cardiac notch
depression in the medial surface of the inferior lobe of the left lung where the apex of the heart is located
cardiac output (CO)
amount of blood pumped by each ventricle during one minute; equals HR multiplied by SV
cardiac plexus
paired complex network of nerve fibers near the base of the heart that receive sympathetic and parasympathetic stimulations to regulate HR
cardiac reflexes
series of autonomic reflexes that enable the cardiovascular centers to regulate heart function based upon sensory information from a variety of visceral sensors
cardiac reserve
difference between maximum and resting CO
cardiac skeleton
(also, skeleton of the heart) reinforced connective tissue located within the atrioventricular septum; includes four rings that surround the openings between the atria and ventricles, and the openings to the pulmonary trunk and aorta; the point of attachment for the heart valves
cardiogenic shock
type of shock that results from the inability of the heart to maintain cardiac output
cardiomyocyte
muscle cell of the heart
carpal bone
one of the eight small bones that form the wrist and base of the hand; these are grouped as a proximal row consisting of (from lateral to medial) the scaphoid, lunate, triquetrum, and pisiform bones, and a distal row containing (from lateral to medial) the trapezium, trapezoid, capitate, and hamate bones
carpal tunnel
passageway between the anterior forearm and hand formed by the carpal bones and flexor retinaculum
carpometacarpal joint
articulation between one of the carpal bones in the distal row and a metacarpal bone of the hand
cartilaginous joint
joint at which the bones are united by hyaline cartilage (synchondrosis) or fibrocartilage (symphysis)
carotid canal
zig-zag tunnel providing passage through the base of the skull for the internal carotid artery to the brain; begins anteromedial to the styloid process and terminates in the middle cranial cavity, near the posterior-lateral base of the sella turcica
carotid sinuses
small pockets near the base of the internal carotid arteries that are the locations of the baroreceptors and chemoreceptors that trigger a reflex that aids in the regulation of vascular homeostasis
cartilage
semi-rigid connective tissue found on the skeleton in areas where flexibility and smooth surfaces support movement
catagen
transitional phase marking the end of the anagen phase of the hair growth cycle
catalyst
substance that increases the rate of a chemical reaction without itself being changed in the process
cation
atom with a positive charge
caudal
describes a position below or lower than another part of the body proper; near or toward the tail (in humans, the coccyx, or lowest part of the spinal column); also referred to as inferior
caval opening
opening in the diaphragm that allows the inferior vena cava to pass through; foramen for the vena cava
cell cycle
life cycle of a single cell, from its birth until its division into two new daughter cells
cell membrane
membrane surrounding all animal cells, composed of a lipid bilayer interspersed with various molecules; also known as plasma membrane
centriole
small, self-replicating organelle that provides the origin for microtubule growth and moves DNA during cell division
central canal
longitudinal channel in the center of each osteon; contains blood vessels, nerves, and lymphatic vessels; also known as the Haversian canal
central chemoreceptor
one of the specialized receptors that are located in the brain that sense changes in hydrogen ion, oxygen, or carbon dioxide concentrations in the brain
cervical curve
posteriorly concave curvature of the cervical vertebral column region; a secondary curve of the vertebral column
cervical vertebrae
seven vertebrae numbered as C1–C7 that are located in the neck region of the vertebral column
central tolerance
B cell tolerance induced in immature B cells of the bone marrow
centromere
region of attachment for two sister chromatids
centrosome
cellular structure that organizes microtubules during cell division
channel protein
membrane-spanning protein that has an inner pore which allows the passage of one or more substances
checkpoint
progress point in the cell cycle during which certain conditions must be met in order for the cell to proceed to a subsequence phase
chemical energy
form of energy that is absorbed as chemical bonds form, stored as they are maintained, and released as they are broken
chemokine
soluble, long-range, cell-to-cell communication molecule
chloride shift
facilitated diffusion that exchanges bicarbonate (HCO3–) with chloride (Cl–) ions
chondrocytes
cells of the cartilage
chordae tendineae
string-like extensions of tough connective tissue that extend from the flaps of the atrioventricular valves to the papillary muscles
chromatin
substance consisting of DNA and associated proteins
chromosome
condensed version of chromatin
chronic inflammation
inflammation occurring for long periods of time
cilia
small appendage on certain cells formed by microtubules and modified for movement of materials across the cellular surface
circular
(also, sphincter) fascicles that are concentrically arranged around an opening
circulatory shock
also simply called shock; a life-threatening medical condition in which the circulatory system is unable to supply enough blood flow to provide adequate oxygen and other nutrients to the tissues to maintain cellular metabolism
circumduction
circular motion of the arm, thigh, hand, thumb, or finger that is produced by the sequential combination of flexion, abduction, extension, and adduction
circumflex artery
branch of the left coronary artery that follows coronary sulcus
class switching
ability of B cells to change the class of antibody they produce without altering the specificity for antigen
clavicle
collarbone; elongated bone that articulates with the manubrium of the sternum medially and the acromion of the scapula laterally
clavicular notch
paired notches located on the superior-lateral sides of the sternal manubrium, for articulation with the clavicle
cleavage furrow
contractile ring that forms around a cell during cytokinesis that pinches the cell into two halves
clone
group of lymphocytes sharing the same antigen receptor
clonal anergy
process whereby B cells that react to soluble antigens in bone marrow are made nonfunctional
clonal deletion
removal of self-reactive B cells by inducing apoptosis
clonal expansion
growth of a clone of selected lymphocytes
clonal selection
stimulating growth of lymphocytes that have specific receptors
closed reduction
manual manipulation of a broken bone to set it into its natural position without surgery
clotting factors
group of 12 identified substances active in coagulation
coagulation
formation of a blood clot; part of the process of hemostasis
coccyx
small bone located at inferior end of the adult vertebral column that is formed by the fusion of four coccygeal vertebrae; also referred to as the “tailbone”
codon
consecutive sequence of three nucleotides on an mRNA molecule that corresponds to a specific amino acid
colloid
liquid mixture in which the solute particles consist of clumps of molecules large enough to scatter light
collagen fiber
flexible fibrous proteins that give connective tissue tensile strength
colony-stimulating factors (CSFs)
glycoproteins that trigger the proliferation and differentiation of myeloblasts into granular leukocytes (basophils, neutrophils, and eosinophils)
common pathway
final coagulation pathway activated either by the intrinsic or the extrinsic pathway, and ending in the formation of a blood clot
compliance
degree to which a blood vessel can stretch as opposed to being rigid
compound
substance composed of two or more different elements joined by chemical bonds
compact bone
dense osseous tissue that can withstand compressive forces
complement
enzymatic cascade of constitutive blood proteins that have antipathogen effects, including the direct killing of bacteria
compressor urethrae
deep perineal muscle in women
concentration
number of particles within a given space
concentration gradient
difference in the concentration of a substance between two regions
concentric contraction
muscle contraction that shortens the muscle to move a load
conducting zone
region of the respiratory system that includes the organs and structures that provide passageways for air and are not directly involved in gas exchange
conduction
transfer of heat through physical contact
condylar process of the mandible
thickened upward projection from posterior margin of mandibular ramus
condyle
oval-shaped process located at the top of the condylar process of the mandible
connective tissue
type of tissue that serves to hold in place, connect, and integrate the body’s organs and systems
connective tissue membrane
connective tissue that encapsulates organs and lines movable joints
connective tissue proper
connective tissue containing a viscous matrix, fibers, and cells.
constant region domain
part of a lymphocyte antigen receptor that does not vary much between different receptor types
contractility
ability to shorten (contract) forcibly
control center
compares values to their normal range; deviations cause the activation of an effector
convection
transfer of heat between the skin and air or water
convergent
fascicles that extend over a broad area and converge on a common attachment site
continuous capillary
most common type of capillary, found in virtually all tissues except epithelia and cartilage; contains very small gaps in the endothelial lining that permit exchange
contraction phase
twitch contraction phase when tension increases
coracobrachialis
muscle that flexes and adducts the arm
coracoclavicular ligament
strong band of connective tissue that anchors the coracoid process of the scapula to the lateral clavicle; provides important indirect support for the acromioclavicular joint
coracoid process
short, hook-like process that projects anteriorly and laterally from the superior margin of the scapula
corona radiata
in an oocyte, a layer of granulosa cells that surrounds the oocyte and that must be penetrated by sperm before fertilization can occur
coronal suture
joint that unites the frontal bone to the right and left parietal bones across the top of the skull
coronary arteries
branches of the ascending aorta that supply blood to the heart; the left coronary artery feeds the left side of the heart, the left atrium and ventricle, and the interventricular septum; the right coronary artery feeds the right atrium, portions of both ventricles, and the heart conduction system
coronary sinus
large, thin-walled vein on the posterior surface of the heart that lies within the atrioventricular sulcus and drains the heart myocardium directly into the right atrium
coronary sulcus
sulcus that marks the boundary between the atria and ventricles
coronary veins
vessels that drain the heart and generally parallel the large surface arteries
coronoid fossa
depression on the anterior surface of the humerus above the trochlea; this space receives the coronoid process of the ulna when the elbow is maximally flexed
coronoid process of the mandible
flattened upward projection from the anterior margin of the mandibular ramus
coronoid process of the ulna
projecting bony lip located on the anterior, proximal ulna; forms the inferior margin of the trochlear notch
corrugator supercilii
prime mover of the eyebrows
cortex
in hair, the second or middle layer of keratinocytes originating from the hair matrix, as seen in a cross-section of the hair bulb
cortical reaction
following fertilization, the release of cortical granules from the oocyte’s plasma membrane into the zona pellucida creating a fertilization membrane that prevents any further attachment or penetration of sperm; part of the slow block to polyspermy
costal cartilage
hyaline cartilage structure attached to the anterior end of each rib that provides for either direct or indirect attachment of most ribs to the sternum
costal facet
site on the lateral sides of a thoracic vertebra for articulation with the head of a rib
costal groove
shallow groove along the inferior margin of a rib that provides passage for blood vessels and a nerve
costoclavicular ligament
band of connective tissue that unites the medial clavicle with the first rib
covalent bond
chemical bond in which two atoms share electrons, thereby completing their valence shells
coracohumeral ligament
intrinsic ligament of the shoulder joint; runs from the coracoid process of the scapula to the anterior humerus
coxal bone
hip bone
cranial
describes a position above or higher than another part of the body proper; also referred to as superior
cranial cavity
division of the posterior (dorsal) cavity that houses the brain
cranial cavity
interior space of the skull that houses the brain
cranium
skull
creatine phosphate
phosphagen used to store energy from ATP and transfer it to muscle
cribriform plate
small, flattened areas with numerous small openings, located to either side of the midline in the floor of the anterior cranial fossa; formed by the ethmoid bone
cricoid cartilage
portion of the larynx composed of a ring of cartilage with a wide posterior region and a thinner anterior region; attached to the esophagus
crista galli
small upward projection located at the midline in the floor of the anterior cranial fossa; formed by the ethmoid bone
cross matching
blood test for identification of blood type using antibodies and small samples of blood
cutaneous membrane
skin; epithelial tissue made up of a stratified squamous epithelial cells that cover the outside of the body
cuboid
tarsal bone that articulates posteriorly with the calcaneus bone, medially with the lateral cuneiform bone, and anteriorly with the fourth and fifth metatarsal bones
cuticle
in hair, the outermost layer of keratinocytes originating from the hair matrix, as seen in a cross-section of the hair bulb
cyclin
one of a group of proteins that function in the progression of the cell cycle
cyclin-dependent kinase (CDK)
one of a group of enzymes associated with cyclins that help them perform their functions
cytokinesis
final stage in cell division, where the cytoplasm divides to form two separate daughter cells
cytokine
soluble, short-range, cell-to-cell communication molecule
cytokines
class of proteins that act as autocrine or paracrine signaling molecules; in the cardiovascular system, they stimulate the proliferation of progenitor cells and help to stimulate both nonspecific and specific resistance to disease
cytoplasm
internal material between the cell membrane and nucleus of a cell, mainly consisting of a water-based fluid called cytosol, within which are all the other organelles and cellular solute and suspended materials
cytoskeleton
“skeleton” of a cell; formed by rod-like proteins that support the cell’s shape and provide, among other functions, locomotive abilities
cytosol
clear, semi-fluid medium of the cytoplasm, made up mostly of water
cytotoxic T cells (Tc)
T lymphocytes with the ability to induce apoptosis in target cells
deep
describes a position farther from the surface of the body
deep anterior compartment
flexor pollicis longus, flexor digitorum profundus, and their associated blood vessels and nerves
deep posterior compartment of the forearm
(deep posterior extensor compartment of the forearm) the abductor pollicis longus, extensor pollicis brevis, extensor pollicis longus, extensor indicis, and their associated blood vessels and nerves
deep transverse perineal
deep perineal muscle in men
defensins
antimicrobial proteins released from neutrophils and macrophages that create openings in the plasma membranes to kill cells
deglutition
swallowing
delayed hypersensitivity
(type IV) T cell-mediated immune response against pathogens infiltrating interstitial tissues, causing cellular infiltrate
deltoid
shoulder muscle that abducts the arm as well as flexes and medially rotates it, and extends and laterally rotates it
deltoid ligament
broad intrinsic ligament located on the medial side of the ankle joint; supports the talus at the talocrural joint and resists excess eversion of the foot
deltoid tuberosity
roughened, V-shaped region located laterally on the mid-shaft of the humerus
denaturation
change in the structure of a molecule through physical or chemical means
dens
bony projection (odontoid process) that extends upward from the body of the C2 (axis) vertebra
dense connective tissue
connective tissue proper that contains many fibers that provide both elasticity and protection
deoxyhemoglobin
molecule of hemoglobin without an oxygen molecule bound to it
deoxyribonucleic acid (DNA)
deoxyribose-containing nucleotide that stores genetic information
depolarize
to reduce the voltage difference between the inside and outside of a cell’s plasma membrane (the sarcolemma for a muscle fiber), making the inside less negative than at rest
depolarize
to reduce the voltage difference between the inside and outside of a cell’s plasma membrane (the sarcolemma for a muscle fiber), making the inside less negative than at rest
depression
downward (inferior) motion of the scapula or mandible
dermal papilla
(plural = dermal papillae) extension of the papillary layer of the dermis that increases surface contact between the epidermis and dermis
dermis
layer of skin between the epidermis and hypodermis, composed mainly of connective tissue and containing blood vessels, hair follicles, sweat glands, and other structures
desmosome
structure that forms an impermeable junction between cells
diarthrosis
freely mobile joint
diapedesis
(also, emigration) process by which leukocytes squeeze through adjacent cells in a blood vessel wall to enter tissues
diaphragm
skeletal muscle that separates the thoracic and abdominal cavities and is dome-shaped at rest
diaphysis
tubular shaft that runs between the proximal and distal ends of a long bone
diastole
period of time when the heart muscle is relaxed and the chambers fill with blood
diastolic pressure
lower number recorded when measuring arterial blood pressure; represents the minimal value corresponding to the pressure that remains during ventricular relaxation
diffusion
movement of a substance from an area of higher concentration to one of lower concentration
digastric
muscle that has anterior and posterior bellies and elevates the hyoid bone and larynx when one swallows; it also depresses the mandible
diploë
layer of spongy bone, that is sandwiched between two the layers of compact bone found in flat bones
diploid
condition marked by the presence of a double complement of genetic material (two sets of chromosomes, one set inherited from each of two parents)
disaccharide
pair of carbohydrate monomers bonded by dehydration synthesis via a glycosidic bond
disulfide bond
covalent bond formed within a polypeptide between sulfide groups of sulfur-containing amino acids, for example, cysteine
distal
describes a position farther from the point of attachment or the trunk of the body
distal radioulnar joint
articulation between the head of the ulna and the ulnar notch of the radius
distal tibiofibular joint
articulation between the distal fibula and the fibular notch of the tibia
DNA polymerase
enzyme that functions in adding new nucleotides to a growing strand of DNA during DNA replication
DNA replication
process of duplicating a molecule of DNA
dorsal
describes the back or direction toward the back of the body; also referred to as posterior
dorsal cavity
posterior body cavity that houses the brain and spinal cord; also referred to the posterior body cavity
dorsal group
region that includes the extensor digitorum brevis
dorsal interossei
muscles that abduct and flex the three middle fingers at the metacarpophalangeal joints and extend them at the interphalangeal joints
dorsal respiratory group (DRG)
region of the medulla oblongata that stimulates the contraction of the diaphragm and intercostal muscles to induce inspiration
dorsiflexion
movement at the ankle that brings the top of the foot toward the anterior leg
dorsum nasi
intermediate portion of the external nose that connects the bridge to the apex and is supported by the nasal bone
ear ossicles
three small bones located in the middle ear cavity that serve to transmit sound vibrations to the inner ear
early induced immune response
includes antimicrobial proteins stimulated during the first several days of an infection
eccentric contraction
muscle contraction that lengthens the muscle as the tension is diminished
eccrine sweat gland
type of sweat gland that is common throughout the skin surface; it produces a hypotonic sweat for thermoregulation
ectoderm
outermost embryonic germ layer from which the epidermis and the nervous tissue derive
effector
organ that can cause a change in a value
effector T cells
immune cells with a direct, adverse effect on a pathogen
ejection fraction
portion of the blood that is pumped or ejected from the heart with each contraction; mathematically represented by SV divided by EDV
elastic artery
(also, conducting artery) artery with abundant elastic fibers located closer to the heart, which maintains the pressure gradient and conducts blood to smaller branches
elastic cartilage
type of cartilage, with elastin as the major protein, characterized by rigid support as well as elasticity
elastic fiber
fibrous protein within connective tissue that contains a high percentage of the protein elastin that allows the fibers to stretch and return to original size
elasticity
ability to stretch and rebound
elastin fibers
fibers made of the protein elastin that increase the elasticity of the dermis
elbow joint
joint located between the upper arm and forearm regions of the upper limb; formed by the articulations between the trochlea of the humerus and the trochlear notch of the ulna, and the capitulum of the humerus and the head of the radius
elbow joint
humeroulnar joint
electrocardiogram (ECG)
surface recording of the electrical activity of the heart that can be used for diagnosis of irregular heart function; also abbreviated as EKG
electron
subatomic particle having a negative charge and nearly no mass; found orbiting the atom’s nucleus
electron shell
area of space a given distance from an atom’s nucleus in which electrons are grouped
electrical gradient
difference in the electrical charge (potential) between two regions
eleiden
clear protein-bound lipid found in the stratum lucidum that is derived from keratohyalin and helps to prevent water loss
element
substance that cannot be created or broken down by ordinary chemical means
elevation
upward (superior) motion of the scapula or mandible
embolus
thrombus that has broken free from the blood vessel wall and entered the circulation
emigration
(also, diapedesis) process by which leukocytes squeeze through adjacent cells in a blood vessel wall to enter tissues
end diastolic volume (EDV)
(also, preload) the amount of blood in the ventricles at the end of atrial systole just prior to ventricular contraction
end systolic volume (ESV)
amount of blood remaining in each ventricle following systole
endocardium
innermost layer of the heart lining the heart chambers and heart valves; composed of endothelium reinforced with a thin layer of connective tissue that binds to the myocardium
endochondral ossification
process in which bone forms by replacing hyaline cartilage
endocytosis
import of material into the cell by formation of a membrane-bound vesicle
endoderm
innermost embryonic germ layer from which most of the digestive system and lower respiratory system derive
endomysium
loose, and well-hydrated connective tissue covering each muscle fiber in a skeletal muscle
endomysium
loose, and well-hydrated connective tissue covering each muscle fiber in a skeletal muscle
endoplasmic reticulum (ER)
cellular organelle that consists of interconnected membrane-bound tubules, which may or may not be associated with ribosomes (rough type or smooth type, respectively)
endosteum
delicate membranous lining of a bone’s medullary cavity
endothelium
layer of smooth, simple squamous epithelium that lines the endocardium and blood vessels
enzyme
protein or RNA that catalyzes chemical reactions
eosinophils
granulocytes that stain with eosin; they release antihistamines and are especially active against parasitic worms
epicardial coronary arteries
surface arteries of the heart that generally follow the sulci
epicardium
innermost layer of the serous pericardium and the outermost layer of the heart wall
epicranial aponeurosis
(also, galea aponeurosis) flat broad tendon that connects the frontalis and occipitalis
epidermis
outermost tissue layer of the skin
epiglottis
leaf-shaped piece of elastic cartilage that is a portion of the larynx that swings to close the trachea during swallowing
epimysium
outer layer of connective tissue around a skeletal muscle
epimysium
outer layer of connective tissue around a skeletal muscle
epiphyseal plate
(also, growth plate) sheet of hyaline cartilage in the metaphysis of an immature bone; replaced by bone tissue as the organ grows in length
epiphyseal line
completely ossified remnant of the epiphyseal plate
epiphysis
wide section at each end of a long bone; filled with spongy bone and red marrow
epithelial membrane
epithelium attached to a layer of connective tissue
epithelial tissue
type of tissue that serves primarily as a covering or lining of body parts, protecting the body; it also functions in absorption, transport, and secretion
eponychium
nail fold that meets the proximal end of the nail body, also called the cuticle
erector spinae group
large muscle mass of the back; primary extensor of the vertebral column
erythroblastosis fetalis
disease of Rh factor-positive newborns in Rh-negative mothers with multiple Rh-positive children; resulting from the action of maternal antibodies against fetal blood
erythrocyte
(also, red blood cell) mature myeloid blood cell that is composed mostly of hemoglobin and functions primarily in the transportation of oxygen and carbon dioxide
erythropoietin (EPO)
glycoprotein that triggers the bone marrow to produce RBCs; secreted by the kidney in response to low oxygen levels
ethmoid air cell
one of several small, air-filled spaces located within the lateral sides of the ethmoid bone, between the orbit and upper nasal cavity
ethmoid bone
unpaired bone that forms the roof and upper, lateral walls of the nasal cavity, portions of the floor of the anterior cranial fossa and medial wall of orbit, and the upper portion of the nasal septum
evaporation
transfer of heat that occurs when water changes from a liquid to a gas
eversion
foot movement involving the intertarsal joints of the foot in which the bottom of the foot is turned laterally, away from the midline
exchange reaction
type of chemical reaction in which bonds are both formed and broken, resulting in the transfer of components
excitability
ability to undergo neural stimulation
excitation-contraction coupling
sequence of events from motor neuron signaling to a skeletal muscle fiber to contraction of the fiber’s sarcomeres
excitation-contraction coupling
sequence of events from motor neuron signaling to a skeletal muscle fiber to contraction of the fiber’s sarcomeres
exocytosis
export of a substance out of a cell by formation of a membrane-bound vesicle
exon
one of the coding regions of an mRNA molecule that remain after splicing
expiration
(also, exhalation) process that causes the air to leave the lungs
expiratory reserve volume (ERV)
amount of air that can be forcefully exhaled after a normal tidal exhalation
extensibility
ability to lengthen (extend)
extension
movement in the sagittal plane that increases the angle of a joint (straightens the joint); motion involving posterior bending of the vertebral column or returning to the upright position from a flexed position
extensor
muscle that increases the angle at the joint
extensor carpi radialis brevis
muscle that extends and abducts the hand at the wrist
extensor carpi ulnaris
muscle that extends and adducts the hand
extensor digiti minimi
muscle that extends the little finger
extensor digitorum
muscle that extends the hand at the wrist and the phalanges
extensor digitorum brevis
muscle that extends the toes
extensor digitorum longus
muscle that is lateral to the tibialis anterior
extensor hallucis longus
muscle that is partly deep to the tibialis anterior and extensor digitorum longus
extensor indicis
muscle that inserts onto the tendon of the extensor digitorum of the index finger
extensor pollicis brevis
muscle that inserts onto the base of the proximal phalanx of the thumb
extensor pollicis longus
muscle that inserts onto the base of the distal phalanx of the thumb
extensor radialis longus
muscle that extends and abducts the hand at the wrist
extensor retinaculum
band of connective tissue that extends over the dorsal surface of the hand
external acoustic meatus
ear canal opening located on the lateral side of the skull
external callus
collar of hyaline cartilage and bone that forms around the outside of a fracture
external elastic membrane
membrane composed of elastic fibers that separates the tunica media from the tunica externa; seen in larger arteries
external intercostal
superficial intercostal muscles that raise the rib cage
external nose
region of the nose that is easily visible to others
external oblique
superficial abdominal muscle with fascicles that extend inferiorly and medially
external occipital protuberance
small bump located at the midline on the posterior skull
external root sheath
outer layer of the hair follicle that is an extension of the epidermis, which encloses the hair root
extracellular fluid (ECF)
fluid exterior to cells; includes the interstitial fluid, blood plasma, and fluid found in other reservoirs in the body
extrinsic eye muscles
originate outside the eye and insert onto the outer surface of the white of the eye, and create eyeball movement
extrinsic pathway
initial coagulation pathway that begins with tissue damage and results in the activation of the common pathway
extrinsic muscles of the hand
muscles that move the wrists, hands, and fingers and originate on the arm
facet
small, flattened area on a bone for an articulation (joint) with another bone, or for muscle attachment
facial bones
fourteen bones that support the facial structures and form the upper and lower jaws and the hard palate
facilitated diffusion
diffusion of a substance with the aid of a membrane protein
false ribs
vertebrochondral ribs 8–12 whose costal cartilage either attaches indirectly to the sternum via the costal cartilage of the next higher rib or does not attach to the sternum at all
fas ligand
molecule expressed on cytotoxic T cells and NK cells that binds to the fas molecule on a target cell and induces it do undergo apoptosis
fascicle
bundle of muscle fibers within a skeletal muscle
fascicle
bundle of muscle fibers within a skeletal muscle
fascicle
muscle fibers bundled by perimysium into a unit
fauces
portion of the posterior oral cavity that connects the oral cavity to the oropharynx
Fc region
in an antibody molecule, the site where the two termini of the heavy chains come together; many cells have receptors for this portion of the antibody, adding functionality to these molecules
femoral triangle
region formed at the junction between the hip and the leg and includes the pectineus, femoral nerve, femoral artery, femoral vein, and deep inguinal lymph nodes
femoropatellar joint
portion of the knee joint consisting of the articulation between the distal femur and the patella
femur
thigh bone; the single bone of the thigh
fenestrated capillary
type of capillary with pores or fenestrations in the endothelium that allow for rapid passage of certain small materials
ferritin
protein-containing storage form of iron found in the bone marrow, liver, and spleen
fertilization
unification of genetic material from male and female haploid gametes
fertilization membrane
impenetrable barrier that coats a nascent zygote; part of the slow block to polyspermy
fibrin
insoluble, filamentous protein that forms the structure of a blood clot
fibrinogen
plasma protein produced in the liver and involved in blood clotting
fibrinolysis
gradual degradation of a blood clot
fibroblast
most abundant cell type in connective tissue, secretes protein fibers and matrix into the extracellular space
fibrocartilage
tough form of cartilage, made of thick bundles of collagen fibers embedded in chondroitin sulfate ground substance
fibrocyte
less active form of fibroblast
fibroelastic membrane
specialized membrane that connects the ends of the C-shape cartilage in the trachea; contains smooth muscle fibers
fibrous joint
joint where the articulating areas of the adjacent bones are connected by fibrous connective tissue
fibula
thin, non-weight-bearing bone found on the lateral side of the leg
fibular collateral ligament
extrinsic ligament of the knee joint that spans from the lateral epicondyle of the femur to the head of the fibula; resists hyperextension and rotation of the extended knee
fibular notch
wide groove on the lateral side of the distal tibia for articulation with the fibula at the distal tibiofibular joint
fibularis brevis
(also, peroneus brevis) muscle that plantar flexes the foot at the ankle and everts it at the intertarsal joints
fibularis longus
(also, peroneus longus) muscle that plantar flexes the foot at the ankle and everts it at the intertarsal joints
fibularis tertius
small muscle that is associated with the extensor digitorum longus
filling time
duration of ventricular diastole during which filling occurs
fixator
synergist that assists an agonist by preventing or reducing movement at another joint, thereby stabilizing the origin of the agonist
flagellum
appendage on certain cells formed by microtubules and modified for movement
flat bone
thin and curved bone; serves as a point of attachment for muscles and protects internal organs
flexion
movement in the sagittal plane that decreases the angle of a joint (bends the joint); motion involving anterior bending of the vertebral column
flexion
movement that decreases the angle of a joint
flexor
muscle that decreases the angle at the joint
flexor carpi radialis
muscle that flexes and abducts the hand at the wrist
flexor carpi ulnaris
muscle that flexes and adducts the hand at the wrist
flexor digiti minimi brevis
muscle that flexes the little finger
flexor digitorum longus
muscle that flexes the four small toes
flexor digitorum profundus
muscle that flexes the phalanges of the fingers and the hand at the wrist
flexor digitorum superficialis
muscle that flexes the hand and the digits
flexor hallucis longus
muscle that flexes the big toe
flexor pollicis brevis
muscle that flexes the thumb
flexor pollicis longus
muscle that flexes the distal phalanx of the thumb
flexor retinaculum
band of connective tissue that extends over the palmar surface of the hand
flexor retinaculum
strong band of connective tissue at the anterior wrist that spans the top of the U-shaped grouping of the carpal bones to form the roof of the carpal tunnel
floating ribs
vertebral ribs 11–12 that do not attach to the sternum or to the costal cartilage of another rib
fluid connective tissue
specialized cells that circulate in a watery fluid containing salts, nutrients, and dissolved proteins
fontanelles
expanded areas of fibrous connective tissue that separate the braincase bones of the skull prior to birth and during the first year after birth
foot
portion of the lower limb located distal to the ankle joint
foramen lacerum
irregular opening in the base of the skull, located inferior to the exit of carotid canal
foramen ovale
opening in the fetal heart that allows blood to flow directly from the right atrium to the left atrium, bypassing the fetal pulmonary circuit
foramen magnum
large opening in the occipital bone of the skull through which the spinal cord emerges and the vertebral arteries enter the cranium
foramen ovale of the middle cranial fossa
oval-shaped opening in the floor of the middle cranial fossa
foramen rotundum
round opening in the floor of the middle cranial fossa, located between the superior orbital fissure and foramen ovale
foramen spinosum
small opening in the floor of the middle cranial fossa, located lateral to the foramen ovale
forced breathing
(also, hyperpnea) mode of breathing that occurs during exercise or by active thought that requires muscle contraction for both inspiration and expiration
forearm
region of the upper limb located between the elbow and wrist joints; contains the radius and ulna bones
formed elements
cellular components of blood; that is, erythrocytes, leukocytes, and platelets
fossa
(plural = fossae) shallow depression on the surface of a bone
fovea capitis
minor indentation on the head of the femur that serves as the site of attachment for the ligament to the head of the femur
fossa ovalis
oval-shaped depression in the interatrial septum that marks the former location of the foramen ovale
fracture
broken bone
fracture hematoma
blood clot that forms at the site of a broken bone
Frank-Starling mechanism
relationship between ventricular stretch and contraction in which the force of heart contraction is directly proportional to the initial length of the muscle fiber
frontal bone
unpaired bone that forms forehead, roof of orbit, and floor of anterior cranial fossa
frontal sinus
air-filled space within the frontal bone; most anterior of the paranasal sinuses
frontal plane
two-dimensional, vertical plane that divides the body or organ into anterior and posterior portions
frontalis
front part of the occipitofrontalis muscle
functional group
group of atoms linked by strong covalent bonds that tends to behave as a distinct unit in chemical reactions with other atoms
functional residual capacity (FRC)
sum of ERV and RV, which is the amount of air that remains in the lungs after a tidal expiration
fusiform
muscle that has fascicles that are spindle-shaped to create large bellies
G0 phase
phase of the cell cycle, usually entered from the G1 phase; characterized by long or permanent periods where the cell does not move forward into the DNA synthesis phase
G1 phase
first phase of the cell cycle, after a new cell is born
G2 phase
third phase of the cell cycle, after the DNA synthesis phase
gastrocnemius
most superficial muscle of the calf
gene
functional length of DNA that provides the genetic information necessary to build a protein
gene expression
active interpretation of the information coded in a gene to produce a functional gene product
genioglossus
muscle that originates on the mandible and allows the tongue to move downward and forward
geniohyoid
muscle that depresses the mandible, and raises and pulls the hyoid bone anteriorly
glassy membrane
layer of connective tissue that surrounds the base of the hair follicle, connecting it to the dermis
glenohumeral joint
shoulder joint; articulation between the glenoid cavity of the scapula and head of the humerus; multiaxial ball-and-socket joint that allows for flexion/extension, abduction/adduction, circumduction, and medial/lateral rotation of the humerus
glenohumeral ligament
one of the three intrinsic ligaments of the shoulder joint that strengthen the anterior articular capsule
glenoid labrum
lip of fibrocartilage located around the outside margin of the glenoid cavity of the scapula
globin
heme-containing globular protein that is a constituent of hemoglobin
globulins
heterogeneous group of plasma proteins that includes transport proteins, clotting factors, immune proteins, and others
glottis
opening between the vocal folds through which air passes when producing speech
gluteal group
muscle group that extends, flexes, rotates, adducts, and abducts the femur
gluteal tuberosity
roughened area on the posterior side of the proximal femur, extending inferiorly from the base of the greater trochanter
gluteus maximus
largest of the gluteus muscles that extends the femur
gluteus medius
muscle deep to the gluteus maximus that abducts the femur at the hip
gluteus minimus
smallest of the gluteal muscles and deep to the gluteus medius
glycocalyx
coating of sugar molecules that surrounds the cell membrane
glycolysis
anaerobic breakdown of glucose to ATP
glycoprotein
protein that has one or more carbohydrates attached
Golgi apparatus
cellular organelle formed by a series of flattened, membrane-bound sacs that functions in protein modification, tagging, packaging, and transport
gracilis
muscle that adducts the thigh and flexes the leg at the knee
graded muscle response
modification of contraction strength
graft-versus-host disease
in bone marrow transplants; occurs when the transplanted cells mount an immune response against the recipient
granular leukocytes
leukocytes with abundant granules in their cytoplasm; specifically, neutrophils, eosinophils, and basophils
granzyme
apoptosis-inducing substance contained in granules of NK cells and cytotoxic T cells
great cardiac vein
vessel that follows the interventricular sulcus on the anterior surface of the heart and flows along the coronary sulcus into the coronary sinus on the posterior surface; parallels the anterior interventricular artery and drains the areas supplied by this vessel
greater pelvis
(also, greater pelvic cavity or false pelvis) broad space above the pelvic brim defined laterally by the fan-like portion of the upper ilium
greater sciatic foramen
pelvic opening formed by the greater sciatic notch of the hip bone, the sacrum, and the sacrospinous ligament
greater sciatic notch
large, U-shaped indentation located on the posterior margin of the ilium, superior to the ischial spine
greater trochanter
large, bony expansion of the femur that projects superiorly from the base of the femoral neck
greater tubercle
enlarged prominence located on the lateral side of the proximal humerus
ground substance
fluid or semi-fluid portion of the matrix
hair
keratinous filament growing out of the epidermis
hair bulb
structure at the base of the hair root that surrounds the dermal papilla
hair follicle
cavity or sac from which hair originates
hair matrix
layer of basal cells from which a strand of hair grows
hair papilla
mass of connective tissue, blood capillaries, and nerve endings at the base of the hair follicle
hair root
part of hair that is below the epidermis anchored to the follicle
hair shaft
part of hair that is above the epidermis but is not anchored to the follicle
Haldane effect
relationship between the partial pressure of oxygen and the affinity of hemoglobin for carbon dioxide
hallux
big toe; digit 1 of the foot
hamate
from the lateral side, the fourth of the four distal carpal bones; articulates with the lunate and triquetrum proximally, the fourth and fifth metacarpals distally, and the capitate laterally
hamstring group
three long muscles on the back of the leg
hand
region of the upper limb distal to the wrist joint
head of the femur
rounded, proximal end of the femur that articulates with the acetabulum of the hip bone to form the hip joint
head of the fibula
small, knob-like, proximal end of the fibula; articulates with the inferior aspect of the lateral condyle of the tibia
head of the humerus
smooth, rounded region on the medial side of the proximal humerus; articulates with the glenoid fossa of the scapula to form the glenohumeral (shoulder) joint
head of the metatarsal bone
expanded, distal end of each metatarsal bone
head of the radius
disc-shaped structure that forms the proximal end of the radius; articulates with the capitulum of the humerus as part of the elbow joint, and with the radial notch of the ulna as part of the proximal radioulnar joint
head of the ulna
small, rounded distal end of the ulna; articulates with the ulnar notch of the distal radius, forming the distal radioulnar joint
heart block
interruption in the normal conduction pathway
heart rate (HR)
number of times the heart contracts (beats) per minute
heart sounds
sounds heard via auscultation with a stethoscope of the closing of the atrioventricular valves (“lub”) and semilunar valves (“dub”)
heavy chain
larger protein chain of an antibody
helicase
enzyme that functions to separate the two DNA strands of a double helix during DNA replication
helper T cells (Th)
T cells that secrete cytokines to enhance other immune responses, involved in activation of both B and T cell lymphocytes
hematocrit
(also, packed cell volume) volume percentage of erythrocytes in a sample of centrifuged blood
heme
red, iron-containing pigment to which oxygen binds in hemoglobin
hemocytoblast
hemopoietic stem cell that gives rise to the formed elements of blood
hemoglobin
oxygen-carrying compound in erythrocytes
hemolysis
destruction (lysis) of erythrocytes and the release of their hemoglobin into circulation
hemolytic disease of the newborn (HDN)
(also, erythroblastosis fetalis) disorder causing agglutination and hemolysis in an Rh+ fetus or newborn of an Rh− mother
hemophilia
genetic disorder characterized by inadequate synthesis of clotting factors
hemopoiesis
production of the formed elements of blood
hemopoietic growth factors
chemical signals including erythropoietin, thrombopoietin, colony-stimulating factors, and interleukins that regulate the differentiation and proliferation of particular blood progenitor cells
hemopoietic stem cell
type of pluripotent stem cell that gives rise to the formed elements of blood (hemocytoblast)
hemorrhage
excessive bleeding
hemosiderin
protein-containing storage form of iron found in the bone marrow, liver, and spleen
hemostasis
physiological process by which bleeding ceases
heparin
short-acting anticoagulant stored in mast cells and released when tissues are injured, opposes prothrombin
hip bone
coxal bone; single bone that forms the pelvic girdle; consists of three areas, the ilium, ischium, and pubis
hip joint
joint located at the proximal end of the lower limb; formed by the articulation between the acetabulum of the hip bone and the head of the femur
histamine
vasoactive mediator in granules of mast cells and is the primary cause of allergies and anaphylactic shock
histone
family of proteins that associate with DNA in the nucleus to form chromatin
homologous
describes two copies of the same chromosome (not identical), one inherited from each parent
hook of the hamate bone
bony extension located on the anterior side of the hamate carpal bone
humeroradial joint
articulation between the capitulum of the humerus and head of the radius
humeroulnar joint
articulation between the trochlea of humerus and the trochlear notch of the ulna; uniaxial hinge joint that allows for flexion/extension of the forearm
humerus
single bone of the upper arm
hyaline cartilage
most common type of cartilage, smooth and made of short collagen fibers embedded in a chondroitin sulfate ground substance
hydrogen bond
dipole-dipole bond in which a hydrogen atom covalently bonded to an electronegative atom is weakly attracted to a second electronegative atom
hydrophilic
describes a substance or structure attracted to water
hydrophobic
describes a substance or structure repelled by water
hyoglossus
muscle that originates on the hyoid bone to move the tongue downward and flatten it
hyperextension
excessive extension of joint, beyond the normal range of movement
hyperflexion
excessive flexion of joint, beyond the normal range of movement
hyperpnea
increased rate and depth of ventilation due to an increase in oxygen demand that does not significantly alter blood oxygen or carbon dioxide levels
hypertension
chronic and persistent blood pressure measurements of 140/90 mm Hg or above
hypertonia
abnormally high muscle tone
hypertonic
describes a solution concentration that is higher than a reference concentration
hypodermis
connective tissue connecting the integument to the underlying bone and muscle
hypotonic
describes a solution concentration that is lower than a reference concentration
hypertrophic cardiomyopathy
pathological enlargement of the heart, generally for no known reason
hypertrophy
addition of structural proteins to muscle fibers
hyperventilation
increased ventilation rate that leads to abnormally low blood carbon dioxide levels and high (alkaline) blood pH
hypervolemia
abnormally high levels of fluid and blood within the body
hyponychium
thickened layer of stratum corneum that lies below the free edge of the nail
hypotonia
abnormally low muscle tone caused by the absence of low-level contractions
hypovolemia
abnormally low levels of fluid and blood within the body
hypovolemic shock
type of circulatory shock caused by excessive loss of blood volume due to hemorrhage or possibly dehydration
hypoxemia
below-normal level of oxygen saturation of blood (typically <95 percent)
hypoxia
lack of oxygen supply to the tissues
hypothenar
group of muscles on the medial aspect of the palm
hypothenar eminence
rounded contour of muscle at the base of the little finger
IgA
antibody whose dimer is secreted by exocrine glands, is especially effective against digestive and respiratory pathogens, and can pass immunity to an infant through breastfeeding
IgD
class of antibody whose only known function is as a receptor on naive B cells; important in B cell activation
IgE
antibody that binds to mast cells and causes antigen-specific degranulation during an allergic response
IgG
main blood antibody of late primary and early secondary responses; passed from mother to unborn child via placenta
IgM
antibody whose monomer is a surface receptor of naive B cells; the pentamer is the first antibody made blood plasma during primary responses
immunoglobulin
protein antibody; occurs as one of five main classes
infrahyoid muscles
anterior neck muscles that are attached to, and inferior to the hyoid bone
infraspinatus
muscle that laterally rotates the arm
iliac crest
curved, superior margin of the ilium
iliac fossa
shallow depression found on the anterior and medial surfaces of the upper ilium
iliacus
muscle that, along with the psoas major, makes up the iliopsoas
iliococcygeus
muscle that makes up the levator ani along with the pubococcygeus
iliocostalis cervicis
muscle of the iliocostalis group associated with the cervical region
iliocostalis group
laterally placed muscles of the erector spinae
iliocostalis lumborum
muscle of the iliocostalis group associated with the lumbar region
iliocostalis thoracis
muscle of the iliocostalis group associated with the thoracic region
iliofemoral ligament
intrinsic ligament spanning from the ilium of the hip bone to the femur, on the superior-anterior aspect of the hip joint
iliopsoas group
muscle group consisting of iliacus and psoas major muscles, that flexes the thigh at the hip, rotates it laterally, and flexes the trunk of the body onto the hip
iliotibial tract
muscle that inserts onto the tibia; made up of the gluteus maximus and connective tissues of the tensor fasciae latae
ilium
superior portion of the hip bone
immunoglobulins
(also, antibodies or gamma globulins) antigen-specific proteins produced by specialized B lymphocytes that protect the body by binding to foreign objects such as bacteria and viruses
immunological memory
ability of the adaptive immune response to mount a stronger and faster immune response upon re-exposure to a pathogen
inferior angle of the scapula
inferior corner of the scapula located where the medial and lateral borders meet
inferior extensor retinaculum
cruciate ligament of the ankle
inferior gemellus
muscle deep to the gluteus maximus on the lateral surface of the thigh that laterally rotates the femur at the hip
inferior pubic ramus
narrow segment of bone that passes inferiorly and laterally from the pubic body; joins with the ischial ramus to form the ischiopubic ramus
inferior rotation
movement of the scapula during upper limb adduction in which the glenoid cavity of the scapula moves in a downward direction as the medial end of the scapular spine moves in an upward direction
inferior vena cava
large systemic vein that returns blood to the heart from the inferior portion of the body
inflammation
basic innate immune response characterized by heat, redness, pain, and swelling
infraglenoid tubercle
small bump or roughened area located on the lateral border of the scapula, near the inferior margin of the glenoid cavity
infraspinous fossa
broad depression located on the posterior scapula, inferior to the spine
innermost intercostal
the deepest intercostal muscles that draw the ribs together
inspiration
(also, inhalation) process that causes air to enter the lungs
inspiratory capacity (IC)
sum of the TV and IRV, which is the amount of air that can maximally be inhaled past a tidal expiration
inspiratory reserve volume (IRV)
amount of air that enters the lungs due to deep inhalation past the tidal volume
integral protein
membrane-associated protein that spans the entire width of the lipid bilayer
integumentary system
skin and its accessory structures
interatrial band
(also, Bachmann’s bundle) group of specialized conducting cells that transmit the impulse directly from the SA node in the right atrium to the left atrium
interatrial septum
cardiac septum located between the two atria; contains the fossa ovalis after birth
intercalated disc
physical junction between adjacent cardiac muscle cells; consisting of desmosomes, specialized linking proteoglycans, and gap junctions that allow passage of ions between the two cells
intercondylar eminence
irregular elevation on the superior end of the tibia, between the articulating surfaces of the medial and lateral condyles
intercondylar fossa
deep depression on the posterior side of the distal femur that separates the medial and lateral condyles
intercostal muscles
muscles that span the spaces between the ribs
interferons
early induced proteins made in virally infected cells that cause nearby cells to make antiviral proteins
internal intercostal
muscles the intermediate intercostal muscles that draw the ribs together
internal oblique
flat, intermediate abdominal muscle with fascicles that run perpendicular to those of the external oblique
intermediate cuneiform
middle of the three cuneiform tarsal bones; articulates posteriorly with the navicular bone, medially with the medial cuneiform bone, laterally with the lateral cuneiform bone, and anteriorly with the second metatarsal bone
intermediate filament
type of cytoskeletal filament made of keratin, characterized by an intermediate thickness, and playing a role in resisting cellular tension
interosseous border of the fibula
small ridge running down the medial side of the fibular shaft; for attachment of the interosseous membrane between the fibula and tibia
interosseous border of the radius
narrow ridge located on the medial side of the radial shaft; for attachment of the interosseous membrane between the ulna and radius bones
interosseous border of the tibia
small ridge running down the lateral side of the tibial shaft; for attachment of the interosseous membrane between the tibia and fibula
interosseous border of the ulna
narrow ridge located on the lateral side of the ulnar shaft; for attachment of the interosseous membrane between the ulna and radius
interosseous membrane of the forearm
sheet of dense connective tissue that unites the radius and ulna bones
interosseous membrane of the leg
sheet of dense connective tissue that unites the shafts of the tibia and fibula bones
interphalangeal joint
articulation between adjacent phalanx bones of the hand or foot digits
interphase
entire life cycle of a cell, excluding mitosis
intertrochanteric crest
short, prominent ridge running between the greater and lesser trochanters on the posterior side of the proximal femur
intertrochanteric line
small ridge running between the greater and lesser trochanters on the anterior side of the proximal femur
intertubercular groove (sulcus)
bicipital groove; narrow groove located between the greater and lesser tubercles of the humerus
interleukins
signaling molecules that may function in hemopoiesis, inflammation, and specific immune responses
intermediate
group of midpalmar muscles
internal elastic membrane
membrane composed of elastic fibers that separates the tunica intima from the tunica media; seen in larger arteries
internal root sheath
innermost layer of keratinocytes in the hair follicle that surround the hair root up to the hair shaft
internodal pathways
specialized conductile cells within the atria that transmit the impulse from the SA node throughout the myocardial cells of the atrium and to the AV node
interventricular septum
cardiac septum located between the two ventricles
interstitial fluid (IF)
fluid in the small spaces between cells not contained within blood vessels
intra-alveolar pressure
(intrapulmonary pressure) pressure of the air within the alveoli
intracellular fluid (ICF)
fluid in the cytosol of cells
intrapleural pressure
pressure of the air within the pleural cavity
intrinsic muscles of the hand
muscles that move the wrists, hands, and fingers and originate in the palm
intrinsic pathway
initial coagulation pathway that begins with vascular damage or contact with foreign substances, and results in the activation of the common pathway
intron
non-coding regions of a pre-mRNA transcript that may be removed during splicing
inversion
foot movement involving the intertarsal joints of the foot in which the bottom of the foot is turned toward the midline
ion
atom with an overall positive or negative charge
ionic bond
attraction between an anion and a cation
ischemia
insufficient blood flow to the tissues
ischial ramus
bony extension projecting anteriorly and superiorly from the ischial tuberosity; joins with the inferior pubic ramus to form the ischiopubic ramus
ischial spine
pointed, bony projection from the posterior margin of the ischium that separates the greater sciatic notch and lesser sciatic notch
ischial tuberosity
large, roughened protuberance that forms the posteroinferior portion of the hip bone; weight-bearing region of the pelvis when sitting
ischiococcygeus
muscle that assists the levator ani and pulls the coccyx anteriorly
ischiofemoral ligament
intrinsic ligament spanning from the ischium of the hip bone to the femur, on the posterior aspect of the hip joint
ischiopubic ramus
narrow extension of bone that connects the ischial tuberosity to the pubic body; formed by the junction of the ischial ramus and inferior pubic ramus
ischium
posteroinferior portion of the hip bone
isometric contraction
muscle contraction that occurs with no change in muscle length
isotonic
describes a solution concentration that is the same as a reference concentration
isotonic contraction
muscle contraction that involves changes in muscle length
isotope
one of the variations of an element in which the number of neutrons differ from each other
isovolumic contraction
(also, isovolumetric contraction) initial phase of ventricular contraction in which tension and pressure in the ventricle increase, but no blood is pumped or ejected from the heart
isovolumic ventricular relaxation phase
initial phase of the ventricular diastole when pressure in the ventricles drops below pressure in the two major arteries, the pulmonary trunk, and the aorta, and blood attempts to flow back into the ventricles, producing the dicrotic notch of the ECG and closing the two semilunar valves
keratin
type of structural protein that gives skin, hair, and nails its hard, water-resistant properties
keratinocyte
cell that produces keratin and is the most predominant type of cell found in the epidermis
keratohyalin
granulated protein found in the stratum granulosum
kinetochore
region of a centromere where microtubules attach to a pair of sister chromatids
lacunae
(singular = lacuna) small spaces in bone or cartilage tissue that cells occupy
lamina propria
areolar connective tissue underlying a mucous membrane
Langerhans cell
specialized dendritic cell found in the stratum spinosum that functions as a macrophage
laryngeal prominence
region where the two lamina of the thyroid cartilage join, forming a protrusion known as “Adam’s apple”
laryngopharynx
portion of the pharynx bordered by the oropharynx superiorly and esophagus and trachea inferiorly; serves as a route for both air and food
larynx
cartilaginous structure that produces the voice, prevents food and beverages from entering the trachea, and regulates the volume of air that enters and leaves the lungs
lateral condyle of the femur
smooth, articulating surface that forms the distal and posterior sides of the lateral expansion of the distal femur
lateral condyle of the tibia
lateral, expanded region of the proximal tibia that includes the smooth surface that articulates with the lateral condyle of the femur as part of the knee joint
lateral cuneiform
most lateral of the three cuneiform tarsal bones; articulates posteriorly with the navicular bone, medially with the intermediate cuneiform bone, laterally with the cuboid bone, and anteriorly with the third metatarsal bone
lateral epicondyle of the femur
roughened area of the femur located on the lateral side of the lateral condyle
lateral epicondyle of the humerus
small projection located on the lateral side of the distal humerus
lateral malleolus
expanded distal end of the fibula
lateral supracondylar ridge
narrow, bony ridge located along the lateral side of the distal humerus, superior to the lateral epicondyle
left atrioventricular valve
(also, mitral valve or bicuspid valve) valve located between the left atrium and ventricle; consists of two flaps of tissue
leg
portion of the lower limb located between the knee and ankle joints
lesser trochanter
small, bony projection on the medial side of the proximal femur, at the base of the femoral neck
lesser tubercle
small, bony prominence located on anterior side of the proximal humerus
leukemia
cancer involving leukocytes
leukocyte
(also, white blood cell) colorless, nucleated blood cell, the chief function of which is to protect the body from disease
leukocytosis
excessive leukocyte proliferation
leukopenia
below-normal production of leukocytes
ligament of the head of the femur
ligament that spans the acetabulum of the hip bone and the fovea capitis of the femoral head
ligand
molecule that binds with specificity to a specific receptor molecule
light chain
small protein chain of an antibody
linea aspera
longitudinally running bony ridge located in the middle third of the posterior femur
lingual tonsil
lymphoid tissue located at the base of the tongue
loose connective tissue
(also, areolar tissue) type of connective tissue proper that shows little specialization with cells dispersed in the matrix
lumen
interior of a tubular structure such as a blood vessel or a portion of the alimentary canal through which blood, chyme, or other substances travel
lunate
from the lateral side, the second of the four proximal carpal bones; articulates with the radius proximally, the capitate and hamate distally, the scaphoid laterally, and the triquetrum medially
lymphocytes
agranular leukocytes of the lymphoid stem cell line, many of which function in specific immunity
lymphoma
form of cancer in which masses of malignant T and/or B lymphocytes collect in lymph nodes, the spleen, the liver, and other tissues
lysosome
membrane-bound cellular organelle originating from the Golgi apparatus and containing digestive enzymes
lysozyme
digestive enzyme with bactericidal properties
mass number
sum of the number of protons and neutrons in the nucleus of an atom
marginal arteries
branches of the right coronary artery that supply blood to the superficial portions of the right ventricle
matrix
extracellular material which is produced by the cells embedded in it, containing ground substance and fibers
matter
physical substance; that which occupies space and has mass
meatus
one of three recesses (superior, middle, and inferior) in the nasal cavity attached to the conchae that increase the surface area of the nasal cavity
medial condyle of the femur
smooth, articulating surface that forms the distal and posterior sides of the medial expansion of the distal femur
medial condyle of the tibia
medial, expanded region of the proximal tibia that includes the smooth surface that articulates with the medial condyle of the femur as part of the knee joint
medial cuneiform
most medial of the three cuneiform tarsal bones; articulates posteriorly with the navicular bone, laterally with the intermediate cuneiform bone, and anteriorly with the first and second metatarsal bones
medial epicondyle of the femur
roughened area of the distal femur located on the medial side of the medial condyle
medial epicondyle of the humerus
enlarged projection located on the medial side of the distal humerus
medial malleolus
bony expansion located on the medial side of the distal tibia
megakaryocyte
bone marrow cell that produces platelets
melanin
pigment that determines the color of hair and skin
melanocyte
cell found in the stratum basale of the epidermis that produces the pigment melanin
melanosome
intercellular vesicle that transfers melanin from melanocytes into keratinocytes of the epidermis
memory cell
type of B or T lymphocyte that forms after exposure to a pathogen
Merkel cell
receptor cell in the stratum basale of the epidermis that responds to the sense of touch
mesenchymal cell
adult stem cell from which most connective tissue cells are derived
mesenchyme
embryonic tissue from which connective tissue cells derive
mesoderm
middle embryonic germ layer from which connective tissue, muscle tissue, and some epithelial tissue derive
mesothelium
simple squamous epithelial portion of serous membranes, such as the superficial portion of the epicardium (the visceral pericardium) and the deepest portion of the pericardium (the parietal pericardium)
metacarpal bone
one of the five long bones that form the palm of the hand; numbered 1–5, starting on the lateral (thumb) side of the hand
metacarpophalangeal joint
articulation between the distal end of a metacarpal bone of the hand and a proximal phalanx bone of the thumb or a finger
metaphase
second stage of mitosis (and meiosis), characterized by the linear alignment of sister chromatids in the center of the cell
metaphase plate
linear alignment of sister chromatids in the center of the cell, which takes place during metaphase
metatarsal bone
one of the five elongated bones that forms the anterior half of the foot; numbered 1–5, starting on the medial side of the foot
metatarsophalangeal joint
articulation between a metatarsal bone of the foot and the proximal phalanx bone of a toe
metarteriole
short vessel arising from a terminal arteriole that branches to supply a capillary bed
MHC polygeny
multiple MHC genes and their proteins found in body cells
MHC polymorphism
multiple alleles for each individual MHC locus
microcirculation
blood flow through the capillaries
microfilament
the thinnest of the cytoskeletal filaments; composed of actin subunits that function in muscle contraction and cellular structural support
microtubule
the thickest of the cytoskeletal filaments, composed of tubulin subunits that function in cellular movement and structural support
midcarpal joint
articulation between the proximal and distal rows of the carpal bones; contributes to movements of the hand at the wrist
middle cardiac vein
vessel that parallels and drains the areas supplied by the posterior interventricular artery; drains into the great cardiac vein
mitochondrion
one of the cellular organelles bound by a double lipid bilayer that function primarily in the production of cellular energy (ATP)
mitosis
division of genetic material, during which the cell nucleus breaks down and two new, fully functional, nuclei are formed
mitotic phase
phase of the cell cycle in which a cell undergoes mitosis
mitotic spindle
network of microtubules, originating from centrioles, that arranges and pulls apart chromosomes during mitosis
mitral valve
(also, left atrioventricular valve or bicuspid valve) valve located between the left atrium and ventricle; consists of two flaps of tissue
moderator band
band of myocardium covered by endocardium that arises from the inferior portion of the interventricular septum in the right ventricle and crosses to the anterior papillary muscle; contains conductile fibers that carry electrical signals followed by contraction of the heart
monocytes
agranular leukocytes of the myeloid stem cell line that circulate in the bloodstream; tissue monocytes are macrophages
mucous connective tissue
specialized loose connective tissue present in the umbilical cord
mucous membrane
tissue membrane that is covered by protective mucous and lines tissue exposed to the outside environment
murmur
unusual heart sound detected by auscultation; typically related to septal or valve defects
muscle tissue
type of tissue that is capable of contracting and generating tension in response to stimulation; produces movement.
muscular artery
(also, distributing artery) artery with abundant smooth muscle in the tunica media that branches to distribute blood to the arteriole network
mutation
change in the nucleotide sequence in a gene within a cell’s DNA
myocardium
thickest layer of the heart composed of cardiac muscle cells built upon a framework of primarily collagenous fibers and blood vessels that supply it and the nervous fibers that help to regulate it
myocardial conducting cells
specialized cells that transmit electrical impulses throughout the heart and trigger contraction by the myocardial contractile cells
myocardial contractile cells
bulk of the cardiac muscle cells in the atria and ventricles that conduct impulses and contract to propel blood
naris
(plural = nares) opening of the nostrils
nasal bone
bone of the skull that lies under the root and bridge of the nose and is connected to the frontal and maxillary bones
nasal septum
wall composed of bone and cartilage that separates the left and right nasal cavities
nasopharynx
portion of the pharynx flanked by the conchae and oropharynx that serves as an airway
natural killer (NK) cells
cytotoxic lymphocytes capable of recognizing cells that do not express “self” proteins on their plasma membrane or that contain foreign or abnormal markers; provide generalized, nonspecific immunity
navicular
tarsal bone that articulates posteriorly with the talus bone, laterally with the cuboid bone, and anteriorly with the medial, intermediate, and lateral cuneiform bones
neck of the femur
narrowed region located inferior to the head of the femur
neck of the radius
narrowed region immediately distal to the head of the radius
negative inotropic factors
factors that negatively impact or lower heart contractility
nervi vasorum
small nerve fibers found in arteries and veins that trigger contraction of the smooth muscle in their walls
nervous tissue
type of tissue that is capable of sending and receiving impulses through electrochemical signals.
neutrophils
granulocytes that stain with a neutral dye and are the most numerous of the leukocytes; especially active against bacteria
nucleus
cell’s central organelle; contains the cell’s DNA
olecranon fossa
large depression located on the posterior side of the distal humerus; this space receives the olecranon process of the ulna when the elbow is fully extended
olecranon process
expanded posterior and superior portions of the proximal ulna; forms the bony tip of the elbow
organelle
any of several different types of membrane-enclosed specialized structures in the cell that perform specific functions for the cell
oropharynx
portion of the pharynx flanked by the nasopharynx, oral cavity, and laryngopharynx that is a passageway for both air and food
osmosis
diffusion of water molecules down their concentration gradient across a selectively permeable membrane
oxyhemoglobin
(Hb–O2) bound form of hemoglobin and oxygen
oxygen–hemoglobin dissociation curve
graph that describes the relationship of partial pressure to the binding and disassociation of oxygen to and from heme
P wave
component of the electrocardiogram that represents the depolarization of the atria
pacemaker
cluster of specialized myocardial cells known as the SA node that initiates the sinus rhythm
packed cell volume (PCV)
(also, hematocrit) volume percentage of erythrocytes present in a sample of centrifuged blood
palatine tonsil
one of the paired structures composed of lymphoid tissue located anterior to the uvula at the roof of isthmus of the fauces
palmar interossei
muscles that abduct and flex each finger at the metacarpophalangeal joints and extend each finger at the interphalangeal joints
palmaris longus
muscle that provides weak flexion of the hand at the wrist
papillary layer
superficial layer of the dermis, made of loose, areolar connective tissue
paranasal sinus
one of the cavities within the skull that is connected to the conchae that serve to warm and humidify incoming air, produce mucus, and lighten the weight of the skull; consists of frontal, maxillary, sphenoidal, and ethmoidal sinuses
parenchyma
functional cells of a gland or organ, in contrast with the supportive or connective tissue of a gland or organ
passive immunity
transfer of immunity to a pathogen to an individual that lacks immunity to this pathogen usually by the injection of antibodies
passive transport
form of transport across the cell membrane that does not require input of cellular energy
patella
kneecap; the largest sesamoid bone of the body; articulates with the distal femur
patellar surface
smooth groove located on the anterior side of the distal femur, between the medial and lateral condyles; site of articulation for the patella
pectoral girdle
shoulder girdle, made up of the clavicle and scapula
pectoralis major
thick, fan-shaped axial muscle that covers much of the superior thorax
pectoralis minor
muscle that moves the scapula and assists in inhalation
perfusion
distribution of blood into the capillaries so the tissues can be supplied
peripheral chemoreceptor
one of the specialized receptors located in the aortic arch and carotid arteries that sense changes in pH, carbon dioxide, or oxygen blood levels
peripheral protein
membrane-associated protein that does not span the width of the lipid bilayer, but is attached peripherally to integral proteins, membrane lipids, or other components of the membrane
peripheral tolerance
mature B cell made tolerant by lack of T cell help
peroxisome
membrane-bound organelle that contains enzymes primarily responsible for detoxifying harmful substances
phagocytosis
endocytosis of large particles
phalanx bone of the foot
(plural = phalanges) one of the 14 bones that form the toes; these include the proximal and distal phalanges of the big toe, and the proximal, middle, and distal phalanx bones of toes two through five
pharyngeal tonsil
structure composed of lymphoid tissue located in the nasopharynx
pharynx
region of the conducting zone that forms a tube of skeletal muscle lined with respiratory epithelium; located between the nasal conchae and the esophagus and trachea
phalanx bone of the hand
(plural = phalanges) one of the 14 bones that form the thumb and fingers; these include the proximal and distal phalanges of the thumb, and the proximal, middle, and distal phalanx bones of the fingers two through five
philtrum
concave surface of the face that connects the apex of the nose to the top lip
pinocytosis
endocytosis of fluid
plasma
in blood, the liquid extracellular matrix composed mostly of water that circulates the formed elements and dissolved materials throughout the cardiovascular system
platelets
(also, thrombocytes) one of the formed elements of blood that consists of cell fragments broken off from megakaryocytes
pneumotaxic center
network of neurons within the pons that inhibit the activity of the neurons in the dorsal respiratory group; controls rate of breathing
polymorphonuclear
having a lobed nucleus, as seen in some leukocytes
positive chemotaxis
process in which a cell is attracted to move in the direction of chemical stimuli
positive inotropic factors
factors that positively impact or increase heart contractility
papillary muscle
extension of the myocardium in the ventricles to which the chordae tendineae attach
pectinate muscles
muscular ridges seen on the anterior surface of the right atrium
pericardial cavity
cavity surrounding the heart filled with a lubricating serous fluid that reduces friction as the heart contracts
pericardial sac
(also, pericardium) membrane that separates the heart from other mediastinal structures; consists of two distinct, fused sublayers: the fibrous pericardium and the parietal pericardium
pericardium
(also, pericardial sac) membrane that separates the heart from other mediastinal structures; consists of two distinct, fused sublayers: the fibrous pericardium and the parietal pericardium
polyspermy
penetration of an oocyte by more than one sperm
posterior cardiac vein
vessel that parallels and drains the areas supplied by the marginal artery branch of the circumflex artery; drains into the great cardiac vein
posterior interventricular artery
(also, posterior descending artery) branch of the right coronary artery that runs along the posterior portion of the interventricular sulcus toward the apex of the heart and gives rise to branches that supply the interventricular septum and portions of both ventricles
posterior interventricular sulcus
sulcus located between the left and right ventricles on the anterior surface of the heart
precapillary sphincters
circular rings of smooth muscle that surround the entrance to a capillary and regulate blood flow into that capillary
preload
(also, end diastolic volume) amount of blood in the ventricles at the end of atrial systole just prior to ventricular contraction
prepotential depolarization
(also, spontaneous depolarization) mechanism that accounts for the autorhythmic property of cardiac muscle; the membrane potential increases as sodium ions diffuse through the always-open sodium ion channels and causes the electrical potential to rise
pronator quadratus
pronator that originates on the ulna and inserts on the radius
pronator teres
pronator that originates on the humerus and inserts on the radius
prophase
first stage of mitosis (and meiosis), characterized by breakdown of the nuclear envelope and condensing of the chromatin to form chromosomes
proximal tibiofibular joint
articulation between the head of the fibula and the inferior aspect of the lateral condyle of the tibia
pulmonary arteries
left and right branches of the pulmonary trunk that carry deoxygenated blood from the heart to each of the lungs
pulmonary capillaries
capillaries surrounding the alveoli of the lungs where gas exchange occurs: carbon dioxide exits the blood and oxygen enters
pulmonary circuit
blood flow to and from the lungs
pulmonary surfactant
substance composed of phospholipids and proteins that reduces the surface tension of the alveoli; made by type II alveolar cells
pulmonary trunk
large arterial vessel that carries blood ejected from the right ventricle; divides into the left and right pulmonary arteries
pulmonary valve
(also, pulmonary semilunar valve, the pulmonic valve, or the right semilunar valve) valve at the base of the pulmonary trunk that prevents backflow of blood into the right ventricle; consists of three flaps
pulmonary veins
veins that carry highly oxygenated blood into the left atrium, which pumps the blood into the left ventricle, which in turn pumps oxygenated blood into the aorta and to the many branches of the systemic circuit
Purkinje fibers
specialized myocardial conduction fibers that arise from the bundle branches and spread the impulse to the myocardial contraction fibers of the ventricles
Rh blood group
blood-type classification based on the presence or absence of the antigen Rh on the erythrocyte membrane surface
S phase
stage of the cell cycle during which DNA replication occurs
sinusoid capillary
rarest type of capillary, which has extremely large intercellular gaps in the basement membrane in addition to clefts and fenestrations; found in areas such as the bone marrow and liver where passage of large molecules occurs
sister chromatid
one of a pair of identical chromosomes, formed during DNA replication
somatic cell
all cells of the body excluding gamete cells
telophase
final stage of mitosis (and meiosis), preceding cytokinesis, characterized by the formation of two new daughter nuclei
thoroughfare channel
continuation of the metarteriole that enables blood to bypass a capillary bed and flow directly into a venule, creating a vascular shunt
tunica externa
(also, tunica adventitia) outermost layer or tunic of a vessel (except capillaries)
tunica intima
(also, tunica interna) innermost lining or tunic of a vessel
tunica media
middle layer or tunic of a vessel (except capillaries)
vasa vasorum
small blood vessels located within the walls or tunics of larger vessels that supply nourishment to and remove wastes from the cells of the vessels
vascular shunt
continuation of the metarteriole and thoroughfare channel that allows blood to bypass the capillary beds to flow directly from the arterial to the venous circulation
vasoconstriction
constriction of the smooth muscle of a blood vessel, resulting in a decreased vascular diameter
vasodilation
relaxation of the smooth muscle in the wall of a blood vessel, resulting in an increased vascular diameter
vasomotion
irregular, pulsating flow of blood through capillaries and related structures
vein
blood vessel that conducts blood toward the heart
venous reserve
volume of blood contained within systemic veins in the integument, bone marrow, and liver that can be returned to the heart for circulation, if needed
venule
small vessel leading from the capillaries to veins
stroke volume (SV)
amount of blood pumped by each ventricle per contraction; also, the difference between EDV and ESV
target heart rate
range in which both the heart and lungs receive the maximum benefit from an aerobic workout
systole
period of time when the heart muscle is contracting
ventricular ejection phase
second phase of ventricular systole during which blood is pumped from the ventricle
QRS complex
component of the electrocardiogram that represents the depolarization of the ventricles and includes, as a component, the repolarization of the atria
sinoatrial (SA) node
known as the pacemaker, a specialized clump of myocardial conducting cells located in the superior portion of the right atrium that has the highest inherent rate of depolarization that then spreads throughout the heart
sinus rhythm
normal contractile pattern of the heart
spontaneous depolarization
(also, prepotential depolarization) the mechanism that accounts for the autorhythmic property of cardiac muscle; the membrane potential increases as sodium ions diffuse through the always-open sodium ion channels and causes the electrical potential to rise
T wave
component of the electrocardiogram that represents the repolarization of the ventricles
right atrioventricular valve
(also, tricuspid valve) valve located between the right atrium and ventricle; consists of three flaps of tissue
semilunar valves
valves located at the base of the pulmonary trunk and at the base of the aorta
septum
(plural = septa) walls or partitions that divide the heart into chambers
septum primum
flap of tissue in the fetus that covers the foramen ovale within a few seconds after birth
small cardiac vein
parallels the right coronary artery and drains blood from the posterior surfaces of the right atrium and ventricle; drains into the great cardiac vein
sulcus
(plural = sulci) fat-filled groove visible on the surface of the heart; coronary vessels are also located in these areas
superior vena cava
large systemic vein that returns blood to the heart from the superior portion of the body
systemic circuit
blood flow to and from virtually all of the tissues of the body
immediate hypersensitivity
(type I) IgE-mediated mast cell degranulation caused by crosslinking of surface IgE by antigen
sensitization
first exposure to an antigen
severe combined immunodeficiency disease (SCID)
genetic mutation that affects both T cell and B cell arms of the immune response
type I hypersensitivity
immediate response mediated by mast cell degranulation caused by the crosslinking of the antigen-specific IgE molecules on the mast cell surface
type II hypersensitivity
cell damage caused by the binding of antibody and the activation of complement, usually against red blood cells
type III hypersensitivity
damage to tissues caused by the deposition of antibody-antigen (immune) complexes followed by the activation of complement
trabeculae carneae
ridges of muscle covered by endocardium located in the ventricles
tricuspid valve
term used most often in clinical settings for the right atrioventricular valve
valve
in the cardiovascular system, a specialized structure located within the heart or vessels that ensures one-way flow of blood
ventricle
one of the primary pumping chambers of the heart located in the lower portion of the heart; the left ventricle is the major pumping chamber on the lower left side of the heart that ejects blood into the systemic circuit via the aorta and receives blood from the left atrium; the right ventricle is the major pumping chamber on the lower right side of the heart that ejects blood into the pulmonary circuit via the pulmonary trunk and receives blood from the right atrium
plasmin
blood protein active in fibrinolysis
platelet plug
accumulation and adhesion of platelets at the site of blood vessel injury
serum
blood plasma that does not contain clotting factors
thrombin
enzyme essential for the final steps in formation of a fibrin clot
thrombosis
excessive clot formation
thrombus
aggregation of fibrin, platelets, and erythrocytes in an intact artery or vein
tissue factor
protein thromboplastin, which initiates the extrinsic pathway when released in response to tissue damage
vascular spasm
initial step in hemostasis, in which the smooth muscle in the walls of the ruptured or damaged blood vessel contracts
T lymphocytes
(also, T cells) lymphocytes that provide cellular-level immunity by physically attacking foreign or diseased cells
thrombocytes
platelets, one of the formed elements of blood that consists of cell fragments broken off from megakaryocytes
thrombocytopenia
condition in which there are too few platelets, resulting in abnormal bleeding (hemophilia)
thrombocytosis
condition in which there are too many platelets, resulting in abnormal clotting (thrombosis)
macrophage
phagocytic cell of the myeloid lineage; a matured monocyte
oxyhemoglobin
molecule of hemoglobin to which oxygen is bound
polycythemia
elevated level of hemoglobin, whether adaptive or pathological
reticulocyte
immature erythrocyte that may still contain fragments of organelles
major histocompatibility complex (MHC)
gene cluster whose proteins present antigens to T cells
memory T cells
long-lived immune cell reserved for future exposure to an pathogen
MHC class I
found on most cells of the body, it binds to the CD8 molecule on T cells
MHC class II
found on macrophages, dendritic cells, and B cells, it binds to CD4 molecules on T cells
negative selection
selection against thymocytes in the thymus that react with self-antigen
polyclonal response
response by multiple clones to a complex antigen with many determinants
primary adaptive response
immune system’s response to the first exposure to a pathogen
positive selection
selection of thymocytes within the thymus that interact with self, but not non-self, MHC molecules
regulatory T cells (Treg)
(also, suppressor T cells) class of CD4 T cells that regulates other T cell responses
secondary adaptive response
immune response observed upon re-exposure to a pathogen, which is stronger and faster than a primary response
T cell tolerance
process during T cell differentiation where most T cells that recognize antigens from one’s own body are destroyed
Th1 cells
cells that secrete cytokines that enhance the activity of macrophages and other cells
Th2 cells
cells that secrete cytokines that induce B cells to differentiate into antibody-secreting plasma cells
variable region domain
part of a lymphocyte antigen receptor that varies considerably between different receptor types
sickle cell disease
(also, sickle cell anemia) inherited blood disorder in which hemoglobin molecules are malformed, leading to the breakdown of RBCs that take on a characteristic sickle shape
thalassemia
inherited blood disorder in which maturation of RBCs does not proceed normally, leading to abnormal formation of hemoglobin and the destruction of RBCs
transferrin
plasma protein that binds reversibly to iron and distributes it throughout the body
lymphoid stem cells
type of hemopoietic stem cells that gives rise to lymphocytes, including various T cells, B cells, and NK cells, all of which function in immunity
myeloid stem cells
type of hemopoietic stem cell that gives rise to some formed elements, including erythrocytes, megakaryocytes that produce platelets, and a myeloblast lineage that gives rise to monocytes and three forms of granular leukocytes (neutrophils, eosinophils, and basophils)
pluripotent stem cell
stem cell that derives from totipotent stem cells and is capable of differentiating into many, but not all, cell types
totipotent stem cell
embryonic stem cell that is capable of differentiating into any and all cells of the body; enabling the full development of an organism
thrombopoietin
hormone secreted by the liver and kidneys that prompts the development of megakaryocytes into thrombocytes (platelets)
red blood cells (RBCs)
(also, erythrocytes) one of the formed elements of blood that transports oxygen
white blood cells (WBCs)
(also, leukocytes) one of the formed elements of blood that provides defense against disease agents and foreign materials
nutrient
chemical obtained from foods and beverages that is critical to human survival
pulmonary ventilation
exchange of gases between the lungs and the atmosphere; breathing
quiet breathing
(also, eupnea) mode of breathing that occurs at rest and does not require the cognitive thought of the individual
residual volume (RV)
amount of air that remains in the lungs after maximum exhalation
respiratory cycle
one sequence of inspiration and expiration
respiratory rate
total number of breaths taken each minute
respiratory volume
varying amounts of air within the lung at a given time
thoracic wall compliance
ability of the thoracic wall to stretch while under pressure
tidal volume (TV)
amount of air that normally enters the lungs during quiet breathing
total dead space
sum of the anatomical dead space and alveolar dead space
total lung capacity (TLC)
total amount of air that can be held in the lungs; sum of TV, ERV, IRV, and RV
transpulmonary pressure
pressure difference between the intrapleural and intra-alveolar pressures
ventral respiratory group (VRG)
region of the medulla oblongata that stimulates the contraction of the accessory muscles involved in respiration to induce forced inspiration and expiration
vital capacity (VC)
sum of TV, ERV, and IRV, which is all the volumes that participate in gas exchange
pressure
force exerted by a substance in contact with another substance
lateral excursion
side-to-side movement of the mandible away from the midline, toward either the right or left side
lateral flexion
bending of the neck or body toward the right or left side
lateral (external) rotation
movement of the arm at the shoulder joint or the thigh at the hip joint that moves the anterior surface of the limb away from the midline of the body
medial excursion
side-to-side movement that returns the mandible to the midline
medial (internal) rotation
movement of the arm at the shoulder joint or the thigh at the hip joint that brings the anterior surface of the limb toward the midline of the body
metabolic rate
amount of energy consumed minus the amount of energy expended by the body
radiation
transfer of heat via infrared waves
thermoneutral
external temperature at which the body does not expend any energy for thermoregulation, about 84 °F
thermoregulation
process of regulating the temperature of the body
opposition
thumb movement that brings the tip of the thumb in contact with the tip of a finger
plantar flexion
foot movement at the ankle in which the heel is lifted off of the ground
pronated position
forearm position in which the palm faces backward
pronation
forearm motion that moves the palm of the hand from the palm forward to the palm backward position
protraction
anterior motion of the scapula or mandible
reposition
movement of the thumb from opposition back to the anatomical position (next to index finger)
retraction
posterior motion of the scapula or mandible
rotation
movement of a bone around a central axis (atlantoaxial joint) or around its long axis (proximal radioulnar joint; shoulder or hip joint); twisting of the vertebral column resulting from the summation of small motions between adjacent vertebrae
superior rotation
movement of the scapula during upper limb abduction in which the glenoid cavity of the scapula moves in an upward direction as the medial end of the scapular spine moves in a downward direction
supinated position
forearm position in which the palm faces anteriorly (anatomical position)
supination
forearm motion that moves the palm of the hand from the palm backward to the palm forward position
sarcopenia
age-related muscle atrophy
gomphosis
type of fibrous joint in which the root of a tooth is anchored into its bony jaw socket by strong periodontal ligaments
interosseous membrane
wide sheet of fibrous connective tissue that fills the gap between two parallel bones, forming a syndesmosis; found between the radius and ulna of the forearm and between the tibia and fibula of the leg
ligament
strong band of dense connective tissue spanning between bones
periodontal ligament
band of dense connective tissue that anchors the root of a tooth into the bony jaw socket
suture
fibrous joint that connects the bones of the skull (except the mandible); an immobile joint (synarthrosis)
syndesmosis
type of fibrous joint in which two separated, parallel bones are connected by an interosseous membrane
synostosis
site at which adjacent bones or bony components have fused together
joint
site at which two or more bones or bone and cartilage come together (articulate)
joint cavity
space enclosed by the articular capsule of a synovial joint that is filled with synovial fluid and contains the articulating surfaces of the adjacent bones
multiaxial joint
type of diarthrosis; a joint that allows for movements within three planes (three axes)
synarthrosis
immobile or nearly immobile joint
synovial joint
joint at which the articulating surfaces of the bones are located within a joint cavity formed by an articular capsule
uniaxial joint
type of diarthrosis; joint that allows for motion within only one plane (one axis)
molecule
two or more atoms covalently bonded together
myogenic response
constriction or dilation in the walls of arterioles in response to pressures related to blood flow; reduces high blood flow or increases low blood flow to help maintain consistent flow to the capillary network
neurogenic shock
type of shock that occurs with cranial or high spinal injuries that damage the cardiovascular centers in the medulla oblongata or the nervous fibers originating from this region
obstructive shock
type of shock that occurs when a significant portion of the vascular system is blocked
macrophage
ameboid phagocyte found in several tissues throughout the body
mast cell
cell found in the skin and the lining of body cells that contains cytoplasmic granules with vasoactive mediators such as histamine
monocyte
precursor to macrophages and dendritic cells seen in the blood
neutrophil
phagocytic white blood cell recruited from the bloodstream to the site of infection via the bloodstream
opsonization
enhancement of phagocytosis by the binding of antibody or antimicrobial protein
pattern recognition receptor (PRR)
leukocyte receptor that binds to specific cell wall components of different bacterial species
perforin
molecule in NK cell and cytotoxic T cell granules that form pores in the membrane of a target cell
phagocytosis
movement of material from the outside to the inside of the cells via vesicles made from invaginations of the plasma membrane
sepsis
(also, septicemia) organismal-level inflammatory response to a massive infection
septic shock
(also, blood poisoning) type of shock that follows a massive infection resulting in organism-wide inflammation
vascular shock
type of shock that occurs when arterioles lose their normal muscular tone and dilate dramatically
polar molecule
molecule with regions that have opposite charges resulting from uneven numbers of electrons in the nuclei of the atoms participating in the covalent bond
psychoneuroimmunology
study of the connections between the immune, nervous, and endocrine systems
tissue typing
typing of MHC molecules between a recipient and donor for use in a potential transplantation procedure
messenger RNA (mRNA)
nucleotide molecule that serves as an intermediate in the genetic code between DNA and protein
polypeptide
chain of amino acids linked by peptide bonds
polyribosome
simultaneous translation of a single mRNA transcript by multiple ribosomes
promoter
region of DNA that signals transcription to begin at that site within the gene
proteome
full complement of proteins produced by a cell (determined by the cell’s specific gene expression)
ribosomal RNA (rRNA)
RNA that makes up the subunits of a ribosome
RNA polymerase
enzyme that unwinds DNA and then adds new nucleotides to a growing strand of RNA for the transcription phase of protein synthesis
spliceosome
complex of enzymes that serves to splice out the introns of a pre-mRNA transcript
splicing
the process of modifying a pre-mRNA transcript by removing certain, typically non-coding, regions
transcription
process of producing an mRNA molecule that is complementary to a particular gene of DNA
transfer RNA (tRNA)
molecules of RNA that serve to bring amino acids to a growing polypeptide strand and properly place them into the sequence
translation
process of producing a protein from the nucleotide sequence code of an mRNA transcript
triplet
consecutive sequence of three nucleotides on a DNA molecule that, when transcribed into an mRNA codon, corresponds to a particular amino acid
lateralis
to the outside
longus
long
maximus
largest
medialis
to the inside
medius
medium
minimus
smallest
oblique
at an angle
rectus
straight
tri
three
neutron
heavy subatomic particle having no electrical charge and found in the atom’s nucleus
periodic table of the elements
arrangement of the elements in a table according to their atomic number; elements having similar properties because of their electron arrangements compose columns in the table, while elements having the same number of valence shells compose rows in the table
proton
heavy subatomic particle having a positive charge and found in the atom’s nucleus
radioactive isotope
unstable, heavy isotope that gives off subatomic particles, or electromagnetic energy, as it decays; also called radioisotopes
valence shell
outermost electron shell of an atom
latent period
the time when a twitch does not produce contraction
motor unit
motor neuron and the group of muscle fibers it innervates
muscle tension
force generated by the contraction of the muscle; tension generated during isotonic contractions and isometric contractions
muscle tone
low levels of muscle contraction that occur when a muscle is not producing movement
myogram
instrument used to measure twitch tension
recruitment
increase in the number of motor units involved in contraction
relaxation phase
period after twitch contraction when tension decreases
tetanus
a continuous fused contraction
treppe
stepwise increase in contraction tension
twitch
single contraction produced by one action potential
wave summation
addition of successive neural stimuli to produce greater contraction
kinetic energy
energy that matter possesses because of its motion
potential energy
stored energy matter possesses because of the positioning or structure of its components
product
one or more substances produced by a chemical reaction
reactant
one or more substances that enter into the reaction
synthesis reaction
type of anabolic reaction in which two or more atoms or molecules bond, resulting in the formation of a larger molecule
inorganic compound
substance that does not contain both carbon and hydrogen
organic compound
substance that contains both carbon and hydrogen
pH
negative logarithm of the hydrogen ion (H+) concentration of a solution
solution
homogeneous liquid mixture in which a solute is dissolved into molecules within a solvent
suspension
liquid mixture in which particles distributed in the liquid settle out over time
lipid
class of nonpolar organic compounds built from hydrocarbons and distinguished by the fact that they are not soluble in water
macromolecule
large molecule formed by covalent bonding
monosaccharide
monomer of carbohydrate; also known as a simple sugar
nucleotide
class of organic compounds composed of one or more phosphate groups, a pentose sugar, and a base
peptide bond
covalent bond formed by dehydration synthesis between two amino acids
phospholipid
a lipid compound in which a phosphate group is combined with a diglyceride
phosphorylation
addition of one or more phosphate groups to an organic compound
polysaccharide
compound consisting of more than two carbohydrate monomers bonded by dehydration synthesis via glycosidic bonds
prostaglandin
lipid compound derived from fatty acid chains and important in regulating several body processes
protein
class of organic compounds that are composed of many amino acids linked together by peptide bonds
purine
nitrogen-containing base with a double ring structure; adenine and guanine
pyrimidine
nitrogen-containing base with a single ring structure; cytosine, thiamine, and uracil
ribonucleic acid (RNA)
ribose-containing nucleotide that helps manifest the genetic code as protein
steroid
(also, sterol) lipid compound composed of four hydrocarbon rings bonded to a variety of other atoms and molecules
substrate
reactant in an enzymatic reaction
triglyceride
lipid compound composed of a glycerol molecule bonded with three fatty acid chains
T cell-dependent antigen
antigen that binds to B cells, which requires signals from T cells to make antibody
T cell-independent antigen
binds to B cells, which do not require signals from T cells to make antibody
receptor
protein molecule that contains a binding site for another specific molecule (called a ligand)
receptor-mediated endocytosis
endocytosis of ligands attached to membrane-bound receptors
selective permeability
feature of any barrier that allows certain substances to cross but excludes others
sodium-potassium pump
(also, Na+/K+ ATP-ase) membrane-embedded protein pump that uses ATP to move Na+ out of a cell and K+ into the cell
vesicle
membrane-bound structure that contains materials within or outside of the cell
reactive oxygen species (ROS)
a group of extremely reactive peroxides and oxygen-containing radicals that may contribute to cellular damage
ribosome
cellular organelle that functions in protein synthesis
latissimus dorsi
broad, triangular axial muscle located on the inferior part of the back
lumbrical
muscle that flexes each finger at the metacarpophalangeal joints and extend each finger at the interphalangeal joints
opponens digiti minimi
muscle that brings the little finger across the palm to meet the thumb
opponens pollicis
muscle that moves the thumb across the palm to meet another finger
retinacula
fibrous bands that sheath the tendons at the wrist
rhomboid major
muscle that attaches the vertebral border of the scapula to the spinous process of the thoracic vertebrae
rhomboid minor
muscle that attaches the vertebral border of the scapula to the spinous process of the thoracic vertebrae
rotator cuff
(also, musculotendinous cuff) the circle of tendons around the shoulder joint
macrophage oxidative metabolism
metabolism turned on in macrophages by T cell signals that help destroy intracellular bacteria
neutralization
inactivation of a virus by the binding of specific antibody
seroconversion
clearance of pathogen in the serum and the simultaneous rise of serum antibody
serratus anterior
large and flat muscle that originates on the ribs and inserts onto the scapula
subclavius
muscle that stabilizes the clavicle during movement
subscapularis
muscle that originates on the anterior scapula and medially rotates the arm
superficial anterior compartment of the forearm
flexor carpi radialis, palmaris longus, flexor carpi ulnaris, flexor digitorum superficialis, and their associated blood vessels and nerves
superficial posterior compartment of the forearm
extensor radialis longus, extensor carpi radialis brevis, extensor digitorum, extensor digiti minimi, extensor carpi ulnaris, and their associated blood vessels and nerves
supinator
muscle that moves the palm and forearm anteriorly
supraspinatus
muscle that abducts the arm
teres major
muscle that extends the arm and assists in adduction and medial rotation of it
teres minor
muscle that laterally rotates and extends the arm
thenar
group of muscles on the lateral aspect of the palm
thenar eminence
rounded contour of muscle at the base of the thumb
trapezius
muscle that stabilizes the upper part of the back
triceps brachii
three-headed muscle that extends the forearm
genome
entire complement of an organism’s DNA; found within virtually every cell
nuclear envelope
membrane that surrounds the nucleus; consisting of a double lipid-bilayer
nuclear pore
one of the small, protein-lined openings found scattered throughout the nuclear envelope
nucleolus
small region of the nucleus that functions in ribosome synthesis
nucleosome
unit of chromatin consisting of a DNA strand wrapped around histone proteins
histology
microscopic study of tissue architecture, organization, and function
Korotkoff sounds
noises created by turbulent blood flow through the vessels
mean arterial pressure (MAP)
average driving force of blood to the tissues; approximated by taking diastolic pressure and adding 1/3 of pulse pressure
pulse
alternating expansion and recoil of an artery as blood moves through the vessel; an indicator of heart rate
pulse pressure
difference between the systolic and diastolic pressures
resistance
any condition or parameter that slows or counteracts the flow of blood
respiratory pump
increase in the volume of the thorax during inhalation that decreases air pressure, enabling venous blood to flow into the thoracic region, then exhalation increases pressure, moving blood into the atria
skeletal muscle pump
effect on increasing blood pressure within veins by compression of the vessel caused by the contraction of nearby skeletal muscle
sphygmomanometer
blood pressure cuff attached to a device that measures blood pressure
systolic pressure
larger number recorded when measuring arterial blood pressure; represents the maximum value following ventricular contraction
respiratory bronchiole
specific type of bronchiole that leads to alveolar sacs
respiratory epithelium
ciliated lining of much of the conducting zone that is specialized to remove debris and pathogens, and produce mucus
respiratory membrane
alveolar and capillary wall together, which form an air-blood barrier that facilitates the simple diffusion of gases
respiratory zone
includes structures of the respiratory system that are directly involved in gas exchange
root
region of the external nose between the eyebrows
thyroid cartilage
largest piece of cartilage that makes up the larynx and consists of two lamina
trachea
tube composed of cartilaginous rings and supporting tissue that connects the lung bronchi and the larynx; provides a route for air to enter and exit the lung
trachealis muscle
smooth muscle located in the fibroelastic membrane of the trachea
true vocal cord
one of the pair of folded, white membranes that have a free inner edge that oscillates as air passes through to produce sound
type I alveolar cell
squamous epithelial cells that are the major cell type in the alveolar wall; highly permeable to gases
type II alveolar cell
cuboidal epithelial cells that are the minor cell type in the alveolar wall; secrete pulmonary surfactant
vestibular fold
part of the folded region of the glottis composed of mucous membrane; supports the epiglottis during swallowing
vascular tone
contractile state of smooth muscle in a blood vessel
serous membrane
type of tissue membrane that lines body cavities and lubricates them with serous fluid
synovial membrane
connective tissue membrane that lines the cavities of freely movable joints, producing synovial fluid for lubrication
tissue
group of cells that are similar in form and perform related functions
tissue membrane
thin layer or sheet of cells that covers the outside of the body, organs, and internal cavities
totipotent
embryonic cells that have the ability to differentiate into any type of cell and organ in the body
gross anatomy
study of the larger structures of the body, typically with the unaided eye; also referred to macroscopic anatomy
homeostasis
steady state of body systems that living organisms maintain
microscopic anatomy
study of very small structures of the body using magnification
physiology
science that studies the chemistry, biochemistry, and physics of the body’s functions
regional anatomy
study of the structures that contribute to specific body regions
systemic anatomy
study of the structures that contribute to specific body systems
reticular fiber
fine fibrous protein, made of collagen subunits, which cross-link to form supporting “nets” within connective tissue
reticular tissue
type of loose connective tissue that provides a supportive framework to soft organs, such as lymphatic tissue, spleen, and the liver
supportive connective tissue
type of connective tissue that provides strength to the body and protects soft tissue
myocyte
muscle cells
skeletal muscle
usually attached to bone, under voluntary control, each cell is a fiber that is multinucleated and striated
smooth muscle
under involuntary control, moves internal organs, cells contain a single nucleus, are spindle-shaped, and do not appear striated; each cell is a fiber
striation
alignment of parallel actin and myosin filaments which form a banded pattern
myelin
layer of lipid inside some neuroglial cells that wraps around the axons of some neurons
neuroglia
supportive neural cells
neuron
excitable neural cell that transfer nerve impulses
oligodendrocyte
neuroglial cell that produces myelin in the brain
Schwann cell
neuroglial cell that produces myelin in the peripheral nervous system
inferior
describes a position below or lower than another part of the body proper; near or toward the tail (in humans, the coccyx, or lowest part of the spinal column); also referred to as caudal
lateral
describes the side or direction toward the side of the body
medial
describes the middle or direction toward the middle of the body
pericardium
sac that encloses the heart
peritoneum
serous membrane that lines the abdominopelvic cavity and covers the organs found there
plane
imaginary two-dimensional surface that passes through the body
pleura
serous membrane that lines the pleural cavity and covers the lungs
posterior
describes the back or direction toward the back of the body; also referred to as dorsal
posterior cavity
posterior body cavity that houses the brain and spinal cord; also referred to as dorsal cavity
prone
face down
proximal
describes a position nearer to the point of attachment or the trunk of the body
sagittal plane
two-dimensional, vertical plane that divides the body or organ into right and left sides
section
in anatomy, a single flat surface of a three-dimensional structure that has been cut through
serous membrane
membrane that covers organs and reduces friction; also referred to as serosa
serosa
membrane that covers organs and reduces friction; also referred to as serous membrane
spinal cavity
division of the dorsal cavity that houses the spinal cord; also referred to as vertebral cavity
superficial
describes a position nearer to the surface of the body
superior
describes a position above or higher than another part of the body proper; also referred to as cranial
supine
face up
thoracic cavity
division of the anterior (ventral) cavity that houses the heart, lungs, esophagus, and trachea
transverse plane
two-dimensional, horizontal plane that divides the body or organ into superior and inferior portions
ventral
describes the front or direction toward the front of the body; also referred to as anterior
ventral cavity
larger body cavity located anterior to the posterior (dorsal) body cavity; includes the serous membrane-lined pleural cavities for the lungs, pericardial cavity for the heart, and peritoneal cavity for the abdominal and pelvic organs; also referred to as anterior body cavity
reticular layer
deeper layer of the dermis; it has a reticulated appearance due to the presence of abundant collagen and elastin fibers
stratum basale
deepest layer of the epidermis, made of epidermal stem cells
stratum corneum
most superficial layer of the epidermis
stratum granulosum
layer of the epidermis superficial to the stratum spinosum
stratum lucidum
layer of the epidermis between the stratum granulosum and stratum corneum, found only in thick skin covering the palms, soles of the feet, and digits
stratum spinosum
layer of the epidermis superficial to the stratum basale, characterized by the presence of desmosomes
vitiligo
skin condition in which melanocytes in certain areas lose the ability to produce melanin, possibly due an autoimmune reaction that leads to loss of color in patches
lunula
basal part of the nail body that consists of a crescent-shaped layer of thick epithelium
medulla
in hair, the innermost layer of keratinocytes originating from the hair matrix
nail bed
layer of epidermis upon which the nail body forms
nail body
main keratinous plate that forms the nail
nail cuticle
fold of epithelium that extends over the nail bed, also called the eponychium
nail fold
fold of epithelium at that extend over the sides of the nail body, holding it in place
nail root
part of the nail that is lodged deep in the epidermis from which the nail grows
sebaceous gland
type of oil gland found in the dermis all over the body and helps to lubricate and waterproof the skin and hair by secreting sebum
sebum
oily substance that is composed of a mixture of lipids that lubricates the skin and hair
sudoriferous gland
sweat gland
telogen
resting phase of the hair growth cycle initiated with catagen and terminated by the beginning of a new anagen phase of hair growth
Meissner corpuscle
(also, tactile corpuscle) receptor in the skin that responds to light touch
Pacinian corpuscle
(also, lamellated corpuscle) receptor in the skin that responds to vibration
rickets
disease in children caused by vitamin D deficiency, which leads to the weakening of bones
vitamin D
compound that aids absorption of calcium and phosphates in the intestine to improve bone health
hematopoiesis
production of blood cells, which occurs in the red marrow of the bones
orthopedist
doctor who specializes in diagnosing and treating musculoskeletal disorders and injuries
osseous tissue
bone tissue; a hard, dense connective tissue that forms the structural elements of the skeleton
red marrow
connective tissue in the interior cavity of a bone where hematopoiesis takes place
skeletal system
organ system composed of bones and cartilage that provides for movement, support, and protection
yellow marrow
connective tissue in the interior cavity of a bone where fat is stored
irregular bone
bone of complex shape; protects internal organs from compressive forces
long bone
cylinder-shaped bone that is longer than it is wide; functions as a lever
sesamoid bone
small, round bone embedded in a tendon; protects the tendon from compressive forces
short bone
cube-shaped bone that is approximately equal in length, width, and thickness; provides limited motion
hole
opening or depression in a bone
lacunae
(singular = lacuna) spaces in a bone that house an osteocyte
medullary cavity
hollow region of the diaphysis; filled with yellow marrow
nutrient foramen
small opening in the middle of the external surface of the diaphysis, through which an artery enters the bone to provide nourishment
osteoblast
cell responsible for forming new bone
osteoclast
cell responsible for resorbing bone
osteocyte
primary cell in mature bone; responsible for maintaining the matrix
osteogenic cell
undifferentiated cell with high mitotic activity; the only bone cells that divide; they differentiate and develop into osteoblasts
osteon
(also, Haversian system) basic structural unit of compact bone; made of concentric layers of calcified matrix
perforating canal
(also, Volkmann’s canal) channel that branches off from the central canal and houses vessels and nerves that extend to the periosteum and endosteum
periosteum
fibrous membrane covering the outer surface of bone and continuous with ligaments
projection
bone markings where part of the surface sticks out above the rest of the surface, where tendons and ligaments attach
spongy bone
(also, cancellous bone) trabeculated osseous tissue that supports shifts in weight distribution
trabeculae
(singular = trabecula) spikes or sections of the lattice-like matrix in spongy bone
intramembranous ossification
process by which bone forms directly from mesenchymal tissue
modeling
process, during bone growth, by which bone is resorbed on one surface of a bone and deposited on another
ossification
(also, osteogenesis) bone formation
ossification center
cluster of osteoblasts found in the early stages of intramembranous ossification
osteoid
uncalcified bone matrix secreted by osteoblasts
perichondrium
membrane that covers cartilage
primary ossification center
region, deep in the periosteal collar, where bone development starts during endochondral ossification
proliferative zone
region of the epiphyseal plate that makes new chondrocytes to replace those that die at the diaphyseal end of the plate and contributes to longitudinal growth of the epiphyseal plate
remodeling
process by which osteoclasts resorb old or damaged bone at the same time as and on the same surface where osteoblasts form new bone to replace that which is resorbed
reserve zone
region of the epiphyseal plate that anchors the plate to the osseous tissue of the epiphysis
secondary ossification center
region of bone development in the epiphyses
zone of calcified matrix
region of the epiphyseal plate closest to the diaphyseal end; functions to connect the epiphyseal plate to the diaphysis
zone of maturation and hypertrophy
region of the epiphyseal plate where chondrocytes from the proliferative zone grow and mature and contribute to the longitudinal growth of the epiphyseal plate
internal callus
fibrocartilaginous matrix, in the endosteal region, between the two ends of a broken bone
open reduction
surgical exposure of a bone to reset a fracture
hyoid bone
small, U-shaped bone located in upper neck that does not contact any other bone
ribs
thin, curved bones of the chest wall
sacrum
single bone located near the inferior end of the adult vertebral column that is formed by the fusion of five sacral vertebrae; forms the posterior portion of the pelvis
skeleton
bones of the body
skull
bony structure that forms the head, face, and jaws, and protects the brain; consists of 22 bones
sternum
flattened bone located at the center of the anterior chest
thoracic cage
consists of 12 pairs of ribs and sternum
vertebra
individual bone in the neck and back regions of the vertebral column
vertebral column
entire sequence of bones that extend from the skull to the tailbone
glabella
slight depression of frontal bone, located at the midline between the eyebrows
greater wings of sphenoid bone
lateral projections of the sphenoid bone that form the anterior wall of the middle cranial fossa and an area of the lateral skull
filtration
in the cardiovascular system, the movement of material from a capillary into the interstitial fluid, moving from an area of higher pressure to lower pressure
interstitial fluid colloidal osmotic pressure (IFCOP)
pressure exerted by the colloids within the interstitial fluid
interstitial fluid hydrostatic pressure (IFHP)
force exerted by the fluid in the tissue spaces
net filtration pressure (NFP)
force driving fluid out of the capillary and into the tissue spaces; equal to the difference of the capillary hydrostatic pressure and the blood colloidal osmotic pressure
reabsorption
in the cardiovascular system, the movement of material from the interstitial fluid into the capillaries
hard palate
bony structure that forms the roof of the mouth and floor of the nasal cavity, formed by the palatine process of the maxillary bones and the horizontal plate of the palatine bones
horizontal plate
medial extension from the palatine bone that forms the posterior quarter of the hard palate
hypoglossal canal
paired openings that pass anteriorly from the anterior-lateral margins of the foramen magnum deep to the occipital condyles
hypophyseal (pituitary) fossa
shallow depression on top of the sella turcica that houses the pituitary (hypophyseal) gland
inferior nasal concha
one of the paired bones that project from the lateral walls of the nasal cavity to form the largest and most inferior of the nasal conchae
infraorbital foramen
opening located on anterior skull, below the orbit
infratemporal fossa
space on lateral side of skull, below the level of the zygomatic arch and deep (medial) to the ramus of the mandible
internal acoustic meatus
opening into petrous ridge, located on the lateral wall of the posterior cranial fossa
jugular foramen
irregularly shaped opening located in the lateral floor of the posterior cranial cavity
lacrimal bone
paired bones that contribute to the anterior-medial wall of each orbit
lacrimal fossa
shallow depression in the anterior-medial wall of the orbit, formed by the lacrimal bone that gives rise to the nasolacrimal canal
lambdoid suture
inverted V-shaped joint that unites the occipital bone to the right and left parietal bones on the posterior skull
lateral pterygoid plate
paired, flattened bony projections of the sphenoid bone located on the inferior skull, lateral to the medial pterygoid plate
lesser wings of the sphenoid bone
lateral extensions of the sphenoid bone that form the bony lip separating the anterior and middle cranial fossae
lingula
small flap of bone located on the inner (medial) surface of mandibular ramus, next to the mandibular foramen
mandible
unpaired bone that forms the lower jaw bone; the only moveable bone of the skull
mandibular foramen
opening located on the inner (medial) surface of the mandibular ramus
mandibular fossa
oval depression located on the inferior surface of the skull
mandibular notch
large U-shaped notch located between the condylar process and coronoid process of the mandible
mastoid process
large bony prominence on the inferior, lateral skull, just behind the earlobe
maxillary bone
(also, maxilla) paired bones that form the upper jaw and anterior portion of the hard palate
maxillary sinus
air-filled space located with each maxillary bone; largest of the paranasal sinuses
medial pterygoid plate
paired, flattened bony projections of the sphenoid bone located on the inferior skull medial to the lateral pterygoid plate; form the posterior portion of the nasal cavity lateral wall
mental foramen
opening located on the anterior-lateral side of the mandibular body
mental protuberance
inferior margin of anterior mandible that forms the chin
middle cranial fossa
centrally located cranial fossa that extends from the lesser wings of the sphenoid bone to the petrous ridge
middle nasal concha
nasal concha formed by the ethmoid bone that is located between the superior and inferior conchae
mylohyoid line
bony ridge located along the inner (medial) surface of the mandibular body
nasal bone
paired bones that form the base of the nose
nasal cavity
opening through skull for passage of air
nasal conchae
curved bony plates that project from the lateral walls of the nasal cavity; include the superior and middle nasal conchae, which are parts of the ethmoid bone, and the independent inferior nasal conchae bone
nasal septum
flat, midline structure that divides the nasal cavity into halves, formed by the perpendicular plate of the ethmoid bone, vomer bone, and septal cartilage
nasolacrimal canal
passage for drainage of tears that extends downward from the medial-anterior orbit to the nasal cavity, terminating behind the inferior nasal conchae
occipital bone
unpaired bone that forms the posterior portions of the brain case and base of the skull
occipital condyle
paired, oval-shaped bony knobs located on the inferior skull, to either side of the foramen magnum
optic canal
opening spanning between middle cranial fossa and posterior orbit
orbit
bony socket that contains the eyeball and associated muscles
palatine bone
paired bones that form the posterior quarter of the hard palate and a small area in floor of the orbit
palatine process
medial projection from the maxilla bone that forms the anterior three quarters of the hard palate
paranasal sinuses
cavities within the skull that are connected to the conchae that serve to warm and humidify incoming air, produce mucus, and lighten the weight of the skull; consist of frontal, maxillary, sphenoidal, and ethmoidal sinuses
parietal bone
paired bones that form the upper, lateral sides of the skull
perpendicular plate of the ethmoid bone
downward, midline extension of the ethmoid bone that forms the superior portion of the nasal septum
petrous ridge
petrous portion of the temporal bone that forms a large, triangular ridge in the floor of the cranial cavity, separating the middle and posterior cranial fossae; houses the middle and inner ear structures
posterior cranial fossa
deepest and most posterior cranial fossa; extends from the petrous ridge to the occipital bone
pterion
H-shaped suture junction region that unites the frontal, parietal, temporal, and sphenoid bones on the lateral side of the skull
ramus of the mandible
vertical portion of the mandible
sagittal suture
joint that unites the right and left parietal bones at the midline along the top of the skull
sella turcica
elevated area of sphenoid bone located at midline of the middle cranial fossa
septal cartilage
flat cartilage structure that forms the anterior portion of the nasal septum
sphenoid bone
unpaired bone that forms the central base of skull
sphenoid sinus
air-filled space located within the sphenoid bone; most posterior of the paranasal sinuses
squamous suture
joint that unites the parietal bone to the squamous portion of the temporal bone on the lateral side of the skull
styloid process
downward projecting, elongated bony process located on the inferior aspect of the skull
stylomastoid foramen
opening located on inferior skull, between the styloid process and mastoid process
superior nasal concha
smallest and most superiorly located of the nasal conchae; formed by the ethmoid bone
superior nuchal line
paired bony lines on the posterior skull that extend laterally from the external occipital protuberance
superior orbital fissure
irregularly shaped opening between the middle cranial fossa and the posterior orbit
supraorbital foramen
opening located on anterior skull, at the superior margin of the orbit
supraorbital margin
superior margin of the orbit
suture
junction line at which adjacent bones of the skull are united by fibrous connective tissue
temporal bone
paired bones that form the lateral, inferior portions of the skull, with squamous, mastoid, and petrous portions
temporal fossa
shallow space on the lateral side of the skull, above the level of the zygomatic arch
temporal process of the zygomatic bone
short extension from the zygomatic bone that forms the anterior portion of the zygomatic arch
vomer bone
unpaired bone that forms the inferior and posterior portions of the nasal septum
zygomatic arch
elongated, free-standing arch on the lateral skull, formed anteriorly by the temporal process of the zygomatic bone and posteriorly by the zygomatic process of the temporal bone
zygomatic bone
cheekbone; paired bones that contribute to the lateral orbit and anterior zygomatic arch
zygomatic process of the temporal bone
extension from the temporal bone that forms the posterior portion of the zygomatic arch
inferior articular process
bony process that extends downward from the vertebral arch of a vertebra that articulates with the superior articular process of the next lower vertebra
intervertebral disc
structure located between the bodies of adjacent vertebrae that strongly joins the vertebrae; provides padding, weight bearing ability, and enables vertebral column movements
intervertebral foramen
opening located between adjacent vertebrae for exit of a spinal nerve
bronchoconstriction
decrease in the size of the bronchiole due to contraction of the muscular wall
bronchodilation
increase in the size of the bronchiole due to contraction of the muscular wall
cardiac notch
indentation on the surface of the left lung that allows space for the heart
hilum
concave structure on the mediastinal surface of the lungs where blood vessels, lymphatic vessels, nerves, and a bronchus enter the lung
lung
organ of the respiratory system that performs gas exchange
parietal pleura
outermost layer of the pleura that connects to the thoracic wall, mediastinum, and diaphragm
pleural cavity
space between the visceral and parietal pleurae
pleural fluid
substance that acts as a lubricant for the visceral and parietal layers of the pleura during the movement of breathing
pulmonary artery
artery that arises from the pulmonary trunk and carries deoxygenated, arterial blood to the alveoli
pulmonary plexus
network of autonomic nervous system fibers found near the hilum of the lung
visceral pleura
innermost layer of the pleura that is superficial to the lungs and extends into the lung fissures
kyphosis
(also, humpback or hunchback) excessive posterior curvature of the thoracic vertebral column region
lamina
portion of the vertebral arch on each vertebra that extends between the transverse and spinous process
lateral sacral crest
paired irregular ridges running down the lateral sides of the posterior sacrum that was formed by the fusion of the transverse processes from the five sacral vertebrae
ligamentum flavum
series of short ligaments that unite the lamina of adjacent vertebrae
lordosis
(also, swayback) excessive anterior curvature of the lumbar vertebral column region
lumbar curve
posteriorly concave curvature of the lumbar vertebral column region; a secondary curve of the vertebral column
lumbar vertebrae
five vertebrae numbered as L1–L5 that are located in lumbar region (lower back) of the vertebral column
median sacral crest
irregular ridge running down the midline of the posterior sacrum that was formed from the fusion of the spinous processes of the five sacral vertebrae
nuchal ligament
expanded portion of the supraspinous ligament within the posterior neck; interconnects the spinous processes of the cervical vertebrae and attaches to the base of the skull
nucleus pulposus
gel-like central region of an intervertebral disc; provides for padding, weight-bearing, and movement between adjacent vertebrae
pedicle
portion of the vertebral arch that extends from the vertebral body to the transverse process
posterior arch
posterior portion of the ring-like C1 (atlas) vertebra
posterior longitudinal ligament
ligament that runs the length of the vertebral column, uniting the posterior sides of the vertebral bodies
posterior (dorsal) sacral foramen
one of the series of paired openings located on the posterior (dorsal) side of the sacrum
primary curve
anteriorly concave curvatures of the thoracic and sacrococcygeal regions that are retained from the original fetal curvature of the vertebral column
lateral pterygoid
muscle that moves the mandible from side to side
longissimus capitis
muscle of the longissimus group associated with the head region
longissimus cervicis
muscle of the longissimus group associated with the cervical region
longissimus group
intermediately placed muscles of the erector spinae
longissimus thoracis
muscle of the longissimus group associated with the thoracic region
masseter
main muscle for chewing that elevates the mandible to close the mouth
mastication
chewing
medial pterygoid
muscle that moves the mandible from side to side
middle scalene
longest scalene muscle, located between the anterior and posterior scalenes
multifidus
muscle of the lumbar region that helps extend and laterally flex the vertebral column
mylohyoid
muscle that lifts the hyoid bone and helps press the tongue to the top of the mouth
occipitalis
posterior part of the occipitofrontalis muscle
occipitofrontalis
muscle that makes up the scalp with a frontal belly and an occipital belly
omohyoid
muscle that has superior and inferior bellies and depresses the hyoid bone
orbicularis oculi
circular muscle that closes the eye
orbicularis oris
circular muscle that moves the lips
palatoglossus
muscle that originates on the soft palate to elevate the back of the tongue
posterior scalene
smallest scalene muscle, located posterior to the middle scalene
scalene muscles
flex, laterally flex, and rotate the head; contribute to deep inhalation
segmental muscle group
interspinales and intertransversarii muscles that bring together the spinous and transverse processes of each consecutive vertebra
semispinalis capitis
transversospinales muscle associated with the head region
semispinalis cervicis
transversospinales muscle associated with the cervical region
semispinalis thoracis
transversospinales muscle associated with the thoracic region
spinalis capitis
muscle of the spinalis group associated with the head region
spinalis cervicis
muscle of the spinalis group associated with the cervical region
spinalis group
medially placed muscles of the erector spinae
spinalis thoracis
muscle of the spinalis group associated with the thoracic region
splenius
posterior neck muscles; includes the splenius capitis and splenius cervicis
splenius capitis
neck muscle that inserts into the head region
splenius cervicis
neck muscle that inserts into the cervical region
sternocleidomastoid
major muscle that laterally flexes and rotates the head
sternohyoid
muscle that depresses the hyoid bone
sternothyroid
muscle that depresses the larynx’s thyroid cartilage
styloglossus
muscle that originates on the styloid bone, and allows upward and backward motion of the tongue
stylohyoid
muscle that elevates the hyoid bone posteriorly
suprahyoid muscles
neck muscles that are superior to the hyoid bone
temporalis
muscle that retracts the mandible
thyrohyoid
muscle that depresses the hyoid bone and elevates the larynx’s thyroid cartilage
transversospinales
muscles that originate at the transverse processes and insert at the spinous processes of the vertebrae
sacral canal
bony tunnel that runs through the sacrum
sacral foramina
series of paired openings for nerve exit located on both the anterior (ventral) and posterior (dorsal) aspects of the sacrum
sacral hiatus
inferior opening and termination of the sacral canal
sacral promontory
anterior lip of the base (superior end) of the sacrum
sacrococcygeal curve
anteriorly concave curvature formed by the sacrum and coccyx; a primary curve of the vertebral column
scoliosis
abnormal lateral curvature of the vertebral column
secondary curve
posteriorly concave curvatures of the cervical and lumbar regions of the vertebral column that develop after the time of birth
spinous process
unpaired bony process that extends posteriorly from the vertebral arch of a vertebra
superior articular process
bony process that extends upward from the vertebral arch of a vertebra that articulates with the inferior articular process of the next higher vertebra
superior articular process of the sacrum
paired processes that extend upward from the sacrum to articulate (join) with the inferior articular processes from the L5 vertebra
supraspinous ligament
ligament that interconnects the spinous processes of the thoracic and lumbar vertebrae
Dalton’s law
statement of the principle that a specific gas type in a mixture exerts its own pressure, as if that specific gas type was not part of a mixture of gases
external respiration
gas exchange that occurs in the alveoli
Henry’s law
statement of the principle that the concentration of gas in a liquid is directly proportional to the solubility and partial pressure of that gas
internal respiration
gas exchange that occurs at the level of body tissues
partial pressure
force exerted by each gas in a mixture of gases
total pressure
sum of all the partial pressures of a gaseous mixture
ventilation
movement of air into and out of the lungs; consists of inspiration and expiration
thoracic curve
anteriorly concave curvature of the thoracic vertebral column region; a primary curve of the vertebral column
thoracic vertebrae
twelve vertebrae numbered as T1–T12 that are located in the thoracic region (upper back) of the vertebral column
transverse foramen
opening found only in the transverse processes of cervical vertebrae
transverse process
paired bony processes that extends laterally from the vertebral arch of a vertebra
vertebral arch
bony arch formed by the posterior portion of each vertebra that surrounds and protects the spinal cord
vertebral (spinal) canal
bony passageway within the vertebral column for the spinal cord that is formed by the series of individual vertebral foramina
vertebral foramen
opening associated with each vertebra defined by the vertebral arch that provides passage for the spinal cord
head of the rib
posterior end of a rib that articulates with the bodies of thoracic vertebrae
jugular (suprasternal) notch
shallow notch located on superior surface of sternal manubrium
manubrium
expanded, superior portion of the sternum
neck of the rib
narrowed region of a rib, next to the rib head
sternal angle
junction line between manubrium and body of the sternum and the site for attachment of the second rib to the sternum
true ribs
vertebrosternal ribs 1–7 that attach via their costal cartilage directly to the sternum
tubercle of the rib
small bump on the posterior side of a rib for articulation with the transverse process of a thoracic vertebra
xiphoid process
small process that forms the inferior tip of the sternum
glenohumeral joint
shoulder joint; formed by the articulation between the glenoid cavity of the scapula and the head of the humerus
glenoid cavity
(also, glenoid fossa) shallow depression located on the lateral scapula, between the superior and lateral borders
lateral border of the scapula
diagonally oriented lateral margin of the scapula
medial border of the scapula
elongated, medial margin of the scapula
pectoral girdle
shoulder girdle; the set of bones, consisting of the scapula and clavicle, which attaches each upper limb to the axial skeleton
scapula
shoulder blade bone located on the posterior side of the shoulder
spine of the scapula
prominent ridge passing mediolaterally across the upper portion of the posterior scapular surface
sternal end of the clavicle
medial end of the clavicle that articulates with the manubrium of the sternum
sternoclavicular joint
articulation between the manubrium of the sternum and the sternal end of the clavicle; forms the only bony attachment between the pectoral girdle of the upper limb and the axial skeleton
subscapular fossa
broad depression located on the anterior (deep) surface of the scapula
superior angle of the scapula
corner of the scapula between the superior and medial borders of the scapula
superior border of the scapula
superior margin of the scapula
supraglenoid tubercle
small bump located at the superior margin of the glenoid cavity
suprascapular notch
small notch located along the superior border of the scapula, medial to the coracoid process
supraspinous fossa
narrow depression located on the posterior scapula, superior to the spine
skeletal muscle
striated, multinucleated muscle that requires signaling from the nervous system to trigger contraction; most skeletal muscles are referred to as voluntary muscles that move bones and produce movement
smooth muscle
nonstriated, mononucleated muscle in the skin that is associated with hair follicles; assists in moving materials in the walls of internal organs, blood vessels, and internal passageways
pisiform
from the lateral side, the fourth of the four proximal carpal bones; articulates with the anterior surface of the triquetrum
pollex
(also, thumb) digit 1 of the hand
proximal radioulnar joint
articulation formed by the radial notch of the ulna and the head of the radius
radial fossa
small depression located on the anterior humerus above the capitulum; this space receives the head of the radius when the elbow is maximally flexed
radial notch of the ulna
small, smooth area on the lateral side of the proximal ulna; articulates with the head of the radius as part of the proximal radioulnar joint
radial tuberosity
oval-shaped, roughened protuberance located on the medial side of the proximal radius
radiocarpal joint
wrist joint, located between the forearm and hand regions of the upper limb; articulation formed proximally by the distal end of the radius and the fibrocartilaginous pad that unites the distal radius and ulna bone, and distally by the scaphoid, lunate, and triquetrum carpal bones
radius
bone located on the lateral side of the forearm
scaphoid
from the lateral side, the first of the four proximal carpal bones; articulates with the radius proximally, the trapezoid, trapezium, and capitate distally, and the lunate medially
shaft of the humerus
narrow, elongated, central region of the humerus
shaft of the radius
narrow, elongated, central region of the radius
shaft of the ulna
narrow, elongated, central region of the ulna
styloid process of the radius
pointed projection located on the lateral end of the distal radius
styloid process of the ulna
short, bony projection located on the medial end of the distal ulna
surgical neck
region of the humerus where the expanded, proximal end joins with the narrower shaft
trapezium
from the lateral side, the first of the four distal carpal bones; articulates with the scaphoid proximally, the first and second metacarpals distally, and the trapezoid medially
trapezoid
from the lateral side, the second of the four distal carpal bones; articulates with the scaphoid proximally, the second metacarpal distally, the trapezium laterally, and the capitate medially
triquetrum
from the lateral side, the third of the four proximal carpal bones; articulates with the lunate laterally, the hamate distally, and has a facet for the pisiform
trochlea
pulley-shaped region located medially at the distal end of the humerus; articulates at the elbow with the trochlear notch of the ulna
trochlear notch
large, C-shaped depression located on the anterior side of the proximal ulna; articulates at the elbow with the trochlea of the humerus
ulna
bone located on the medial side of the forearm
ulnar notch of the radius
shallow, smooth area located on the medial side of the distal radius; articulates with the head of the ulna at the distal radioulnar joint
ulnar tuberosity
roughened area located on the anterior, proximal ulna inferior to the coronoid process
lesser pelvis
(also, lesser pelvic cavity or true pelvis) narrow space located within the pelvis, defined superiorly by the pelvic brim (pelvic inlet) and inferiorly by the pelvic outlet
lesser sciatic foramen
pelvic opening formed by the lesser sciatic notch of the hip bone, the sacrospinous ligament, and the sacrotuberous ligament
lesser sciatic notch
shallow indentation along the posterior margin of the ischium, inferior to the ischial spine
obturator foramen
large opening located in the anterior hip bone, between the pubis and ischium regions
lateral compartment of the leg
region that includes the fibularis (peroneus) longus and the fibularis (peroneus) brevis and their associated blood vessels and nerves
medial compartment of the thigh
a region that includes the adductor longus, adductor brevis, adductor magnus, pectineus, gracilis, and their associated blood vessels and nerves
obturator externus
muscle deep to the gluteus maximus on the lateral surface of the thigh that laterally rotates the femur at the hip
obturator internus
muscle deep to the gluteus maximus on the lateral surface of the thigh that laterally rotates the femur at the hip
patellar ligament
extension of the quadriceps tendon below the patella
pectineus
muscle that abducts and flexes the femur at the hip
pelvic girdle
hips, a foundation for the lower limb
piriformis
muscle deep to the gluteus maximus on the lateral surface of the thigh that laterally rotates the femur at the hip
plantar aponeurosis
muscle that supports the longitudinal arch of the foot
plantar group
four-layered group of intrinsic foot muscles
plantaris
muscle that runs obliquely between the gastrocnemius and the soleus
popliteal fossa
diamond-shaped space at the back of the knee
popliteus
muscle that flexes the leg at the knee and creates the floor of the popliteal fossa
posterior compartment of the leg
region that includes the superficial gastrocnemius, soleus, and plantaris, and the deep popliteus, flexor digitorum longus, flexor hallucis longus, and tibialis posterior
posterior compartment of the thigh
region that includes muscles that flex the leg and extend the thigh
psoas major
muscle that, along with the iliacus, makes up the iliopsoas
quadratus femoris
muscle deep to the gluteus maximus on the lateral surface of the thigh that laterally rotates the femur at the hip
quadriceps femoris group
four muscles, that extend and stabilize the knee
quadriceps tendon
(also, patellar tendon) tendon common to all four quadriceps muscles, inserts into the patella
rectus femoris
quadricep muscle on the anterior aspect of the thigh
sartorius
band-like muscle that flexes, abducts, and laterally rotates the leg at the hip
semimembranosus
hamstring muscle
semitendinosus
hamstring muscle
soleus
wide, flat muscle deep to the gastrocnemius
superior extensor retinaculum
transverse ligament of the ankle
superior gemellus
muscle deep to the gluteus maximus on the lateral surface of the thigh that laterally rotates the femur at the hip
tensor fascia lata
muscle that flexes and abducts the thigh
tibialis anterior
muscle located on the lateral surface of the tibia
tibialis posterior
muscle that plantar flexes and inverts the foot
vastus intermedius
quadricep muscle that is between the vastus lateralis and vastus medialis and is deep to the rectus femoris
vastus lateralis
quadricep muscle on the lateral aspect of the thigh
vastus medialis
quadricep muscle on the medial aspect of the thigh
pectineal line
narrow ridge located on the superior surface of the superior pubic ramus
pelvic brim
pelvic inlet; the dividing line between the greater and lesser pelvic regions; formed by the superior margin of the pubic symphysis, the pectineal lines of each pubis, the arcuate lines of each ilium, and the sacral promontory
pelvic girdle
hip girdle; consists of a single hip bone, which attaches a lower limb to the sacrum of the axial skeleton
pelvic inlet
pelvic brim
pelvic outlet
inferior opening of the lesser pelvis; formed by the inferior margin of the pubic symphysis, right and left ischiopubic rami and sacrotuberous ligaments, and the tip of the coccyx
pelvis
ring of bone consisting of the right and left hip bones, the sacrum, and the coccyx
posterior inferior iliac spine
small, bony projection located at the inferior margin of the auricular surface on the posterior ilium
posterior sacroiliac ligament
strong ligament spanning the sacrum and ilium of the hip bone that supports the posterior side of the sacroiliac joint
posterior superior iliac spine
rounded, posterior end of the iliac crest
pubic arch
bony structure formed by the pubic symphysis, and the bodies and inferior pubic rami of the right and left pubic bones
pubic body
enlarged, medial portion of the pubis region of the hip bone
pubic symphysis
joint formed by the articulation between the pubic bodies of the right and left hip bones
pubic tubercle
small bump located on the superior aspect of the pubic body
pubis
anterior portion of the hip bone
lactic acid
product of anaerobic glycolysis
oxygen debt
amount of oxygen needed to compensate for ATP produced without oxygen during muscle contraction
power stroke
action of myosin pulling actin inward (toward the M line)
afferent lymphatic vessels
lead into a lymph node
antibody
antigen-specific protein secreted by plasma cells; immunoglobulin
antigen
molecule recognized by the receptors of B and T lymphocytes
barrier defenses
antipathogen defenses deriving from a barrier that physically prevents pathogens from entering the body to establish an infection
B cells
lymphocytes that act by differentiating into an antibody-secreting plasma cell
bone marrow
tissue found inside bones; the site of all blood cell differentiation and maturation of B lymphocytes
bronchus-associated lymphoid tissue (BALT)
lymphoid nodule associated with the respiratory tract
chyle
lipid-rich lymph inside the lymphatic capillaries of the small intestine
cisterna chyli
bag-like vessel that forms the beginning of the thoracic duct
efferent lymphatic vessels
lead out of a lymph node
germinal centers
clusters of rapidly proliferating B cells found in secondary lymphoid tissues
high endothelial venules
vessels containing unique endothelial cells specialized to allow migration of lymphocytes from the blood to the lymph node
immune system
series of barriers, cells, and soluble mediators that combine to response to infections of the body with pathogenic organisms
innate immune response
rapid but relatively nonspecific immune response
lymph
fluid contained within the lymphatic system
lymph node
one of the bean-shaped organs found associated with the lymphatic vessels
lymphatic capillaries
smallest of the lymphatic vessels and the origin of lymph flow
lymphatic system
network of lymphatic vessels, lymph nodes, and ducts that carries lymph from the tissues and back to the bloodstream.
lymphatic trunks
large lymphatics that collect lymph from smaller lymphatic vessels and empties into the blood via lymphatic ducts
lymphocytes
white blood cells characterized by a large nucleus and small rim of cytoplasm
lymphoid nodules
unencapsulated patches of lymphoid tissue found throughout the body
mucosa-associated lymphoid tissue (MALT)
lymphoid nodule associated with the mucosa
naïve lymphocyte
mature B or T cell that has not yet encountered antigen for the first time
natural killer cell (NK)
cytotoxic lymphocyte of innate immune response
plasma cell
differentiated B cell that is actively secreting antibody
primary lymphoid organ
site where lymphocytes mature and proliferate; red bone marrow and thymus gland
right lymphatic duct
drains lymph fluid from the upper right side of body into the right subclavian vein
secondary lymphoid organs
sites where lymphocytes mount adaptive immune responses; examples include lymph nodes and spleen
spleen
secondary lymphoid organ that filters pathogens from the blood (white pulp) and removes degenerating or damaged blood cells (red pulp)
T cell
lymphocyte that acts by secreting molecules that regulate the immune system or by causing the destruction of foreign cells, viruses, and cancer cells
thoracic duct
large duct that drains lymph from the lower limbs, left thorax, left upper limb, and the left side of the head
thymocyte
immature T cell found in the thymus
thymus
primary lymphoid organ; where T lymphocytes proliferate and mature
tonsils
lymphoid nodules associated with the nasopharynx
pyruvic acid
product of glycolysis that can be used in aerobic respiration or converted to lactic acid
sacroiliac joint
joint formed by the articulation between the auricular surfaces of the sacrum and ilium
sacrospinous ligament
ligament that spans the sacrum to the ischial spine of the hip bone
sacrotuberous ligament
ligament that spans the sacrum to the ischial tuberosity of the hip bone
subpubic angle
inverted V-shape formed by the convergence of the right and left ischiopubic rami; this angle is greater than 80 degrees in females and less than 70 degrees in males
superior pubic ramus
narrow segment of bone that passes laterally from the pubic body to join the ilium
knee joint
joint that separates the thigh and leg portions of the lower limb; formed by the articulations between the medial and lateral condyles of the femur, and the medial and lateral condyles of the tibia
shaft of the femur
cylindrically shaped region that forms the central portion of the femur
shaft of the fibula
elongated, slender portion located between the expanded ends of the fibula
shaft of the tibia
triangular-shaped, central portion of the tibia
soleal line
small, diagonally running ridge located on the posterior side of the proximal tibia
sustentaculum tali
bony ledge extending from the medial side of the calcaneus bone
talus
tarsal bone that articulates superiorly with the tibia and fibula at the ankle joint; also articulates inferiorly with the calcaneus bone and anteriorly with the navicular bone
tarsal bone
one of the seven bones that make up the posterior foot; includes the calcaneus, talus, navicular, cuboid, medial cuneiform, intermediate cuneiform, and lateral cuneiform bones
thigh
portion of the lower limb located between the hip and knee joints
tibia
shin bone; the large, weight-bearing bone located on the medial side of the leg
tibial tuberosity
elevated area on the anterior surface of the proximal tibia
apical ectodermal ridge
enlarged ridge of ectoderm at the distal end of a limb bud that stimulates growth and elongation of the limb
limb bud
small elevation that appears on the lateral side of the embryo during the fourth or fifth week of development, which gives rise to an upper or lower limb
negative feedback
homeostatic mechanism that tends to stabilize an upset in the body’s physiological condition by preventing an excessive response to a stimulus, typically as the stimulus is removed
normal range
range of values around the set point that do not cause a reaction by the control center
positive feedback
mechanism that intensifies a change in the body’s physiological condition in response to a stimulus
sensor
(also, receptor) reports a monitored physiological value to the control center
set point
ideal value for a physiological parameter; the level or small range within which a physiological parameter such as blood pressure is stable and optimally healthful, that is, within its parameters of homeostasis
levator ani
pelvic muscle that resists intra-abdominal pressure and supports the pelvic viscera
linea alba
white, fibrous band that runs along the midline of the trunk
pelvic diaphragm
muscular sheet that comprises the levator ani and the ischiococcygeus
perineum
diamond-shaped region between the pubic symphysis, coccyx, and ischial tuberosities
pubococcygeus
muscle that makes up the levator ani along with the iliococcygeus
quadratus lumborum
posterior part of the abdominal wall that helps with posture and stabilization of the body
rectus abdominis
long, linear muscle that extends along the middle of the trunk
rectus sheaths
tissue that makes up the linea alba
sphincter urethrovaginalis
deep perineal muscle in women
tendinous intersections
three transverse bands of collagen fibers that divide the rectus abdominis into segments
transversus abdominis
deep layer of the abdomen that has fascicles arranged transversely around the abdomen
urogenital triangle
anterior triangle of the perineum that includes the external genitals
lateral meniscus
C-shaped fibrocartilage articular disc located at the knee, between the lateral condyle of the femur and the lateral condyle of the tibia
lateral tibiofemoral joint
portion of the knee consisting of the articulation between the lateral condyle of the tibia and the lateral condyle of the femur; allows for flexion/extension at the knee
ligament of the head of the femur
intracapsular ligament that runs from the acetabulum of the hip bone to the head of the femur
medial meniscus
C-shaped fibrocartilage articular disc located at the knee, between the medial condyle of the femur and medial condyle of the tibia
medial tibiofemoral joint
portion of the knee consisting of the articulation between the medial condyle of the tibia and the medial condyle of the femur; allows for flexion/extension at the knee
patellar ligament
ligament spanning from the patella to the anterior tibia; serves as the final attachment for the quadriceps femoris muscle
posterior cruciate ligament
intracapsular ligament of the knee; extends from the posterior, superior surface of the tibia to the inner aspect of the medial condyle of the femur; prevents anterior displacement of the femur when the knee is flexed and weight bearing
posterior talofibular ligament
intrinsic ligament located on the lateral side of the ankle joint, between the talus bone and lateral malleolus of the fibula; supports the talus at the talocrural joint and resists excess inversion of the foot
pubofemoral ligament
intrinsic ligament spanning from the pubis of the hip bone to the femur, on the anterior-inferior aspect of the hip joint
radial collateral ligament
intrinsic ligament on the lateral side of the elbow joint; runs from the lateral epicondyle of humerus to merge with the annular ligament
rotator cuff
strong connective tissue structure formed by the fusion of four rotator cuff muscle tendons to the articular capsule of the shoulder joint; surrounds and supports superior, anterior, lateral, and posterior sides of the humeral head
subacromial bursa
bursa that protects the supraspinatus muscle tendon and superior end of the humerus from rubbing against the acromion of the scapula
subscapular bursa
bursa that prevents rubbing of the subscapularis muscle tendon against the scapula
subtalar joint
articulation between the talus and calcaneus bones of the foot; allows motions that contribute to inversion/eversion of the foot
talocrural joint
ankle joint; articulation between the talus bone of the foot and medial malleolus of the tibia, distal tibia, and lateral malleolus of the fibula; a uniaxial hinge joint that allows only for dorsiflexion and plantar flexion of the foot
temporomandibular joint (TMJ)
articulation between the condyle of the mandible and the mandibular fossa and articular tubercle of the temporal bone of the skull; allows for depression/elevation (opening/closing of mouth), protraction/retraction, and side-to-side motions of the mandible
tibial collateral ligament
extrinsic ligament of knee joint that spans from the medial epicondyle of the femur to the medial tibia; resists hyperextension and rotation of extended knee
ulnar collateral ligament
intrinsic ligament on the medial side of the elbow joint; spans from the medial epicondyle of the humerus to the medial ulna
zygapophysial joints
facet joints; plane joints between the superior and inferior articular processes of adjacent vertebrae that provide for only limited motions between the vertebrae
insertion
end of a skeletal muscle that is attached to the structure (usually a bone) that is moved when the muscle contracts
multipennate
pennate muscle that has a tendon branching within it
origin
end of a skeletal muscle that is attached to another structure (usually a bone) in a fixed position
parallel
fascicles that extend in the same direction as the long axis of the muscle
pennate
fascicles that are arranged differently based on their angles to the tendon
prime mover
(also, agonist) principle muscle involved in an action
synergist
muscle whose contraction helps a prime mover in an action
unipennate
pennate muscle that has fascicles located on one side of the tendon
motor end-plate
sarcolemma of muscle fiber at the neuromuscular junction, with receptors for the neurotransmitter acetylcholine
myofibril
long, cylindrical organelle that runs parallel within the muscle fiber and contains the sarcomeres
myosin
protein that makes up most of the thick cylindrical myofilament within a sarcomere muscle fiber
neuromuscular junction (NMJ)
synapse between the axon terminal of a motor neuron and the section of the membrane of a muscle fiber with receptors for the acetylcholine released by the terminal
neurotransmitter
signaling chemical released by nerve terminals that bind to and activate receptors on target cells
motor end-plate
sarcolemma of muscle fiber at the neuromuscular junction, with receptors for the neurotransmitter acetylcholine
myofibril
long, cylindrical organelle that runs parallel within the muscle fiber and contains the sarcomeres
myosin
protein that makes up most of the thick cylindrical myofilament within a sarcomere muscle fiber
neuromuscular junction (NMJ)
synapse between the axon terminal of a motor neuron and the section of the membrane of a muscle fiber with receptors for the acetylcholine released by the terminal
neurotransmitter
signaling chemical released by nerve terminals that bind to and activate receptors on target cells
perimysium
connective tissue that bundles skeletal muscle fibers into fascicles within a skeletal muscle
sarcomere
longitudinally, repeating functional unit of skeletal muscle, with all of the contractile and associated proteins involved in contraction
sarcolemma
plasma membrane of a skeletal muscle fiber
sarcoplasm
cytoplasm of a muscle cell
sarcoplasmic reticulum (SR)
specialized smooth endoplasmic reticulum, which stores, releases, and retrieves Ca++
synaptic cleft
space between a nerve (axon) terminal and a motor end-plate
T-tubule
projection of the sarcolemma into the interior of the cell
thick filament
the thick myosin strands and their multiple heads projecting from the center of the sarcomere toward, but not all to way to, the Z-discs
thin filament
thin strands of actin and its troponin-tropomyosin complex projecting from the Z-discs toward the center of the sarcomere
triad
the grouping of one T-tubule and two terminal cisternae
troponin
regulatory protein that binds to actin, tropomyosin, and calcium
tropomyosin
regulatory protein that covers myosin-binding sites to prevent actin from binding to myosin
voltage-gated sodium channels
membrane proteins that open sodium channels in response to a sufficient voltage change, and initiate and transmit the action potential as Na+ enters through the channel
perimysium
connective tissue that bundles skeletal muscle fibers into fascicles within a skeletal muscle
sarcomere
longitudinally, repeating functional unit of skeletal muscle, with all of the contractile and associated proteins involved in contraction
sarcolemma
plasma membrane of a skeletal muscle fiber
sarcoplasm
cytoplasm of a muscle cell
sarcoplasmic reticulum (SR)
specialized smooth endoplasmic reticulum, which stores, releases, and retrieves Ca++
synaptic cleft
space between a nerve (axon) terminal and a motor end-plate
T-tubule
projection of the sarcolemma into the interior of the cell
thick filament
the thick myosin strands and their multiple heads projecting from the center of the sarcomere toward, but not all to way to, the Z-discs
thin filament
thin strands of actin and its troponin-tropomyosin complex projecting from the Z-discs toward the center of the sarcomere
triad
the grouping of one T-tubule and two terminal cisternae
troponin
regulatory protein that binds to actin, tropomyosin, and calcium
tropomyosin
regulatory protein that covers myosin-binding sites to prevent actin from binding to myosin
voltage-gated sodium channels
membrane proteins that open sodium channels in response to a sufficient voltage change, and initiate and transmit the action potential as Na+ enters through the channel
universal donor
individual with type O− blood
universal recipient
individual with type AB+ blood
abdominal aorta
portion of the aorta inferior to the aortic hiatus and superior to the common iliac arteries
adrenal artery
branch of the abdominal aorta; supplies blood to the adrenal (suprarenal) glands
adrenal vein
drains the adrenal or suprarenal glands that are immediately superior to the kidneys; the right adrenal vein enters the inferior vena cava directly and the left adrenal vein enters the left renal vein
anterior cerebral artery
arises from the internal carotid artery; supplies the frontal lobe of the cerebrum
anterior communicating artery
anastomosis of the right and left internal carotid arteries; supplies blood to the brain
anterior tibial artery
branches from the popliteal artery; supplies blood to the anterior tibial region; becomes the dorsalis pedis artery
anterior tibial vein
forms from the dorsal venous arch; drains the area near the tibialis anterior muscle and leads to the popliteal vein
aorta
largest artery in the body, originating from the left ventricle and descending to the abdominal region where it bifurcates into the common iliac arteries at the level of the fourth lumbar vertebra; arteries originating from the aorta distribute blood to virtually all tissues of the body
aortic arch
arc that connects the ascending aorta to the descending aorta; ends at the intervertebral disk between the fourth and fifth thoracic vertebrae
aortic hiatus
opening in the diaphragm that allows passage of the thoracic aorta into the abdominal region where it becomes the abdominal aorta
arterial circle
(also, circle of Willis) anastomosis located at the base of the brain that ensures continual blood supply; formed from branches of the internal carotid and vertebral arteries; supplies blood to the brain
ascending aorta
initial portion of the aorta, rising from the left ventricle for a distance of approximately 5 cm
axillary artery
continuation of the subclavian artery as it penetrates the body wall and enters the axillary region; supplies blood to the region near the head of the humerus (humeral circumflex arteries); the majority of the vessel continues into the brachium and becomes the brachial artery
axillary vein
major vein in the axillary region; drains the upper limb and becomes the subclavian vein
azygos vein
originates in the lumbar region and passes through the diaphragm into the thoracic cavity on the right side of the vertebral column; drains blood from the intercostal veins, esophageal veins, bronchial veins, and other veins draining the mediastinal region; leads to the superior vena cava
basilar artery
formed from the fusion of the two vertebral arteries; sends branches to the cerebellum, brain stem, and the posterior cerebral arteries; the main blood supply to the brain stem
basilic vein
superficial vein of the arm that arises from the palmar venous arches, intersects with the median cubital vein, parallels the ulnar vein, and continues into the upper arm; along with the brachial vein, it leads to the axillary vein
brachial artery
continuation of the axillary artery in the brachium; supplies blood to much of the brachial region; gives off several smaller branches that provide blood to the posterior surface of the arm in the region of the elbow; bifurcates into the radial and ulnar arteries at the coronoid fossa
brachial vein
deeper vein of the arm that forms from the radial and ulnar veins in the lower arm; leads to the axillary vein
brachiocephalic artery
single vessel located on the right side of the body; the first vessel branching from the aortic arch; gives rise to the right subclavian artery and the right common carotid artery; supplies blood to the head, neck, upper limb, and wall of the thoracic region
brachiocephalic vein
one of a pair of veins that form from a fusion of the external and internal jugular veins and the subclavian vein; subclavian, external and internal jugulars, vertebral, and internal thoracic veins lead to it; drains the upper thoracic region and flows into the superior vena cava
bronchial artery
systemic branch from the aorta that provides oxygenated blood to the lungs in addition to the pulmonary circuit
bronchial vein
drains the systemic circulation from the lungs and leads to the azygos vein
cavernous sinus
enlarged vein that receives blood from most of the other cerebral veins and the eye socket, and leads to the petrosal sinus
celiac trunk
(also, celiac artery) major branch of the abdominal aorta; gives rise to the left gastric artery, the splenic artery, and the common hepatic artery that forms the hepatic artery to the liver, the right gastric artery to the stomach, and the cystic artery to the gall bladder
cephalic vein
superficial vessel in the upper arm; leads to the axillary vein
cerebrovascular accident (CVA)
blockage of blood flow to the brain; also called a stroke
circle of Willis
(also, arterial circle) anastomosis located at the base of the brain that ensures continual blood supply; formed from branches of the internal carotid and vertebral arteries; supplies blood to the brain
common carotid artery
right common carotid artery arises from the brachiocephalic artery, and the left common carotid arises from the aortic arch; gives rise to the external and internal carotid arteries; supplies the respective sides of the head and neck
common hepatic artery
branch of the celiac trunk that forms the hepatic artery, the right gastric artery, and the cystic artery
common iliac artery
branch of the aorta that leads to the internal and external iliac arteries
common iliac vein
one of a pair of veins that flows into the inferior vena cava at the level of L5; the left common iliac vein drains the sacral region; divides into external and internal iliac veins near the inferior portion of the sacroiliac joint
cystic artery
branch of the common hepatic artery; supplies blood to the gall bladder
deep femoral artery
branch of the femoral artery; gives rise to the lateral circumflex arteries
deep femoral vein
drains blood from the deeper portions of the thigh and leads to the femoral vein
descending aorta
portion of the aorta that continues downward past the end of the aortic arch; subdivided into the thoracic aorta and the abdominal aorta
digital arteries
formed from the superficial and deep palmar arches; supply blood to the digits
digital veins
drain the digits and feed into the palmar arches of the hand and dorsal venous arch of the foot
dorsal arch
(also, arcuate arch) formed from the anastomosis of the dorsalis pedis artery and medial and plantar arteries; branches supply the distal portions of the foot and digits
dorsal venous arch
drains blood from digital veins and vessels on the superior surface of the foot
dorsalis pedis artery
forms from the anterior tibial artery; branches repeatedly to supply blood to the tarsal and dorsal regions of the foot
esophageal artery
branch of the thoracic aorta; supplies blood to the esophagus
esophageal vein
drains the inferior portions of the esophagus and leads to the azygos vein
external carotid artery
arises from the common carotid artery; supplies blood to numerous structures within the face, lower jaw, neck, esophagus, and larynx
external iliac artery
branch of the common iliac artery that leaves the body cavity and becomes a femoral artery; supplies blood to the lower limbs
external iliac vein
formed when the femoral vein passes into the body cavity; drains the legs and leads to the common iliac vein
external jugular vein
one of a pair of major veins located in the superficial neck region that drains blood from the more superficial portions of the head, scalp, and cranial regions, and leads to the subclavian vein
femoral artery
continuation of the external iliac artery after it passes through the body cavity; divides into several smaller branches, the lateral deep femoral artery, and the genicular artery; becomes the popliteal artery as it passes posterior to the knee
femoral circumflex vein
forms a loop around the femur just inferior to the trochanters; drains blood from the areas around the head and neck of the femur; leads to the femoral vein
femoral vein
drains the upper leg; receives blood from the great saphenous vein, the deep femoral vein, and the femoral circumflex vein; becomes the external iliac vein when it crosses the body wall
fibular vein
drains the muscles and integument near the fibula and leads to the popliteal vein
genicular artery
branch of the femoral artery; supplies blood to the region of the knee
gonadal artery
branch of the abdominal aorta; supplies blood to the gonads or reproductive organs; also described as ovarian arteries or testicular arteries, depending upon the sex of the individual
gonadal vein
generic term for a vein draining a reproductive organ; may be either an ovarian vein or a testicular vein, depending on the sex of the individual
great cerebral vein
receives most of the smaller vessels from the inferior cerebral veins and leads to the straight sinus
great saphenous vein
prominent surface vessel located on the medial surface of the leg and thigh; drains the superficial portions of these areas and leads to the femoral vein
hemiazygos vein
smaller vein complementary to the azygos vein; drains the esophageal veins from the esophagus and the left intercostal veins, and leads to the brachiocephalic vein via the superior intercostal vein
hepatic artery proper
branch of the common hepatic artery; supplies systemic blood to the liver
hepatic portal system
specialized circulatory pathway that carries blood from digestive organs to the liver for processing before being sent to the systemic circulation
hepatic vein
drains systemic blood from the liver and flows into the inferior vena cava
inferior mesenteric artery
branch of the abdominal aorta; supplies blood to the distal segment of the large intestine and rectum
inferior phrenic artery
branch of the abdominal aorta; supplies blood to the inferior surface of the diaphragm
inferior vena cava
large systemic vein that drains blood from areas largely inferior to the diaphragm; empties into the right atrium
intercostal artery
branch of the thoracic aorta; supplies blood to the muscles of the thoracic cavity and vertebral column
intercostal vein
drains the muscles of the thoracic wall and leads to the azygos vein
internal carotid artery
arises from the common carotid artery and begins with the carotid sinus; goes through the carotid canal of the temporal bone to the base of the brain; combines with branches of the vertebral artery forming the arterial circle; supplies blood to the brain
internal iliac artery
branch from the common iliac arteries; supplies blood to the urinary bladder, walls of the pelvis, external genitalia, and the medial portion of the femoral region; in females, also provide blood to the uterus and vagina
internal iliac vein
drains the pelvic organs and integument; formed from several smaller veins in the region; leads to the common iliac vein
internal jugular vein
one of a pair of major veins located in the neck region that passes through the jugular foramen and canal, flows parallel to the common carotid artery that is more or less its counterpart; primarily drains blood from the brain, receives the superficial facial vein, and empties into the subclavian vein
internal thoracic artery
(also, mammary artery) arises from the subclavian artery; supplies blood to the thymus, pericardium of the heart, and the anterior chest wall
internal thoracic vein
(also, internal mammary vein) drains the anterior surface of the chest wall and leads to the brachiocephalic vein
lateral circumflex artery
branch of the deep femoral artery; supplies blood to the deep muscles of the thigh and the ventral and lateral regions of the integument
lateral plantar artery
arises from the bifurcation of the posterior tibial arteries; supplies blood to the lateral plantar surfaces of the foot
left gastric artery
branch of the celiac trunk; supplies blood to the stomach
lumbar arteries
branches of the abdominal aorta; supply blood to the lumbar region, the abdominal wall, and spinal cord
lumbar veins
drain the lumbar portion of the abdominal wall and spinal cord; the superior lumbar veins drain into the azygos vein on the right or the hemiazygos vein on the left; blood from these vessels is returned to the superior vena cava rather than the inferior vena cava
maxillary vein
drains blood from the maxillary region and leads to the external jugular vein
medial plantar artery
arises from the bifurcation of the posterior tibial arteries; supplies blood to the medial plantar surfaces of the foot
median antebrachial vein
vein that parallels the ulnar vein but is more medial in location; intertwines with the palmar venous arches
median cubital vein
superficial vessel located in the antecubital region that links the cephalic vein to the basilic vein in the form of a v; a frequent site for a blood draw
median sacral artery
continuation of the aorta into the sacrum
mediastinal artery
branch of the thoracic aorta; supplies blood to the mediastinum
middle cerebral artery
another branch of the internal carotid artery; supplies blood to the temporal and parietal lobes of the cerebrum
middle sacral vein
drains the sacral region and leads to the left common iliac vein
occipital sinus
enlarged vein that drains the occipital region near the falx cerebelli and flows into the left and right transverse sinuses, and also into the vertebral veins
ophthalmic artery
branch of the internal carotid artery; supplies blood to the eyes
ovarian artery
branch of the abdominal aorta; supplies blood to the ovary, uterine (Fallopian) tube, and uterus
ovarian vein
drains the ovary; the right ovarian vein leads to the inferior vena cava and the left ovarian vein leads to the left renal vein
palmar arches
superficial and deep arches formed from anastomoses of the radial and ulnar arteries; supply blood to the hand and digital arteries
palmar venous arches
drain the hand and digits, and feed into the radial and ulnar veins
parietal branches
(also, somatic branches) group of arterial branches of the thoracic aorta; includes those that supply blood to the thoracic cavity, vertebral column, and the superior surface of the diaphragm
pericardial artery
branch of the thoracic aorta; supplies blood to the pericardium
petrosal sinus
enlarged vein that receives blood from the cavernous sinus and flows into the internal jugular vein
phrenic vein
drains the diaphragm; the right phrenic vein flows into the inferior vena cava and the left phrenic vein leads to the left renal vein
plantar arch
formed from the anastomosis of the dorsalis pedis artery and medial and plantar arteries; branches supply the distal portions of the foot and digits
plantar veins
drain the foot and lead to the plantar venous arch
plantar venous arch
formed from the plantar veins; leads to the anterior and posterior tibial veins through anastomoses
popliteal artery
continuation of the femoral artery posterior to the knee; branches into the anterior and posterior tibial arteries
popliteal vein
continuation of the femoral vein behind the knee; drains the region behind the knee and forms from the fusion of the fibular and anterior and posterior tibial veins
posterior cerebral artery
branch of the basilar artery that forms a portion of the posterior segment of the arterial circle; supplies blood to the posterior portion of the cerebrum and brain stem
posterior communicating artery
branch of the posterior cerebral artery that forms part of the posterior portion of the arterial circle; supplies blood to the brain
posterior tibial artery
branch from the popliteal artery that gives rise to the fibular or peroneal artery; supplies blood to the posterior tibial region
posterior tibial vein
forms from the dorsal venous arch; drains the area near the posterior surface of the tibia and leads to the popliteal vein
pulmonary artery
one of two branches, left and right, that divides off from the pulmonary trunk and leads to smaller arterioles and eventually to the pulmonary capillaries
pulmonary circuit
system of blood vessels that provide gas exchange via a network of arteries, veins, and capillaries that run from the heart, through the body, and back to the lungs
pulmonary trunk
single large vessel exiting the right ventricle that divides to form the right and left pulmonary arteries
pulmonary veins
two sets of paired vessels, one pair on each side, that are formed from the small venules leading away from the pulmonary capillaries that flow into the left atrium
radial artery
formed at the bifurcation of the brachial artery; parallels the radius; gives off smaller branches until it reaches the carpal region where it fuses with the ulnar artery to form the superficial and deep palmar arches; supplies blood to the lower arm and carpal region
radial vein
parallels the radius and radial artery; arises from the palmar venous arches and leads to the brachial vein
renal artery
branch of the abdominal aorta; supplies each kidney
renal vein
largest vein entering the inferior vena cava; drains the kidneys and leads to the inferior vena cava
right gastric artery
branch of the common hepatic artery; supplies blood to the stomach
sigmoid sinuses
enlarged veins that receive blood from the transverse sinuses; flow through the jugular foramen and into the internal jugular vein
small saphenous vein
located on the lateral surface of the leg; drains blood from the superficial regions of the lower leg and foot, and leads to the popliteal vein
splenic artery
branch of the celiac trunk; supplies blood to the spleen
straight sinus
enlarged vein that drains blood from the brain; receives most of the blood from the great cerebral vein and flows into the left or right transverse sinus
subclavian artery
right subclavian arises from the brachiocephalic artery, whereas the left subclavian artery arises from the aortic arch; gives rise to the internal thoracic, vertebral, and thyrocervical arteries; supplies blood to the arms, chest, shoulders, back, and central nervous system
subclavian vein
located deep in the thoracic cavity; becomes the axillary vein as it enters the axillary region; drains the axillary and smaller local veins near the scapular region; leads to the brachiocephalic vein
subscapular vein
drains blood from the subscapular region and leads to the axillary vein
superior mesenteric artery
branch of the abdominal aorta; supplies blood to the small intestine (duodenum, jejunum, and ileum), the pancreas, and a majority of the large intestine
superior phrenic artery
branch of the thoracic aorta; supplies blood to the superior surface of the diaphragm
superior sagittal sinus
enlarged vein located midsagittally between the meningeal and periosteal layers of the dura mater within the falx cerebri; receives most of the blood drained from the superior surface of the cerebrum and leads to the inferior jugular vein and the vertebral vein
superior vena cava
large systemic vein; drains blood from most areas superior to the diaphragm; empties into the right atrium
temporal vein
drains blood from the temporal region and leads to the external jugular vein
testicular artery
branch of the abdominal aorta; will ultimately travel outside the body cavity to the testes and form one component of the spermatic cord
testicular vein
drains the testes and forms part of the spermatic cord; the right testicular vein empties directly into the inferior vena cava and the left testicular vein empties into the left renal vein
thoracic aorta
portion of the descending aorta superior to the aortic hiatus
thyrocervical artery
arises from the subclavian artery; supplies blood to the thyroid, the cervical region, the upper back, and shoulder
transient ischemic attack (TIA)
temporary loss of neurological function caused by a brief interruption in blood flow; also known as a mini-stroke
transverse sinuses
pair of enlarged veins near the lambdoid suture that drain the occipital, sagittal, and straight sinuses, and leads to the sigmoid sinuses
trunk
large vessel that gives rise to smaller vessels
ulnar artery
formed at the bifurcation of the brachial artery; parallels the ulna; gives off smaller branches until it reaches the carpal region where it fuses with the radial artery to form the superficial and deep palmar arches; supplies blood to the lower arm and carpal region
ulnar vein
parallels the ulna and ulnar artery; arises from the palmar venous arches and leads to the brachial vein
vertebral artery
arises from the subclavian artery and passes through the vertebral foramen through the foramen magnum to the brain; joins with the internal carotid artery to form the arterial circle; supplies blood to the brain and spinal cord
vertebral vein
arises from the base of the brain and the cervical region of the spinal cord; passes through the intervertebral foramina in the cervical vertebrae; drains smaller veins from the cranium, spinal cord, and vertebrae, and leads to the brachiocephalic vein; counterpart of the vertebral artery
visceral branches
branches of the descending aorta that supply blood to the viscera
]]> 2827 0 3 0 1203 Glossary https://pressbooks.bccampus.ca/apdouglas2017/back-matter/1203-glossary/ Wed, 02 Aug 2017 23:32:23 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=back-matter&p=2828
antrum
fluid-filled chamber that characterizes a mature tertiary (antral) follicle
abducens nerve
sixth cranial nerve; responsible for contraction of one of the extraocular muscles
absolute refractory period
time during an action period when another action potential cannot be generated because the voltage-gated Na+ channel is inactivated
absorption
passage of digested products from the intestinal lumen through mucosal cells and into the bloodstream or lacteals
absorptive state
also called the fed state; the metabolic state occurring during the first few hours after ingesting food in which the body is digesting food and absorbing the nutrients
accessory digestive organ
includes teeth, tongue, salivary glands, gallbladder, liver, and pancreas
accessory duct
(also, duct of Santorini) duct that runs from the pancreas into the duodenum
acetyl coenzyme A (acetyl CoA)
starting molecule of the Krebs cycle
acetylcholine (ACh)
neurotransmitter that binds at a motor end-plate to trigger depolarization
acid
compound that releases hydrogen ions in solution
 
acinus
cluster of glandular epithelial cells in the pancreas that secretes pancreatic juice in the pancreas
acromegaly
disorder in adults caused when abnormally high levels of GH trigger growth of bones in the face, hands, and feet
acrosome
cap-like vesicle located at the anterior-most region of a sperm that is rich with lysosomal enzymes capable of digesting the protective layers surrounding the oocyte
acrosomal reaction
release of digestive enzymes by sperm that enables them to burrow through the corona radiata and penetrate the zona pellucida of an oocyte prior to fertilization
action potential change in voltage of a cell membrane in response to a stimulus that results in transmission of an electrical signal; unique to neurons and muscle fibers
activation gate
part of the voltage-gated Na+ channel that opens when the membrane voltage reaches threshold
adenylyl cyclase
membrane-bound enzyme that converts ATP to cyclic AMP, creating cAMP, as a result of G-protein activation
adenosine triphosphate (ATP)
nucleotide containing ribose and an adenine base that is essential in energy transfer
adrenal cortex
outer region of the adrenal glands consisting of multiple layers of epithelial cells and capillary networks that produces mineralocorticoids and glucocorticoids
adrenal glands
endocrine glands located at the top of each kidney that are important for the regulation of the stress response, blood pressure and blood volume, water homeostasis, and electrolyte levels
adrenal medulla
inner layer of the adrenal glands that plays an important role in the stress response by producing epinephrine and norepinephrine
adrenal medulla
interior portion of the adrenal (or suprarenal) gland that releases epinephrine and norepinephrine into the bloodstream as hormones
adrenergic
synapse where norepinephrine is released, which binds to α- or β-adrenergic receptors
adrenocorticotropic hormone (ACTH) anterior pituitary hormone that stimulates the adrenal cortex to secrete corticosteroid hormones (also called corticotropin) alar plate developmental region of the spinal cord that gives rise to the posterior horn of the gray matter
alimentary canal
continuous muscular digestive tube that extends from the mouth to the anus
alkaloid
substance, usually from a plant source, that is chemically basic with respect to pH and will stimulate bitter receptors
alpha (α)-adrenergic receptor
one of the receptors to which epinephrine and norepinephrine bind, which comes in three subtypes: α1, α2, and α3
alarm reaction
the short-term stress, or the fight-or-flight response, of stage one of the general adaptation syndrome mediated by the hormones epinephrine and norepinephrine
aldosterone
hormone produced and secreted by the adrenal cortex that stimulates sodium and fluid retention and increases blood volume and blood pressure
allantois
finger-like outpocketing of yolk sac forms the primitive excretory duct of the embryo; precursor to the urinary bladder
allele
alternative forms of a gene that occupy a specific locus on a specific gene
alpha cell
pancreatic islet cell type that produces the hormone glucagon
alveoli
(of the breast) milk-secreting cells in the mammary gland
amacrine cell
type of cell in the retina that connects to the bipolar cells near the outer synaptic layer and provides the basis for early image processing within the retina
amino acid
building block of proteins; characterized by an amino and carboxyl functional groups and a variable side-chain
aminopeptidase
brush border enzyme that acts on proteins
amnion
transparent membranous sac that encloses the developing fetus and fills with amniotic fluid
amniotic cavity
cavity that opens up between the inner cell mass and the trophoblast; develops into amnion
ampulla
in the ear, the structure at the base of a semicircular canal that contains the hair cells and cupula for transduction of rotational movement of the head
ampulla
(of the uterine tube) middle portion of the uterine tube in which fertilization often occurs
amygdala
nucleus deep in the temporal lobe of the cerebrum that is related to memory and emotional behavior
anabolic hormones
hormones that stimulate the synthesis of new, larger molecules
anabolic reactions
reactions that build smaller molecules into larger molecules
anal canal
final segment of the large intestine
anal column
long fold of mucosa in the anal canal
anal sinus
recess between anal columns
anaphase
third stage of mitosis (and meiosis), during which sister chromatids separate into two new nuclear regions of a dividing cell
anatomical sphincter
smooth or skeletal muscle surrounding the lumen of a vessel or hollow organ that can restrict flow when contracted
anchoring junction
mechanically attaches adjacent cells to each other or to the basement membrane
angiotensin-converting enzyme (ACE)
enzyme produced by the lungs that catalyzes the reaction of inactive angiotensin I into active angiotensin II
angiotensin I
protein produced by the enzymatic action of renin on angiotensinogen; inactive precursor of angiotensin II
angiotensin II
protein produced by the enzymatic action of ACE on inactive angiotensin I; actively causes vasoconstriction and stimulates aldosterone release by the adrenal cortex
angiotensin-converting enzyme
the enzyme that converts angiotensin I to angiotensin II
angiotensinogen
inactive protein in the circulation produced by the liver; precursor of angiotensin I; must be modified by the enzymes renin and ACE to be activated
anion
atom with a negative charge
anosmia
loss of the sense of smell; usually the result of physical disruption of the first cranial nerve
anterior column
white matter between the anterior horns of the spinal cord composed of many different groups of axons of both ascending and descending tracts
anterior horn
gray matter of the spinal cord containing multipolar motor neurons, sometimes referred to as the ventral horn
anterior median fissure
deep midline feature of the anterior spinal cord, marking the separation between the right and left sides of the cord
anterior spinal artery
blood vessel from the merged branches of the vertebral arteries that runs along the anterior surface of the spinal cord
antidiuretic hormone (ADH)
also known as vasopressin, a hormone that increases the volume of water reabsorbed from the collecting tubules of the kidney
antidiuretic hormone (ADH)
hypothalamic hormone that is stored by the posterior pituitary and that signals the kidneys to reabsorb water
antrum
fluid-filled chamber that characterizes a mature tertiary (antral) follicle
anuria
absence of urine produced; production of 50 mL or less per day
apical
that part of a cell or tissue which, in general, faces an open space
apocrine secretion
release of a substance along with the apical portion of the cell
appendix
(vermiform appendix) coiled tube attached to the cecum
aquaporin
protein-forming water channels through the lipid bilayer of the cell; allows water to cross; activation in the collecting ducts is under the control of ADH
aqueous humor
watery fluid that fills the anterior chamber containing the cornea, iris, ciliary body, and lens of the eye
arachnoid granulation
outpocket of the arachnoid membrane into the dural sinuses that allows for reabsorption of CSF into the blood
arachnoid mater
middle layer of the meninges named for the spider-web–like trabeculae that extend between it and the pia mater
arachnoid trabeculae
filaments between the arachnoid and pia mater within the subarachnoid space
areola
highly pigmented, circular area surrounding the raised nipple and containing areolar glands that secrete fluid important for lubrication during suckling
ascending colon
first region of the colon
ascending tract
central nervous system fibers carrying sensory information from the spinal cord or periphery to the brain
astrocyte
glial cell type of the CNS that provides support for neurons and maintains the blood-brain barrier
ATP synthase
protein pore complex that creates ATP
atrial natriuretic peptide (ANP)
peptide hormone produced by the walls of the atria in response to high blood pressure, blood volume, or blood sodium that reduces the reabsorption of sodium and water in the kidneys and promotes vasodilation
audition
sense of hearing
auricle
fleshy external structure of the ear
autocrine
chemical signal that elicits a response in the same cell that secreted it
autonomic nervous system (ANS)
functional division of the nervous system that is responsible for homeostatic reflexes that coordinate control of cardiac and smooth muscle, as well as glandular tissue
autosomal chromosome
in humans, the 22 pairs of chromosomes that are not the sex chromosomes (XX or XY)
autosomal dominant
pattern of dominant inheritance that corresponds to a gene on one of the 22 autosomal chromosomes
autosomal recessive
pattern of recessive inheritance that corresponds to a gene on one of the 22 autosomal chromosomes
axillary nerve
systemic nerve of the arm that arises from the brachial plexus
axon
single process of the neuron that carries an electrical signal (action potential) away from the cell body toward a target cell
axon hillock
tapering of the neuron cell body that gives rise to the axon
axon segment
single stretch of the axon insulated by myelin and bounded by nodes of Ranvier at either end (except for the first, which is after the initial segment, and the last, which is followed by the axon terminal)
axon terminal
end of the axon, where there are usually several branches extending toward the target cell
axoplasm
cytoplasm of an axon, which is different in composition than the cytoplasm of the neuronal cell body
bacterial flora
bacteria in the large intestine
Bartholin’s glands
(also, greater vestibular glands) glands that produce a thick mucus that maintains moisture in the vulva area; also referred to as the greater vestibular glands
basal forebrain
nuclei of the cerebrum related to modulation of sensory stimuli and attention through broad projections to the cerebral cortex, loss of which is related to Alzheimer’s disease
basal lamina
thin extracellular layer that lies underneath epithelial cells and separates them from other tissues
basal nuclei
nuclei of the cerebrum (with a few components in the upper brain stem and diencephalon) that are responsible for assessing cortical movement commands and comparing them with the general state of the individual through broad modulatory activity of dopamine neurons; largely related to motor functions, as evidenced through the symptoms of Parkinson’s and Huntington’s diseases
basal plate
developmental region of the spinal cord that gives rise to the lateral and anterior horns of gray matter
base
compound that accepts hydrogen ions (H+) in solution
basement membrane
in epithelial tissue, a thin layer of fibrous material that anchors the epithelial tissue to the underlying connective tissue; made up of the basal lamina and reticular lamina
basilar artery
blood vessel from the merged vertebral arteries that runs along the dorsal surface of the brain stem
basilar membrane
in the ear, the floor of the cochlear duct on which the organ of Corti sits
beta (β)-adrenergic receptor
one of the receptors to which epinephrine and norepinephrine bind, which comes in two subtypes: β1 and β2
beta (β)-hydroxybutyrate
primary ketone body produced in the body
beta (β)-oxidation
fatty acid oxidation
beta cell
pancreatic islet cell type that produces the hormone insulin
bile
alkaline solution produced by the liver and important for the emulsification of lipids
bile canaliculus
small duct between hepatocytes that collects bile
bile salts
salts that are released from the liver in response to lipid ingestion and surround the insoluble triglycerides to aid in their conversion to monoglycerides and free fatty acids
bilirubin
main bile pigment, which is responsible for the brown color of feces
biosynthesis reactions
reactions that create new molecules, also called anabolic reactions
bipolar cell
cell type in the retina that connects the photoreceptors to the RGCs
body
mid-portion of the stomach
body mass index (BMI)
relative amount of body weight compared to the overall height; a BMI ranging from 18–24.9 is considered normal weight, 25–29.9 is considered overweight, and greater than 30 is considered obese
body of uterus
middle section of the uterus
bolus
mass of chewed food
bond
electrical force linking atoms
Bowman’s capsule
cup-shaped sack lined by a simple squamous epithelium (parietal surface) and specialized cells called podocytes (visceral surface) that participate in the filtration process; receives the filtrate which then passes on to the PCTs
biogenic amine
class of neurotransmitters that are enzymatically derived from amino acids but no longer contain a carboxyl group
bipolar
shape of a neuron with two processes extending from the neuron cell body—the axon and one dendrite
blastocoel
fluid-filled cavity of the blastocyst
blastocyst
term for the conceptus at the developmental stage that consists of about 100 cells shaped into an inner cell mass that is fated to become the embryo and an outer trophoblast that is fated to become the associated fetal membranes and placenta
blastomere
daughter cell of a cleavage
blood-brain barrier (BBB)
physiological barrier between the circulatory system and the central nervous system that establishes a privileged blood supply, restricting the flow of substances into the CNS
blood–testis barrier
tight junctions between Sertoli cells that prevent bloodborne pathogens from gaining access to later stages of spermatogenesis and prevent the potential for an autoimmune reaction to haploid sperm
brachial plexus
nerve plexus associated with the lower cervical spinal nerves and first thoracic spinal nerve
brain
the large organ of the central nervous system composed of white and gray matter, contained within the cranium and continuous with the spinal cord
broad ligament
wide ligament that supports the uterus by attaching laterally to both sides of the uterus and pelvic wall
Broca’s area
region of the frontal lobe associated with the motor commands necessary for speech production and located only in the cerebral hemisphere responsible for language production, which is the left side in approximately 95 percent of the population
Brodmann’s areas
mapping of regions of the cerebral cortex based on microscopic anatomy that relates specific areas to functional differences, as described by Brodmann in the early 1900s
brush border
fuzzy appearance of the small intestinal mucosa created by microvilli
brush border
formed by microvilli on the surface of certain cuboidal cells; in the kidney it is found in the PCT; increases surface area for absorption in the kidney
buffer
solution containing a weak acid or a weak base that opposes wide fluctuations in the pH of body fluids
bulbourethral glands
(also, Cowper’s glands) glands that secrete a lubricating mucus that cleans and lubricates the urethra prior to and during ejaculation
calcitonin
peptide hormone produced and secreted by the parafollicular cells (C cells) of the thyroid gland that functions to decrease blood calcium levels
calorie
amount of heat it takes to raise 1 kg (1000 g) of water by 1 °C
calyces
cup-like structures receiving urine from the collecting ducts where it passes on to the renal pelvis and ureter
capsaicin
molecule that activates nociceptors by interacting with a temperature-sensitive ion channel and is the basis for “hot” sensations in spicy food
capacitation
process that occurs in the female reproductive tract in which sperm are prepared for fertilization; leads to increased motility and changes in their outer membrane that improve their ability to release enzymes capable of digesting an oocyte’s outer layers
carbohydrate
class of organic compounds built from sugars, molecules containing carbon, hydrogen, and oxygen in a 1-2-1 ratio
cardia
(also, cardiac region) part of the stomach surrounding the cardiac orifice (esophageal hiatus)
carotid canal
opening in the temporal bone through which the internal carotid artery enters the cranium
carrier
heterozygous individual who does not display symptoms of a recessive genetic disorder but can transmit the disorder to his or her offspring
catabolic hormones
hormones that stimulate the breakdown of larger molecules
catabolic reactions
reactions that break down larger molecules into their constituent parts
cation
atom with a positive charge
cauda equina
bundle of spinal nerve roots that descend from the lower spinal cord below the first lumbar vertebra and lie within the vertebral cavity; has the appearance of a horse's tail
caudate
nucleus deep in the cerebrum that is part of the basal nuclei; along with the putamen, it is part of the striatum
cecum
pouch forming the beginning of the large intestine
celiac ganglion
one of the collateral ganglia of the sympathetic system that projects to the digestive system
cell cycle
life cycle of a single cell, from its birth until its division into two new daughter cells
cell junction
point of cell-to-cell contact that connects one cell to another in a tissue
cellular respiration
production of ATP from glucose oxidation via glycolysis, the Krebs cycle, and oxidative phosphorylation
cementum
bone-like tissue covering the root of a tooth
central canal
hollow space within the spinal cord that is the remnant of the center of the neural tube
central nervous system (CNS)
anatomical division of the nervous system located within the cranial and vertebral cavities, namely the brain and spinal cord
central neuron
specifically referring to the cell body of a neuron in the autonomic system that is located in the central nervous system, specifically the lateral horn of the spinal cord or a brain stem nucleus
central sulcus
surface landmark of the cerebral cortex that marks the boundary between the frontal and parietal lobes
central vein
vein that receives blood from hepatic sinusoids
centromere
region of attachment for two sister chromatids
centrosome
cellular structure that organizes microtubules during cell division
cephalic phase
(also, reflex phase) initial phase of gastric secretion that occurs before food enters the stomach
cerebral aqueduct
connection of the ventricular system between the third and fourth ventricles located in the midbrain
cerebral cortex
outer gray matter covering the forebrain, marked by wrinkles and folds known as gyri and sulci
cerebrum
region of the adult brain that develops from the telencephalon and is responsible for higher neurological functions such as memory, emotion, and consciousness
cerebellum
region of the adult brain connected primarily to the pons that developed from the metencephalon (along with the pons) and is largely responsible for comparing information from the cerebrum with sensory feedback from the periphery through the spinal cord
cerebral hemisphere
one half of the bilaterally symmetrical cerebrum
cerebrospinal fluid (CSF)
circulatory medium within the CNS that is produced by ependymal cells in the choroid plexus filtering the blood
cervical plexus
nerve plexus associated with the upper cervical spinal nerves
cervix
elongate inferior end of the uterus where it connects to the vagina
chief cell
gastric gland cell that secretes pepsinogen
checkpoint
progress point in the cell cycle during which certain conditions must be met in order for the cell to proceed to a subsequence phase
chemical digestion
enzymatic breakdown of food
chemical synapse
connection between two neurons, or between a neuron and its target, where a neurotransmitter diffuses across a very short distance
chemoreceptor
sensory receptor cell that is sensitive to chemical stimuli, such as in taste, smell, or pain
cholecystokinin (CCK)
hormone that stimulates the release of pancreatic lipase and the contraction of the gallbladder to release bile salts
cholinergic
synapse at which acetylcholine is released and binds to the nicotinic or muscarinic receptor
cholinergic system
neurotransmitter system of acetylcholine, which includes its receptors and the enzyme acetylcholinesterase
choroid
highly vascular tissue in the wall of the eye that supplies the outer retina with blood
choroid plexus
specialized structure containing ependymal cells that line blood capillaries and filter blood to produce CSF in the four ventricles of the brain
chorion
membrane that develops from the syncytiotrophoblast, cytotrophoblast, and mesoderm; surrounds the embryo and forms the fetal portion of the placenta through the chorionic villi
chorionic membrane
precursor to the chorion; forms from extra-embryonic mesoderm cells
chorionic villi
projections of the chorionic membrane that burrow into the endometrium and develop into the placenta
chromaffin
neuroendocrine cells of the adrenal medulla
chromaffin cells
neuroendocrine cells of the adrenal medulla that release epinephrine and norepinephrine into the bloodstream as part of sympathetic system activity
chyme
soupy liquid created when food is mixed with digestive juices
chylomicrons
vesicles containing cholesterol and triglycerides that transport lipids out of the intestinal cells and into the lymphatic and circulatory systems
chymotrypsin
pancreatic enzyme that digests protein
chymotrypsinogen
proenzyme that is activated by trypsin into chymotrypsin
cleavage furrow
contractile ring that forms around a cell during cytokinesis that pinches the cell into two halves
cranial nerve
one of twelve nerves connected to the brain that are responsible for sensory or motor functions of the head and neck
cranial nerve ganglion
sensory ganglion of cranial nerves
cyclin
one of a group of proteins that function in the progression of the cell cycle
cyclin-dependent kinase (CDK)
one of a group of enzymes associated with cyclins that help them perform their functions
cytokinesis
final stage in cell division, where the cytoplasm divides to form two separate daughter cells
ciliary body
smooth muscle structure on the interior surface of the iris that controls the shape of the lens through the zonule fibers
ciliary ganglion
one of the terminal ganglia of the parasympathetic system, located in the posterior orbit, axons from which project to the iris
circle of Willis
unique anatomical arrangement of blood vessels around the base of the brain that maintains perfusion of blood into the brain even if one component of the structure is blocked or narrowed
circular fold
(also, plica circulare) deep fold in the mucosa and submucosa of the small intestine
citric acid cycle
also called the Krebs cycle or the tricarboxylic acid cycle; converts pyruvate into CO2 and high-energy FADH2, NADH, and ATP molecules
cleavage
form of mitotic cell division in which the cell divides but the total volume remains unchanged; this process serves to produce smaller and smaller cells
clitoris
(also, glans clitoris) nerve-rich area of the vulva that contributes to sexual sensation during intercourse
cochlea
auditory portion of the inner ear containing structures to transduce sound stimuli
cochlear duct
space within the auditory portion of the inner ear that contains the organ of Corti and is adjacent to the scala tympani and scala vestibuli on either side
codominance
pattern of inheritance that corresponds to the equal, distinct, and simultaneous expression of two different alleles
collateral ganglia
ganglia outside of the sympathetic chain that are targets of sympathetic preganglionic fibers, which are the celiac, inferior mesenteric, and superior mesenteric ganglia
colloid
liquid mixture in which the solute particles consist of clumps of molecules large enough to scatter light
colloid
viscous fluid in the central cavity of thyroid follicles, containing the glycoprotein thyroglobulin
colon
part of the large intestine between the cecum and the rectum
common bile duct
structure formed by the union of the common hepatic duct and the gallbladder’s cystic duct
common carotid artery
blood vessel that branches off the aorta (or the brachiocephalic artery on the right) and supplies blood to the head and neck
common hepatic duct
duct formed by the merger of the two hepatic ducts
conceptus
pre-implantation stage of a fertilized egg and its associated membranes
cone photoreceptor
one of the two types of retinal receptor cell that is specialized for color vision through the use of three photopigments distributed through three separate populations of cells
continuous conduction
slow propagation of an action potential along an unmyelinated axon owing to voltage-gated Na+ channels located along the entire length of the cell membrane
contralateral
word meaning “on the opposite side,” as in axons that cross the midline in a fiber tract
cornea
fibrous covering of the anterior region of the eye that is transparent so that light can pass through it
corona radiata
in an oocyte, a layer of granulosa cells that surrounds the oocyte and that must be penetrated by sperm before fertilization can occur
corpus albicans
nonfunctional structure remaining in the ovarian stroma following structural and functional regression of the corpus luteum
corpus callosum
large white matter structure that connects the right and left cerebral hemispheres
corpus cavernosum
either of two columns of erectile tissue in the penis that fill with blood during an erection
corpus luteum
transformed follicle after ovulation that secretes progesterone
corpus spongiosum
(plural = corpora cavernosa) column of erectile tissue in the penis that fills with blood during an erection and surrounds the penile urethra on the ventral portion of the penis
cortical nephrons
nephrons with loops of Henle that do not extend into the renal medulla
cortical reaction
following fertilization, the release of cortical granules from the oocyte’s plasma membrane into the zona pellucida creating a fertilization membrane that prevents any further attachment or penetration of sperm; part of the slow block to polyspermy
cortisol
glucocorticoid important in gluconeogenesis, the catabolism of glycogen, and downregulation of the immune system
countercurrent multiplier system
involves the descending and ascending loops of Henle directing forming urine in opposing directions to create a concentration gradient when combined with variable permeability and sodium pumping
covalent bond
chemical bond in which two atoms share electrons, thereby completing their valence shells
craniosacral system
alternate name for the parasympathetic division of the autonomic nervous system that is based on the anatomical location of central neurons in brain-stem nuclei and the lateral horn of the sacral spinal cord; also referred to as craniosacral outflow
crown
portion of tooth visible superior to the gum line
cupula
specialized structure within the base of a semicircular canal that bends the stereocilia of hair cells when the head rotates by way of the relative movement of the enclosed fluid
cuspid
(also, canine) pointed tooth used for tearing and shredding food
cystic duct
duct through which bile drains and enters the gallbladder
cyclic adenosine monophosphate (cAMP)
second messenger that, in response to adenylyl cyclase activation, triggers a phosphorylation cascade
chylomicron
large lipid-transport compound made up of triglycerides, phospholipids, cholesterol, and proteins
defecation
elimination of undigested substances from the body in the form of feces
delta cell
minor cell type in the pancreas that secretes the hormone somatostatin
denaturation
change in the structure of a molecule through physical or chemical means
dendrite
one of many branchlike processes that extends from the neuron cell body and functions as a contact for incoming signals (synapses) from other neurons or sensory cells
deoxyribonuclease
pancreatic enzyme that digests DNA
deoxyribonucleic acid (DNA)
deoxyribose-containing nucleotide that stores genetic information
depolarization
change in a cell membrane potential from rest toward zero
deglutition
three-stage process of swallowing
dehydration
state of containing insufficient water in blood and other tissues
dens
tooth
dentin
bone-like tissue immediately deep to the enamel of the crown or cementum of the root of a tooth
dentition
set of teeth
descending colon
part of the colon between the transverse colon and the sigmoid colon
descending tract
central nervous system fibers carrying motor commands from the brain to the spinal cord or periphery
deciduous tooth
one of 20 “baby teeth”
detrusor muscle
smooth muscle in the bladder wall; fibers run in all directions to reduce the size of the organ when emptying it of urine
diabetes mellitus
condition caused by destruction or dysfunction of the beta cells of the pancreas or cellular resistance to insulin that results in abnormally high blood glucose levels
diacylglycerol (DAG)
molecule that, like cAMP, activates protein kinases, thereby initiating a phosphorylation cascade
dihydroxyvitamin D
active form of vitamin D required by the intestinal epithelial cells for the absorption of calcium
dipeptidase
brush border enzyme that acts on proteins
diploid
condition marked by the presence of a double complement of genetic material (two sets of chromosomes, one set inherited from each of two parents)
direct pathway
connections within the basal nuclei from the striatum to the globus pallidus internal segment and substantia nigra pars reticulata that disinhibit the thalamus to increase cortical control of movement
disaccharide
pair of carbohydrate monomers bonded by dehydration synthesis via a glycosidic bond
disinhibition
disynaptic connection in which the first synapse inhibits the second cell, which then stops inhibiting the final target
distal convoluted tubules
portions of the nephron distal to the loop of Henle that receive hyposmotic filtrate from the loop of Henle and empty into collecting ducts
disulfide bond
covalent bond formed within a polypeptide between sulfide groups of sulfur-containing amino acids, for example, cysteine
diuresis
excess production of urine
dominant
describes a trait that is expressed both in homozygous and heterozygous form
dominant lethal
inheritance pattern in which individuals with one or two copies of a lethal allele do not survive in utero or have a shortened life span
dorsal (posterior) nerve root
axons entering the posterior horn of the spinal cord
dorsal nucleus of the vagus nerve
location of parasympathetic neurons that project through the vagus nerve to terminal ganglia in the thoracic and abdominal cavities
dorsal (posterior) root ganglion
sensory ganglion attached to the posterior nerve root of a spinal nerve
downregulation
decrease in the number of hormone receptors, typically in response to chronically excessive levels of a hormone
ductus arteriosus
shunt in the pulmonary trunk that diverts oxygenated blood back to the aorta
ductus deferens
(also, vas deferens) duct that transports sperm from the epididymis through the spermatic cord and into the ejaculatory duct; also referred as the vas deferens
ductus venosus
shunt that causes oxygenated blood to bypass the fetal liver on its way to the inferior vena cava
duodenal gland
(also, Brunner’s gland) mucous-secreting gland in the duodenal submucosa
duodenum
first part of the small intestine, which starts at the pyloric sphincter and ends at the jejunum
dura mater
tough, fibrous, outer layer of the meninges that is attached to the inner surface of the cranium and vertebral column and surrounds the entire CNS
dural sinus
any of the venous structures surrounding the brain, enclosed within the dura mater, which drain blood from the CNS to the common venous return of the jugular veins
ectoderm
primary germ layer that develops into the central and peripheral nervous systems, sensory organs, epidermis, hair, and nails
ectopic pregnancy
implantation of an embryo outside of the uterus
Eddinger–Westphal nucleus
location of parasympathetic neurons that project to the ciliary ganglion
effector protein
enzyme that catalyzes the generation of a new molecule, which acts as the intracellular mediator of the signal that binds to the receptor
efferent arteriole
arteriole carrying blood from the glomerulus to the capillary beds around the convoluted tubules and loop of Henle; portion of the portal system
elastase
pancreatic enzyme that digests protein
electrical synapse
connection between two neurons, or any two electrically active cells, where ions flow directly through channels spanning their adjacent cell membranes
electron transport chain (ETC)
ATP production pathway in which electrons are passed through a series of oxidation-reduction reactions that forms water and produces a proton gradient
embryo
developing human during weeks 3–8
embryonic folding
process by which an embryo develops from a flat disc of cells to a three-dimensional shape resembling a cylinder
endocrine gland
groups of cells that release chemical signals into the intercellular fluid to be picked up and transported to their target organs by blood
endoderm
primary germ layer that goes on to form the gastrointestinal tract, liver, pancreas, and lungs
endoneurium
innermost layer of connective tissue that surrounds individual axons within a nerve
endothelium
tissue that lines vessels of the lymphatic and cardiovascular system, made up of a simple squamous epithelium
enamel
covering of the dentin of the crown of a tooth
encapsulated ending
configuration of a sensory receptor neuron with dendrites surrounded by specialized structures to aid in transduction of a particular type of sensation, such as the lamellated corpuscles in the deep dermis and subcutaneous tissue
endocrine gland
tissue or organ that secretes hormones into the blood and lymph without ducts such that they may be transported to organs distant from the site of secretion
endocrine system
cells, tissues, and organs that secrete hormones as a primary or secondary function and play an integral role in normal bodily processes
endogenous
describes substance made in the human body
endometrium
inner lining of the uterus, part of which builds up during the secretory phase of the menstrual cycle and then sheds with menses
energy-consuming phase
first phase of glycolysis, in which two molecules of ATP are necessary to start the reaction
energy-yielding phase
second phase of glycolysis, during which energy is produced
enteric nervous system (ENS)
neural tissue associated with the digestive system that is responsible for nervous control through autonomic connections
enteroendocrine cell
gastric gland cell that releases hormones
enteric nervous system
peripheral structures, namely ganglia and nerves, that are incorporated into the digestive system organs
enteric plexus
neuronal plexus in the wall of the intestines, which is part of the enteric nervous system
enterohepatic circulation
recycling mechanism that conserves bile salts
enterokinase
enzyme located in the wall of the small intestine that activates trypsin
enteropeptidase
intestinal brush-border enzyme that activates trypsinogen to trypsin
electrochemical exclusion
principle of selectively allowing ions through a channel on the basis of their charge
ejaculatory duct
duct that connects the ampulla of the ductus deferens with the duct of the seminal vesicle at the prostatic urethra
ependymal cell
glial cell type in the CNS responsible for producing cerebrospinal fluid
epiblast
upper layer of cells of the embryonic disc that forms from the inner cell mass; gives rise to all three germ layers
epididymis
(plural = epididymides) coiled tubular structure in which sperm start to mature and are stored until ejaculation
epiploic appendage
small sac of fat-filled visceral peritoneum attached to teniae coli
epinephrine
signaling molecule released from the adrenal medulla into the bloodstream as part of the sympathetic response
epinephrine
primary and most potent catecholamine hormone secreted by the adrenal medulla in response to short-term stress; also called adrenaline
epineurium
outermost layer of connective tissue that surrounds an entire nerve
epithalamus
region of the diecephalon containing the pineal gland
equilibrium
sense of balance that includes sensations of position and movement of the head
erythropoietin (EPO)
protein hormone secreted in response to low oxygen levels that triggers the bone marrow to produce red blood cells]
esophageal plexus
neuronal plexus in the wall of the esophagus that is part of the enteric nervous system
esophagus
muscular tube that runs from the pharynx to the stomach
estrogens
class of predominantly female sex hormones important for the development and growth of the female reproductive tract, secondary sex characteristics, the female reproductive cycle, and the maintenance of pregnancy
excitable membrane
cell membrane that regulates the movement of ions so that an electrical signal can be generated
excitatory postsynaptic potential (EPSP)
graded potential in the postsynaptic membrane that is the result of depolarization and makes an action potential more likely to occur
exocrine gland
group of epithelial cells that secrete substances through ducts that open to the skin or to internal body surfaces that lead to the exterior of the body
exocrine system
cells, tissues, and organs that secrete substances directly to target tissues via glandular ducts
exogenous
describes substance made outside of the human body
external anal sphincter
voluntary skeletal muscle sphincter in the anal canal
external ear
structures on the lateral surface of the head, including the auricle and the ear canal back to the tympanic membrane
external urinary sphincter
skeletal muscle; must be relaxed consciously to void urine
exteroceptor
sensory receptor that is positioned to interpret stimuli from the external environment, such as photoreceptors in the eye or somatosensory receptors in the skin
extracellular fluid (ECF)
fluid exterior to cells; includes the interstitial fluid, blood plasma, and fluids found in other reservoirs in the body
extraocular muscle
one of six muscles originating out of the bones of the orbit and inserting into the surface of the eye which are responsible for moving the eye
extraocular muscles
six skeletal muscles that control eye movement within the orbit
FADH2
high-energy molecule needed for glycolysis
facial nerve
seventh cranial nerve; responsible for contraction of the facial muscles and for part of the sense of taste, as well as causing saliva production
fascicle
small bundles of nerve or muscle fibers enclosed by connective tissue
fatty acid oxidation
breakdown of fatty acids into smaller chain fatty acids and acetyl CoA
fauces
opening between the oral cavity and the oropharynx
feces
semisolid waste product of digestion
femoral nerve
systemic nerve of the anterior leg that arises from the lumbar plexus
fenestrations
small windows through a cell, allowing rapid filtration based on size; formed in such a way as to allow substances to cross through a cell without mixing with cell contents
fertilization
unification of genetic material from male and female haploid gametes
fertilization membrane
impenetrable barrier that coats a nascent zygote; part of the slow block to polyspermy
fetus
developing human during the time from the end of the embryonic period (week 9) to birth
fibrous tunic
outer layer of the eye primarily composed of connective tissue known as the sclera and cornea
fibular nerve
systemic nerve of the posterior leg that begins as part of the sciatic nerve
fight-or-flight response
set of responses induced by sympathetic activity that lead to either fleeing a threat or standing up to it, which in the modern world is often associated with anxious feelings
filtration slits
formed by pedicels of podocytes; substances filter between the pedicels based on size
fimbriae
fingerlike projections on the distal uterine tubes
first messenger
hormone that binds to a cell membrane hormone receptor and triggers activation of a second messenger system
flatus
gas in the intestine
flavin adenine dinucleotide (FAD)
coenzyme used to produce FADH2
fluid compartment
fluid inside all cells of the body constitutes a compartment system that is largely segregated from other systems
follicle
ovarian structure of one oocyte and surrounding granulosa (and later theca) cells
follicle-stimulating hormone (FSH)
anterior pituitary hormone that stimulates the production and maturation of sex cells
folliculogenesis
development of ovarian follicles from primordial to tertiary under the stimulation of gonadotropins
foramen magnum
large opening in the occipital bone of the skull through which the spinal cord emerges and the vertebral arteries enter the cranium
foramen ovale
shunt that directly connects the right and left atria and helps divert oxygenated blood from the fetal pulmonary circuit
forming urine
filtrate undergoing modifications through secretion and reabsorption before true urine is produced
fourth ventricle
the portion of the ventricular system that is in the region of the brain stem and opens into the subarachnoid space through the median and lateral apertures
fovea
exact center of the retina at which visual stimuli are focused for maximal acuity, where the retina is thinnest, at which there is nothing but photoreceptors
free nerve ending
configuration of a sensory receptor neuron with dendrites in the connective tissue of the organ, such as in the dermis of the skin, that are most often sensitive to chemical, thermal, and mechanical stimuli
frontal eye field
region of the frontal lobe associated with motor commands to orient the eyes toward an object of visual attention
frontal lobe
region of the cerebral cortex directly beneath the frontal bone of the cranium
functional group
group of atoms linked by strong covalent bonds that tends to behave as a distinct unit in chemical reactions with other atoms
fundus
dome-shaped region of the stomach above and to the left of the cardia
fundus
(of the uterus) domed portion of the uterus that is superior to the uterine tubes
G0 phase
phase of the cell cycle, usually entered from the G1 phase; characterized by long or permanent periods where the cell does not move forward into the DNA synthesis phase
G1 phase
first phase of the cell cycle, after a new cell is born
G2 phase
third phase of the cell cycle, after the DNA synthesis phase
G cell
gastrin-secreting enteroendocrine cell
G protein–coupled receptor
membrane protein complex that consists of a receptor protein that binds to a signaling molecule—a G protein—that is activated by that binding and in turn activates an effector protein (enzyme) that creates a second-messenger molecule in the cytoplasm of the target cell
gallbladder
accessory digestive organ that stores and concentrates bile
gamete
haploid reproductive cell that contributes genetic material to form an offspring
gap junction
allows cytoplasmic communications to occur between cells
ganglion
localized collection of neuron cell bodies in the peripheral nervous system
ganglionic neuron
specifically refers to the cell body of a neuron in the autonomic system that is located in a ganglion
gastric emptying
process by which mixing waves gradually cause the release of chyme into the duodenum
gastric gland
gland in the stomach mucosal epithelium that produces gastric juice
gastric phase
phase of gastric secretion that begins when food enters the stomach
gastric plexuses
neuronal networks in the wall of the stomach that are part of the enteric nervous system
gastric pit
narrow channel formed by the epithelial lining of the stomach mucosa
gastrin
peptide hormone that stimulates secretion of hydrochloric acid and gut motility
gastrocolic reflex
propulsive movement in the colon activated by the presence of food in the stomach
gastroileal reflex
long reflex that increases the strength of segmentation in the ileum
gastrulation
process of cell migration and differentiation into three primary germ layers following cleavage and implantation
gated
property of a channel that determines how it opens under specific conditions, such as voltage change or physical deformation
general adaptation syndrome (GAS)
the human body’s three-stage response pattern to short- and long-term stress
general sense
any sensory system that is distributed throughout the body and incorporated into organs of multiple other systems, such as the walls of the digestive organs or the skin
generator potential
graded potential from dendrites of a unipolar cell which generates the action potential in the initial segment of that cell’s axon
genotype
complete genetic makeup of an individual
gestation
in human development, the period required for embryonic and fetal development in utero; pregnancy
G protein
guanosine triphosphate (GTP) hydrolase that physically moves from the receptor protein to the effector protein to activate the latter
G protein
protein associated with a cell membrane hormone receptor that initiates the next step in a second messenger system upon activation by hormone–receptor binding
gigantism
disorder in children caused when abnormally high levels of GH prompt excessive growth
gingiva
gum
glans penis
bulbous end of the penis that contains a large number of nerve endings
glial cell
one of the various types of neural tissue cells responsible for maintenance of the tissue, and largely responsible for supporting neurons
globus pallidus
nuclei deep in the cerebrum that are part of the basal nuclei and can be divided into the internal and external segments
glomerular filtration rate (GFR)
rate of renal filtration
glomerulus
tuft of capillaries surrounded by Bowman’s capsule; filters the blood based on size
glossopharyngeal nerve
ninth cranial nerve; responsible for contraction of muscles in the tongue and throat and for part of the sense of taste, as well as causing saliva production
glucagon
pancreatic hormone that stimulates the catabolism of glycogen to glucose, thereby increasing blood glucose levels
glucocorticoids
hormones produced by the zona fasciculata of the adrenal cortex that influence glucose metabolism
glucokinase
cellular enzyme, found in the liver, which converts glucose into glucose-6-phosphate upon uptake into the cell
gluconeogenesis
process of glucose synthesis from pyruvate or other molecules
glucose-6-phosphate
phosphorylated glucose produced in the first step of glycolysis
glycogen
form that glucose assumes when it is stored
glycolysis
series of metabolic reactions that breaks down glucose into pyruvate and produces ATP
glycosuria
presence of glucose in the urine; caused by high blood glucose levels that exceed the ability of the kidneys to reabsorb the glucose; usually the result of untreated or poorly controlled diabetes mellitus
goblet cell
unicellular gland found in columnar epithelium that secretes mucous
goiter
enlargement of the thyroid gland either as a result of iodine deficiency or hyperthyroidism
gonadotropin-releasing hormone (GnRH)
hormone released by the hypothalamus that regulates the production of follicle-stimulating hormone and luteinizing hormone from the pituitary gland
gonadotropins
hormones that regulate the function of the gonads
gonads
reproductive organs (testes in men and ovaries in women) that produce gametes and reproductive hormones
graded potential
change in the membrane potential that varies in size, depending on the size of the stimulus that elicits it
granulosa cells
supportive cells in the ovarian follicle that produce estrogen
gray rami communicantes
(singular = ramus communicans) unmyelinated structures that provide a short connection from a sympathetic chain ganglion to the spinal nerve that contains the postganglionic sympathetic fiber
greater splanchnic nerve
nerve that contains fibers of the central sympathetic neurons that do not synapse in the chain ganglia but project onto the celiac ganglion
growth hormone (GH)
anterior pituitary hormone that promotes tissue building and influences nutrient metabolism (also called somatotropin)
gustation
sense of taste
gustatory receptor cells
sensory cells in the taste bud that transduce the chemical stimuli of gustation
gray matter
regions of the nervous system containing cell bodies of neurons with few or no myelinated axons; actually may be more pink or tan in color, but called gray in contrast to white matter
gyrus
ridge formed by convolutions on the surface of the cerebrum or cerebellum
hair cells
mechanoreceptor cells found in the inner ear that transduce stimuli for the senses of hearing and balance
haustrum
small pouch in the colon created by tonic contractions of teniae coli
haustral contraction
slow segmentation in the large intestine
hepatic artery
artery that supplies oxygenated blood to the liver
hepatic lobule
hexagonal-shaped structure composed of hepatocytes that radiate outward from a central vein
hepatic portal vein
vein that supplies deoxygenated nutrient-rich blood to the liver
hepatic sinusoid
blood capillaries between rows of hepatocytes that receive blood from the hepatic portal vein and the branches of the hepatic artery
hepatic vein
vein that drains into the inferior vena cava
hepatocytes
major functional cells of the liver
hepatopancreatic ampulla
(also, ampulla of Vater) bulb-like point in the wall of the duodenum where the bile duct and main pancreatic duct unite
hepatopancreatic sphincter
(also, sphincter of Oddi) sphincter regulating the flow of bile and pancreatic juice into the duodenum
heterozygous
having two different alleles for a given gene
hexokinase
cellular enzyme, found in most tissues, that converts glucose into glucose-6-phosphate upon uptake into the cell
hippocampus
gray matter deep in the temporal lobe that is very important for long-term memory formation
holocrine secretion
release of a substance caused by the rupture of a gland cell, which becomes part of the secretion
homologous
describes two copies of the same chromosome (not identical), one inherited from each parent
homozygous
having two identical alleles for a given gene
hormone
secretion of an endocrine organ that travels via the bloodstream or lymphatics to induce a response in target cells or tissues in another part of the body
hormone receptor
protein within a cell or on the cell membrane that binds a hormone, initiating the target cell response
human chorionic gonadotropin (hCG)
hormone that directs the corpus luteum to survive, enlarge, and continue producing progesterone and estrogen to suppress menses and secure an environment suitable for the developing embryo
hydrochloric acid (HCl)
digestive acid secreted by parietal cells in the stomach
hydrogen bond
dipole-dipole bond in which a hydrogen atom covalently bonded to an electronegative atom is weakly attracted to a second electronegative atom
hypoblast
lower layer of cells of the embryonic disc that extend into the blastocoel to form the yolk sac
hydrostatic pressure
pressure exerted by a fluid against a wall, caused by its own weight or pumping force
hydroxymethylglutaryl CoA (HMG CoA)
molecule created in the first step of the creation of ketone bodies from acetyl CoA
hymen
membrane that covers part of the opening of the vagina
hypercalcemia
abnormally increased blood levels of calcium
hypercapnia
abnormally elevated blood levels of CO2
hyperchloremia
higher-than-normal blood chloride levels
hyperglycemia
abnormally high blood glucose levels
hyperkalemia
higher-than-normal blood potassium levels
hypernatremia
abnormal increase in blood sodium levels
hyperphosphatemia
abnormally increased blood phosphate levels
hyperthyroidism
clinically abnormal, elevated level of thyroid hormone in the blood; characterized by an increased metabolic rate, excess body heat, sweating, diarrhea, weight loss, and increased heart rate
hyperparathyroidism
disorder caused by overproduction of PTH that results in abnormally elevated blood calcium
hypocalcemia
abnormally low blood levels of calcium
hypocapnia
abnormally low blood levels of CO2
hypochloremia
lower-than-normal blood chloride levels
hypoglossal nerve
twelfth cranial nerve; responsible for contraction of muscles of the tongue
hypokalemia
abnormally decreased blood levels of potassium
hyponatremia
lower-than-normal levels of sodium in the blood
hypoparathyroidism
disorder caused by underproduction of PTH that results in abnormally low blood calcium
hypophosphatemia
abnormally low blood phosphate levels
hypophyseal portal system
network of blood vessels that enables hypothalamic hormones to travel into the anterior lobe of the pituitary without entering the systemic circulation
hypothalamus
region of the diencephalon inferior to the thalamus that functions in neural and endocrine signaling
hypothalamus
major region of the diencephalon that is responsible for coordinating autonomic and endocrine control of homeostasis
hypothyroidism
clinically abnormal, low level of thyroid hormone in the blood; characterized by low metabolic rate, weight gain, cold extremities, constipation, and reduced mental activity
ileocecal sphincter
sphincter located where the small intestine joins with the large intestine
ileum
end of the small intestine between the jejunum and the large intestine
implantation
process by which a blastocyst embeds itself in the uterine endometrium
inactivation gate
part of a voltage-gated Na+ channel that closes when the membrane potential reaches +30 mV
inactive proenzymes
forms in which proteases are stored and released to prevent the inappropriate digestion of the native proteins of the stomach, pancreas, and small intestine
incomplete dominance
pattern of inheritance in which a heterozygous genotype expresses a phenotype intermediate between dominant and recessive phenotypes
incontinence
loss of ability to control micturition
incus
(also, anvil) ossicle of the middle ear that connects the malleus to the stapes
indirect pathway
connections within the basal nuclei from the striatum through the globus pallidus external segment and subthalamic nucleus to the globus pallidus internal segment/substantia nigra pars compacta that result in inhibition of the thalamus to decrease cortical control of movement
inferior colliculus
half of the midbrain tectum that is part of the brain stem auditory pathway
inferior mesenteric ganglion
one of the collateral ganglia of the sympathetic system that projects to the digestive system
inferior oblique
extraocular muscle responsible for lateral rotation of the eye
inferior rectus
extraocular muscle responsible for looking down
inferior olive
nucleus in the medulla that is involved in processing information related to motor control
infundibulum
stalk containing vasculature and neural tissue that connects the pituitary gland to the hypothalamus (also called the pituitary stalk)
infundibulum
(of the uterine tube) wide, distal portion of the uterine tube terminating in fimbriae
ingestion
taking food into the GI tract through the mouth
inguinal canal
opening in abdominal wall that connects the testes to the abdominal cavity
inhibin
hormone secreted by the male and female gonads that inhibits FSH production by the anterior pituitary
inhibitory postsynaptic potential (IPSP)
graded potential in the postsynaptic membrane that is the result of hyperpolarization and makes an action potential less likely to occur
interphase
entire life cycle of a cell, excluding mitosis
initial segment
first part of the axon as it emerges from the axon hillock, where the electrical signals known as action potentials are generated
inner cell mass
cluster of cells within the blastocyst that is fated to become the embryo
inner ear
structure within the temporal bone that contains the sensory apparati of hearing and balance
inner segment
in the eye, the section of a photoreceptor that contains the nucleus and other major organelles for normal cellular functions
inner synaptic layer
layer in the retina where bipolar cells connect to RGCs
inorganic compound
substance that does not contain both carbon and hydrogen
inositol triphosphate (IP3)
molecule that initiates the release of calcium ions from intracellular stores
incisor
midline, chisel-shaped tooth used for cutting into food
insulin
pancreatic hormone that enhances the cellular uptake and utilization of glucose, thereby decreasing blood glucose levels
insulin
hormone secreted by the pancreas that stimulates the uptake of glucose into the cells
insulin-like growth factors (IGF)
protein that enhances cellular proliferation, inhibits apoptosis, and stimulates the cellular uptake of amino acids for protein synthesis
integration
nervous system function that combines sensory perceptions and higher cognitive functions (memories, learning, emotion, etc.) to produce a response
intercalated cell
specialized cell of the collecting ducts that secrete or absorb acid or bicarbonate; important in acid–base balance
intercostal nerve
systemic nerve in the thoracic cavity that is found between two ribs
internal anal sphincter
involuntary smooth muscle sphincter in the anal canal
internal carotid artery
branch from the common carotid artery that enters the cranium and supplies blood to the brain
internal urinary sphincter
smooth muscle at the juncture of the bladder and urethra; relaxes as the bladder fills to allow urine into the urethra
interoceptor
sensory receptor that is positioned to interpret stimuli from internal organs, such as stretch receptors in the wall of blood vessels
interventricular foramina
openings between the lateral ventricles and third ventricle allowing for the passage of CSF
intestinal gland
(also, crypt of Lieberkühn) gland in the small intestinal mucosa that secretes intestinal juice
intestinal juice
mixture of water and mucus that helps absorb nutrients from chyme
interstitial fluid (IF)
fluid in the small spaces between cells not contained within blood vessels
intracellular fluid (ICF)
fluid in the cytosol of cells
intramural ganglia
terminal ganglia of the parasympathetic system that are found within the walls of the target effector
intrinsic factor
glycoprotein required for vitamin B12 absorption in the small intestine
intestinal phase
phase of gastric secretion that begins when chyme enters the intestine
inulin
plant polysaccharide injected to determine GFR; is neither secreted nor absorbed by the kidney, so its appearance in the urine is directly proportional to its filtration rate
ion
atom with an overall positive or negative charge
ionic bond
attraction between an anion and a cation
ionotropic receptor
neurotransmitter receptor that acts as an ion channel gate, and opens by the binding of the neurotransmitter
ipsilateral
word meaning on the same side, as in axons that do not cross the midline in a fiber tract
iris
colored portion of the anterior eye that surrounds the pupil
isthmus
narrow, medial portion of the uterine tube that joins the uterus
jejunum
middle part of the small intestine between the duodenum and the ileum
jugular veins
blood vessels that return “used” blood from the head and neck
juxtamedullary nephrons
nephrons adjacent to the border of the cortex and medulla with loops of Henle that extend into the renal medulla
juxtaglomerular apparatus (JGA)
located at the juncture of the DCT and the afferent and efferent arterioles of the glomerulus; plays a role in the regulation of renal blood flow and GFR
juxtaglomerular cell
modified smooth muscle cells of the afferent arteriole; secretes renin in response to a drop in blood pressure
karyotype
systematic arrangement of images of chromosomes into homologous pairs
ketone bodies
alternative source of energy when glucose is limited, created when too much acetyl CoA is created during fatty acid oxidation
kinesthesia
general sensory perception of movement of the body
kinesthesia
sense of body movement based on sensations in skeletal muscles, tendons, joints, and the skin
kinetochore
region of a centromere where microtubules attach to a pair of sister chromatids
Krebs cycle
also called the citric acid cycle or the tricarboxylic acid cycle, converts pyruvate into CO2 and high-energy FADH2, NADH, and ATP molecules
labium
lip
labial frenulum
midline mucous membrane fold that attaches the inner surface of the lips to the gums
labia majora
hair-covered folds of skin located behind the mons pubis
labia minora
thin, pigmented, hairless flaps of skin located medial and deep to the labia majora
lacrimal duct
duct in the medial corner of the orbit that drains tears into the nasal cavity
lacrimal gland
gland lateral to the orbit that produces tears to wash across the surface of the eye
lactase
brush border enzyme that breaks down lactose into glucose and galactose
lacteal
lymphatic capillary in the villi
lactiferous ducts
ducts that connect the mammary glands to the nipple and allow for the transport of milk
lactiferous sinus
area of milk collection between alveoli and lactiferous duct
lanugo
silk-like hairs that coat the fetus; shed later in fetal development
large intestine
terminal portion of the alimentary canal
lateral horn
region of the spinal cord gray matter in the thoracic, upper lumbar, and sacral regions that is the central component of the sympathetic division of the autonomic nervous system
lateral apertures
pair of openings from the fourth ventricle to the subarachnoid space on either side and between the medulla and cerebellum
lateral column
white matter of the spinal cord between the posterior horn on one side and the axons from the anterior horn on the same side; composed of many different groups of axons, of both ascending and descending tracts, carrying motor commands to and from the brain
lateral rectus
extraocular muscle responsible for abduction of the eye
lateral sulcus
surface landmark of the cerebral cortex that marks the boundary between the temporal lobe and the frontal and parietal lobes
lateral ventricles
portions of the ventricular system that are in the region of the cerebrum
laryngopharynx
part of the pharynx that functions in respiration and digestion
leakage channel
ion channel that opens randomly and is not gated to a specific event, also known as a non-gated channel
leaky tight junctions
tight junctions in which the sealing strands of proteins between the membranes of adjacent cells are fewer in number and incomplete; allows limited intercellular movement of solvent and solutes
left colic flexure
(also, splenic flexure) point where the transverse colon curves below the inferior end of the spleen
lens
component of the eye that focuses light on the retina
leptin
protein hormone secreted by adipose tissues in response to food consumption that promotes satiety
lesser splanchnic nerve
nerve that contains fibers of the central sympathetic neurons that do not synapse in the chain ganglia but project onto the inferior mesenteric ganglion
leukocyte esterase
enzyme produced by leukocytes that can be detected in the urine and that serves as an indirect indicator of urinary tract infection
levator palpebrae superioris
muscle that causes elevation of the upper eyelid, controlled by fibers in the oculomotor nerve
Leydig cells
cells between the seminiferous tubules of the testes that produce testosterone; a type of interstitial cell
ligand-gated cation channel
ion channel, such as the nicotinic receptor, that is specific to positively charged ions and opens when a molecule such as a neurotransmitter binds to it
ligand-gated channels
another name for an ionotropic receptor for which a neurotransmitter is the ligand
limbic cortex
collection of structures of the cerebral cortex that are involved in emotion, memory, and behavior and are part of the larger limbic system
limbic system
structures at the edge (limit) of the boundary between the forebrain and hindbrain that are most associated with emotional behavior and memory formation
lingual frenulum
mucous membrane fold that attaches the bottom of the tongue to the floor of the mouth
lingual lipase
digestive enzyme from glands in the tongue that acts on triglycerides
lipid
class of nonpolar organic compounds built from hydrocarbons and distinguished by the fact that they are not soluble in water
lipogenesis
synthesis of lipids that occurs in the liver or adipose tissues
lipolysis
breakdown of triglycerides into glycerol and fatty acids
lipoprotein lipase
enzyme that breaks down triglycerides in chylomicrons into fatty acids and monoglycerides
liver
largest gland in the body whose main digestive function is the production of bile
longitudinal fissure
large separation along the midline between the two cerebral hemispheres
loop of Henle
descending and ascending portions between the proximal and distal convoluted tubules; those of cortical nephrons do not extend into the medulla, whereas those of juxtamedullary nephrons do extend into the medulla
lower esophageal sphincter
smooth muscle sphincter that regulates food movement from the esophagus to the stomach
lower motor neuron
second neuron in the motor command pathway that is directly connected to the skeletal muscle
lumbar plexus
nerve plexus associated with the lumbar spinal nerves
lumbar puncture
procedure used to withdraw CSF from the lower lumbar region of the vertebral column that avoids the risk of damaging CNS tissue because the spinal cord ends at the upper lumbar vertebrae
luteinizing hormone (LH)
anterior pituitary hormone that triggers ovulation and the production of ovarian hormones in females, and the production of testosterone in males
macromolecule
large molecule formed by covalent bonding
macula
enlargement at the base of a semicircular canal at which transduction of equilibrium stimuli takes place within the ampulla
macula densa
cells found in the part of the DCT forming the JGA; sense Na+ concentration in the forming urine
main pancreatic duct
(also, duct of Wirsung) duct through which pancreatic juice drains from the pancreas
major duodenal papilla
point at which the hepatopancreatic ampulla opens into the duodenum
malleus
(also, hammer) ossicle that is directly attached to the tympanic membrane
maltase
brush border enzyme that breaks down maltose and maltotriose into two and three molecules of glucose, respectively
mammary glands
glands inside the breast that secrete milk
mass movement
long, slow, peristaltic wave in the large intestine
mastication
chewing
mechanical digestion
chewing, mixing, and segmentation that prepares food for chemical digestion
mechanoreceptor
receptor cell that transduces mechanical stimuli into an electrochemical signal
mechanically gated channel
ion channel that opens when a physical event directly affects the structure of the protein
meconium
fetal wastes consisting of ingested amniotic fluid, cellular debris, mucus, and bile
medial rectus
extraocular muscle responsible for adduction of the eye
median aperture
singular opening from the fourth ventricle into the subarachnoid space at the midline between the medulla and cerebellum
median nerve
systemic nerve of the arm, located between the ulnar and radial nerves
medulla
inner region of kidney containing the renal pyramids
melatonin
amino acid–derived hormone that is secreted in response to low light and causes drowsiness
membrane potential
distribution of charge across the cell membrane, based on the charges of ions
menarche
first menstruation in a pubertal female
meninges
protective outer coverings of the CNS composed of connective tissue
mesangial
contractile cells found in the glomerulus; can contract or relax to regulate filtration rate
mesenteric plexus
nervous tissue within the wall of the digestive tract that contains neurons that are the targets of autonomic preganglionic fibers and that project to the smooth muscle and glandular tissues in the digestive organ
menses
shedding of the inner portion of the endometrium out though the vagina; also referred to as menstruation
menses phase
phase of the menstrual cycle in which the endometrial lining is shed
menstrual cycle
approximately 28-day cycle of changes in the uterus consisting of a menses phase, a proliferative phase, and a secretory phase
merocrine secretion
release of a substance from a gland via exocytosis
mesoappendix
mesentery of the appendix
mesoderm
primary germ layer that becomes the skeleton, muscles, connective tissue, heart, blood vessels, and kidneys
mesothelium
simple squamous epithelial tissue which covers the major body cavities and is the epithelial portion of serous membranes
metabolic acidosis
condition wherein a deficiency of bicarbonate causes the blood to be overly acidic
metabolic alkalosis
condition wherein an excess of bicarbonate causes the blood to be overly alkaline
metabolism
sum of all catabolic and anabolic reactions that take place in the body
metabotropic receptor
neurotransmitter receptor that involves a complex of proteins that cause metabolic changes in a cell
metaphase
second stage of mitosis (and meiosis), characterized by the linear alignment of sister chromatids in the center of the cell
metaphase plate
linear alignment of sister chromatids in the center of the cell, which takes place during metaphase
micelle
tiny lipid-transport compound composed of bile salts and phospholipids with a fatty acid and monoacylglyceride core
micturition
also called urination or voiding
microglia
glial cell type in the CNS that serves as the resident component of the immune system
microvillus
small projection of the plasma membrane of the absorptive cells of the small intestinal mucosa
middle ear
space within the temporal bone between the ear canal and bony labyrinth where the ossicles amplify sound waves from the tympanic membrane to the oval window
migrating motility complex
form of peristalsis in the small intestine
mineralocorticoids
hormones produced by the zona glomerulosa cells of the adrenal cortex that influence fluid and electrolyte balance
minerals
inorganic compounds required by the body to ensure proper function of the body
mitosis
division of genetic material, during which the cell nucleus breaks down and two new, fully functional, nuclei are formed
mitotic phase
phase of the cell cycle in which a cell undergoes mitosis
mitotic spindle
network of microtubules, originating from centrioles, that arranges and pulls apart chromosomes during mitosis
mixing wave
unique type of peristalsis that occurs in the stomach
monoglyceride molecules
lipid consisting of a single fatty acid chain attached to a glycerol backbone
monosaccharide
monomer of carbohydrate; also known as a simple sugar
molar
tooth used for crushing and grinding food
molecule
two or more atoms covalently bonded together
monosaccharide
smallest, monomeric sugar molecule
mons pubis
mound of fatty tissue located at the front of the vulva
morula
tightly packed sphere of blastomeres that has reached the uterus but has not yet implanted itself
motilin
hormone that initiates migrating motility complexes
motility
movement of food through the GI tract
mucosa
innermost lining of the alimentary canal
mucosal barrier
protective barrier that prevents gastric juice from destroying the stomach itself
mucous gland
group of cells that secrete mucous, a thick, slippery substance that keeps tissues moist and acts as a lubricant
mucous neck cell
gastric gland cell that secretes a uniquely acidic mucus
multipolar
shape of a neuron that has multiple processes—the axon and two or more dendrites
muscularis
muscle (skeletal or smooth) layer of the alimentary canal wall
muscarinic receptor
type of acetylcholine receptor protein that is characterized by also binding to muscarine and is a metabotropic receptor
muscarinic receptor
type of acetylcholine receptor protein that is characterized by also binding to muscarine and is a metabotropic receptor
mutation
change in the nucleotide sequence of DNA
myelin
lipid-rich insulating substance surrounding the axons of many neurons, allowing for faster transmission of electrical signals
myelin sheath
lipid-rich layer of insulation that surrounds an axon, formed by oligodendrocytes in the CNS and Schwann cells in the PNS; facilitates the transmission of electrical signals
myenteric plexus
(plexus of Auerbach) major nerve supply to alimentary canal wall; controls motility
myometrium
smooth muscle layer of uterus that allows for uterine contractions during labor and expulsion of menstrual blood
NADH
high-energy molecule needed for glycolysis
neonatal hypothyroidism
condition characterized by cognitive deficits, short stature, and other signs and symptoms in people born to women who were iodine-deficient during pregnancy
nephrons
functional units of the kidney that carry out all filtration and modification to produce urine; consist of renal corpuscles, proximal and distal convoluted tubules, and descending and ascending loops of Henle; drain into collecting ducts
nerve
cord-like bundle of axons located in the peripheral nervous system that transmits sensory input and response output to and from the central nervous system
nerve plexus
network of nerves without neuronal cell bodies included
neural plate
thickened layer of neuroepithelium that runs longitudinally along the dorsal surface of an embryo and gives rise to nervous system tissue
neural fold
elevated edge of the neural groove
neural tube
precursor to structures of the central nervous system, formed by the invagination and separation of neuroepithelium
neural tunic layer of the eye that contains nervous tissue, namely the retina neuropeptide neurotransmitter type that includes protein molecules and shorter chains of amino acids neuron neural tissue cell that is primarily responsible for generating and propagating electrical signals into, within, and out of the nervous system
neurotransmitter
chemical signal that is released from the synaptic end bulb of a neuron to cause a change in the target cell
neurulation
embryonic process that establishes the central nervous system
nicotinic receptor
type of acetylcholine receptor protein that is characterized by also binding to nicotine and is an ionotropic receptor
nicotinic receptor
type of acetylcholine receptor protein that is characterized by also binding to nicotine and is an ionotropic receptor
nicotinamide adenine dinucleotide (NAD)
coenzyme used to produce NADH
nociceptor
receptor cell that senses pain stimuli
node of Ranvier
gap between two myelinated regions of an axon, allowing for strengthening of the electrical signal as it propagates down the axon
nonspecific channel
channel that is not specific to one ion over another, such as a nonspecific cation channel that allows any positively charged ion across the membrane
norepinephrine
signaling molecule released as a neurotransmitter by most postganglionic sympathetic fibers as part of the sympathetic response, or as a hormone into the bloodstream from the adrenal medulla
norepinephrine
secondary catecholamine hormone secreted by the adrenal medulla in response to short-term stress; also called noradrenaline
notochord
rod-shaped, mesoderm-derived structure that provides support for growing fetus
nucleosidase
brush border enzyme that digests nucleotides
nucleotide
class of organic compounds composed of one or more phosphate groups, a pentose sugar, and a base
nucleus
in the nervous system, a localized collection of neuron cell bodies that are functionally related; a “center” of neural function
nucleus ambiguus
brain-stem nucleus that contains neurons that project through the vagus nerve to terminal ganglia in the thoracic cavity; specifically associated with the heart
occipital lobe
region of the cerebral cortex directly beneath the occipital bone of the cranium
occipital sinuses
dural sinuses along the edge of the occipital lobes of the cerebrum
oculomotor nerve
third cranial nerve; responsible for contraction of four of the extraocular muscles, the muscle in the upper eyelid, and pupillary constriction
odorant molecules
volatile chemicals that bind to receptor proteins in olfactory neurons to stimulate the sense of smell
olfaction
sense of smell
olfaction
special sense responsible for smell, which has a unique, direct connection to the cerebrum
olfactory bulb
central target of the first cranial nerve; located on the ventral surface of the frontal lobe in the cerebrum
olfactory epithelium
region of the nasal epithelium where olfactory neurons are located
olfactory nerve
first cranial nerve; responsible for the sense of smell
olfactory sensory neuron
receptor cell of the olfactory system, sensitive to the chemical stimuli of smell, the axons of which compose the first cranial nerve
oligodendrocyte
glial cell type in the CNS that provides the myelin insulation for axons in tracts
oliguria
below normal urine production of 400–500 mL/day
oocyte
cell that results from the division of the oogonium and undergoes meiosis I at the LH surge and meiosis II at fertilization to become a haploid ovum
oogenesis
process by which oogonia divide by mitosis to primary oocytes, which undergo meiosis to produce the secondary oocyte and, upon fertilization, the ovum
oogonia
ovarian stem cells that undergo mitosis during female fetal development to form primary oocytes
opsin
protein that contains the photosensitive cofactor retinal for phototransduction
optic disc
spot on the retina at which RGC axons leave the eye and blood vessels of the inner retina pass
optic nerve
second cranial nerve, which is responsible visual sensation
oral cavity
(also, buccal cavity) mouth
oral vestibule
part of the mouth bounded externally by the cheeks and lips, and internally by the gums and teeth
organ of Corti
structure in the cochlea in which hair cells transduce movements from sound waves into electrochemical signals
organic compound
substance that contains both carbon and hydrogen
organogenesis
development of the rudimentary structures of all of an embryo’s organs from the germ layers
oropharynx
part of the pharynx continuous with the oral cavity that functions in respiration and digestion
orthostatic reflex
sympathetic function that maintains blood pressure when standing to offset the increased effect of gravity
osmoreceptor
receptor cell that senses differences in the concentrations of bodily fluids on the basis of osmotic pressure
osmoreceptor
hypothalamic sensory receptor that is stimulated by changes in solute concentration (osmotic pressure) in the blood
ossicles
three small bones in the middle ear
otolith
layer of calcium carbonate crystals located on top of the otolithic membrane
otolithic membrane
gelatinous substance in the utricle and saccule of the inner ear that contains calcium carbonate crystals and into which the stereocilia of hair cells are embedded
outer segment
in the eye, the section of a photoreceptor that contains opsin molecules that transduce light stimuli
outer synaptic layer
layer in the retina at which photoreceptors connect to bipolar cells
oval window
membrane at the base of the cochlea where the stapes attaches, marking the beginning of the scala vestibuli
ovarian cycle
approximately 28-day cycle of changes in the ovary consisting of a follicular phase and a luteal phase
ovaries
female gonads that produce oocytes and sex steroid hormones (notably estrogen and progesterone)
ovulation
release of a secondary oocyte and associated granulosa cells from an ovary
ovum
haploid female gamete resulting from completion of meiosis II at fertilization
oxidation
loss of an electron
oxidation-reduction reaction
(also, redox reaction) pair of reactions in which an electron is passed from one molecule to another, oxidizing one and reducing the other
oxidative phosphorylation
process that converts high-energy NADH and FADH2 into ATP
oxytocin
hypothalamic hormone stored in the posterior pituitary gland and important in stimulating uterine contractions in labor, milk ejection during breastfeeding, and feelings of attachment (also produced in males)
palatoglossal arch
muscular fold that extends from the lateral side of the soft palate to the base of the tongue
palatopharyngeal arch
muscular fold that extends from the lateral side of the soft palate to the side of the pharynx
palpebral conjunctiva
membrane attached to the inner surface of the eyelids that covers the anterior surface of the cornea
pancreas
organ with both exocrine and endocrine functions located posterior to the stomach that is important for digestion and the regulation of blood glucose
pancreas
accessory digestive organ that secretes pancreatic juice
pancreatic amylase
enzyme secreted by the pancreas that completes the chemical digestion of carbohydrates in the small intestine
pancreatic islets
specialized clusters of pancreatic cells that have endocrine functions; also called islets of Langerhans
pancreatic lipase
enzyme secreted by the pancreas that participates in lipid digestion
pancreatic lipases
enzymes released from the pancreas that digest lipids in the diet
pancreatic nuclease
enzyme secreted by the pancreas that participates in nucleic acid digestion
pancreatic juice
secretion of the pancreas containing digestive enzymes and bicarbonate
papilla
for gustation, a bump-like projection on the surface of the tongue that contains taste buds
paracrine
chemical signal that elicits a response in neighboring cells; also called paracrine factor
perineurium
layer of connective tissue surrounding fascicles within a nerve
parasympathetic division
division of the autonomic nervous system responsible for restful and digestive functions
parathyroid glands
small, round glands embedded in the posterior thyroid gland that produce parathyroid hormone (PTH)
parathyroid hormone (PTH)
peptide hormone produced and secreted by the parathyroid glands in response to low blood calcium levels
paravertebral ganglia
autonomic ganglia superior to the sympathetic chain ganglia
parietal cell
gastric gland cell that secretes hydrochloric acid and intrinsic factor
parietal lobe
region of the cerebral cortex directly beneath the parietal bone of the cranium
parieto-occipital sulcus
groove in the cerebral cortex representing the border between the parietal and occipital cortices
parotid gland
one of a pair of major salivary glands located inferior and anterior to the ears
pectinate line
horizontal line that runs like a ring, perpendicular to the inferior margins of the anal sinuses
pedicels
finger-like projections of podocytes surrounding glomerular capillaries; interdigitate to form a filtration membrane
penis
male organ of copulation
pepsin
enzyme that begins to break down proteins in the stomach
pepsinogen
inactive form of pepsin
peptide bond
covalent bond formed by dehydration synthesis between two amino acids
peripheral nervous system (PNS)
anatomical division of the nervous system that is largely outside the cranial and vertebral cavities, namely all parts except the brain and spinal cord
peristalsis
muscular contractions and relaxations that propel food through the GI tract
peritubular capillaries
second capillary bed of the renal portal system; surround the proximal and distal convoluted tubules; associated with the vasa recta
permanent tooth
one of 32 adult teeth
perimetrium
outer epithelial layer of uterine wall
pH
negative logarithm of the hydrogen ion (H+) concentration of a solution
pharynx
throat
phenotype
physical or biochemical manifestation of the genotype; expression of the alleles
phosphatase
brush border enzyme that digests nucleotides
phosphodiesterase (PDE)
cytosolic enzyme that deactivates and degrades cAMP
phospholipid
a lipid compound in which a phosphate group is combined with a diglyceride
phosphorylation
addition of one or more phosphate groups to an organic compound
phosphorylation cascade
signaling event in which multiple protein kinases phosphorylate the next protein substrate by transferring a phosphate group from ATP to the protein
photoisomerization
chemical change in the retinal molecule that alters the bonding so that it switches from the 11-cis-retinal isomer to the all-trans-retinal isomer
photon
individual “packet” of light
photoreceptor
receptor cell specialized to respond to light stimuli
phrenic nerve
systemic nerve from the cervical plexus that enervates the diaphragm
physiological sphincter
sphincter consisting of circular smooth muscle indistinguishable from adjacent muscle but possessing differential innervations, permitting its function as a sphincter; structurally weak
pia mater
thin, innermost membrane of the meninges that directly covers the surface of the CNS
pineal gland
endocrine gland that secretes melatonin, which is important in regulating the sleep-wake cycle
pinealocyte
cell of the pineal gland that produces and secretes the hormone melatonin
pituitary dwarfism
disorder in children caused when abnormally low levels of GH result in growth retardation
pituitary gland
bean-sized organ suspended from the hypothalamus that produces, stores, and secretes hormones in response to hypothalamic stimulation (also called hypophysis)
placenta
organ that forms during pregnancy to nourish the developing fetus; also regulates waste and gas exchange between mother and fetus
placenta previa
low placement of fetus within uterus causes placenta to partially or completely cover the opening of the cervix as it grows
placentation
formation of the placenta; complete by weeks 14–16 of pregnancy
plasma osmolality
ratio of solutes to a volume of solvent in the plasma; plasma osmolality reflects a person’s state of hydration
plexus
network of nerves or nervous tissue
podocytes
cells forming finger-like processes; form the visceral layer of Bowman’s capsule; pedicels of the podocytes interdigitate to form a filtration membrane
polar body
smaller cell produced during the process of meiosis in oogenesis
polar molecule
molecule with regions that have opposite charges resulting from uneven numbers of electrons in the nuclei of the atoms participating in the covalent bond
polyspermy
penetration of an oocyte by more than one sperm
polysaccharides
complex carbohydrates made up of many monosaccharides
polyuria
urine production in excess of 2.5 L/day; may be caused by diabetes insipidus, diabetes mellitus, or excessive use of diuretics
porta hepatis
“gateway to the liver” where the hepatic artery and hepatic portal vein enter the liver
portal triad
bile duct, hepatic artery branch, and hepatic portal vein branch
postabsorptive state
also called the fasting state; the metabolic state occurring after digestion when food is no longer the body’s source of energy and it must rely on stored glycogen
postganglionic fiber
axon from a ganglionic neuron in the autonomic nervous system that projects to and synapses with the target effector; sometimes referred to as a postganglionic neuron
postsynaptic potential (PSP)
graded potential in the postsynaptic membrane caused by the binding of neurotransmitter to protein receptors
postcentral gyrus
ridge just posterior to the central sulcus, in the parietal lobe, where somatosensory processing initially takes place in the cerebrum
posterior columns
white matter of the spinal cord that lies between the posterior horns of the gray matter, sometimes referred to as the dorsal column; composed of axons of ascending tracts that carry sensory information up to the brain
posterior horn
gray matter region of the spinal cord in which sensory input arrives, sometimes referred to as the dorsal horn
posterior median sulcus
midline feature of the posterior spinal cord, marking the separation between right and left sides of the cord
posterolateral sulcus
feature of the posterior spinal cord marking the entry of posterior nerve roots and the separation between the posterior and lateral columns of the white matter
PP cell
minor cell type in the pancreas that secretes the hormone pancreatic polypeptide
precentral gyrus
primary motor cortex located in the frontal lobe of the cerebral cortex
precentral gyrus of the frontal cortex
region of the cerebral cortex responsible for generating motor commands, where the upper motor neuron cell body is located
prefrontal lobe
specific region of the frontal lobe anterior to the more specific motor function areas, which can be related to the early planning of movements and intentions to the point of being personality-type functions
preganglionic fiber
axon from a central neuron in the autonomic nervous system that projects to and synapses with a ganglionic neuron; sometimes referred to as a preganglionic neuron
prepuce
(also, foreskin) flap of skin that forms a collar around, and thus protects and lubricates, the glans penis; also referred as the foreskin
premolar
(also, bicuspid) transitional tooth used for mastication, crushing, and grinding food
premotor area
region of the frontal lobe responsible for planning movements that will be executed through the primary motor cortex
prevertebral ganglia
autonomic ganglia that are anterior to the vertebral column and functionally related to the sympathetic chain ganglia
proliferative phase
phase of the menstrual cycle in which the endometrium proliferates
primary follicles
ovarian follicles with a primary oocyte and one layer of cuboidal granulosa cells
primitive streak
indentation along the dorsal surface of the epiblast through which cells migrate to form the endoderm and mesoderm during gastrulation
primordial follicles
least developed ovarian follicles that consist of a single oocyte and a single layer of flat (squamous) granulosa cells
principal cell
found in collecting ducts and possess channels for the recovery or loss of sodium and potassium; under the control of aldosterone; also have aquaporin channels under ADH control to regulate recovery of water
process
in cells, an extension of a cell body; in the case of neurons, this includes the axon and dendrites
progesterone
predominantly female sex hormone important in regulating the female reproductive cycle and the maintenance of pregnancy
prolactin (PRL)
anterior pituitary hormone that promotes development of the mammary glands and the production of breast milk
propagation
movement of an action potential along the length of an axon
prophase
first stage of mitosis (and meiosis), characterized by breakdown of the nuclear envelope and condensing of the chromatin to form chromosomes
propulsion
voluntary process of swallowing and the involuntary process of peristalsis that moves food through the digestive tract
proprioception
general sensory perceptions providing information about location and movement of body parts; the “sense of the self”
proprioception
sense of position and movement of the body
proprioceptor
receptor cell that senses changes in the position and kinesthetic aspects of the body
prostate gland
doughnut-shaped gland at the base of the bladder surrounding the urethra and contributing fluid to semen during ejaculation
protein kinase
enzyme that initiates a phosphorylation cascade upon activation
proteolysis
process of breaking proteins into smaller peptides
proximal convoluted tubules (PCTs)
tortuous tubules receiving filtrate from Bowman’s capsule; most active part of the nephron in reabsorption and secretion
pseudostratified columnar epithelium
tissue that consists of a single layer of irregularly shaped and sized cells that give the appearance of multiple layers; found in ducts of certain glands and the upper respiratory tract
pulp cavity
deepest portion of a tooth, containing nerve endings and blood vessels
pupil
open hole at the center of the iris that light passes through into the eye
putamen
nucleus deep in the cerebrum that is part of the basal nuclei; along with the caudate, it is part of the striatum
polysaccharide
compound consisting of more than two carbohydrate monomers bonded by dehydration synthesis via glycosidic bonds
prostaglandin
lipid compound derived from fatty acid chains and important in regulating several body processes
protein
class of organic compounds that are composed of many amino acids linked together by peptide bonds
Punnett square
grid used to display all possible combinations of alleles transmitted by parents to offspring and predict the mathematical probability of offspring inheriting a given genotype
purine
nitrogen-containing base with a double ring structure; adenine and guanine
pyloric antrum
wider, more superior part of the pylorus
pyloric canal
narrow, more inferior part of the pylorus
pyloric sphincter
sphincter that controls stomach emptying
pylorus
lower, funnel-shaped part of the stomach that is continuous with the duodenum
pyrimidine
nitrogen-containing base with a single ring structure; cytosine, thiamine, and uracil
pyruvate
three-carbon end product of glycolysis and starting material that is converted into acetyl CoA that enters the Krebs cycle
quickening
fetal movements that are strong enough to be felt by the mother
radial nerve
systemic nerve of the arm, the distal component of which is located near the radial bone
recessive
describes a trait that is only expressed in homozygous form and is masked in heterozygous form
recessive lethal
inheritance pattern in which individuals with two copies of a lethal allele do not survive in utero or have a shortened life span
receptor cell
cell that transduces environmental stimuli into neural signals
receptor potential
graded potential in a specialized sensory cell that directly causes the release of neurotransmitter without an intervening action potential
rectal valve
one of three transverse folds in the rectum where feces is separated from flatus
rectum
part of the large intestine between the sigmoid colon and anal canal
reduction
gaining of an electron
refractory period
time after the initiation of an action potential when another action potential cannot be generated
relative refractory period
time during the refractory period when a new action potential can only be initiated by a stronger stimulus than the current action potential because voltage-gated K+ channels are not closed
renal columns
extensions of the renal cortex into the renal medulla; separates the renal pyramids; contains blood vessels and connective tissues
renal corpuscle
consists of the glomerulus and Bowman’s capsule
renal cortex
outer part of kidney containing all of the nephrons; some nephrons have loops of Henle extending into the medulla
renal fat pad
adipose tissue between the renal fascia and the renal capsule that provides protective cushioning to the kidney
renal hilum
recessed medial area of the kidney through which the renal artery, renal vein, ureters, lymphatics, and nerves pass
renal papillae
medullary area of the renal pyramids where collecting ducts empty urine into the minor calyces
renal pyramids
six to eight cone-shaped tissues in the medulla of the kidney containing collecting ducts and the loops of Henle of juxtamedullary nephrons
renin
enzyme produced by juxtaglomerular cells in response to decreased blood pressure or sympathetic nervous activity; catalyzes the conversion of angiotensinogen into angiotensin I
repolarization
return of the membrane potential to its normally negative voltage at the end of the action potential
resistance
property of an axon that relates to the ability of particles to diffuse through the cytoplasm; this is inversely proportional to the fiber diameter
response
nervous system function that causes a target tissue (muscle or gland) to produce an event as a consequence to stimuli
rest and digest
set of functions associated with the parasympathetic system that lead to restful actions and digestion
resting membrane potential
the difference in voltage measured across a cell membrane under steady-state conditions, typically -70 mV
respiratory acidosis
condition wherein an excess of carbonic acid or CO2 causes the blood to be overly acidic
respiratory alkalosis
condition wherein a deficiency of carbonic acid/CO2 levels causes the blood to be overly alkaline
reticular formation
diffuse region of gray matter throughout the brain stem that regulates sleep, wakefulness, and states of consciousness
reticular lamina
matrix containing collagen and elastin secreted by connective tissue; a component of the basement membrane
reticuloendothelial cell
(also, Kupffer cell) phagocyte in hepatic sinusoids that filters out material from venous blood from the alimentary canal
retina
nervous tissue of the eye at which phototransduction takes place
retinal
cofactor in an opsin molecule that undergoes a biochemical change when struck by a photon (pronounced with a stress on the last syllable)
retinal ganglion cell (RGC)
neuron of the retina that projects along the second cranial nerve
retroperitoneal
located posterior to the peritoneum
retroperitoneal
outside the peritoneal cavity; in the case of the kidney and ureters, between the parietal peritoneum and the abdominal wall
ribonuclease
pancreatic enzyme that digests RNA
ribonucleic acid (RNA)
ribose-containing nucleotide that helps manifest the genetic code as protein
right colic flexure
(also, hepatic flexure) point, at the inferior surface of the liver, where the ascending colon turns abruptly to the left
rhodopsin
photopigment molecule found in the rod photoreceptors
rod photoreceptor
one of the two types of retinal receptor cell that is specialized for low-light vision
root
portion of a tooth embedded in the alveolar processes beneath the gum line
round window
membrane that marks the end of the scala tympani
ruga
fold of alimentary canal mucosa and submucosa in the empty stomach and other organs
rugae
(of the vagina) folds of skin in the vagina that allow it to stretch during intercourse and childbirth
S phase
stage of the cell cycle during which DNA replication occurs
saccharolytic fermentation
anaerobic decomposition of carbohydrates
saccule
structure of the inner ear responsible for transducing linear acceleration in the vertical plane
sacral micturition center
group of neurons in the sacral region of the spinal cord that controls urination; acts reflexively unless its action is modified by higher brain centers to allow voluntary urination
sacral plexus
nerve plexus associated with the lower lumbar and sacral spinal nerves
saliva
aqueous solution of proteins and ions secreted into the mouth by the salivary glands
salivary amylase
digestive enzyme in saliva that acts on starch
salivary amylase
digestive enzyme that is found in the saliva and begins the digestion of carbohydrates in the mouth
salivary gland
an exocrine gland that secretes a digestive fluid called saliva
salivation
secretion of saliva
saltatory conduction
quick propagation of the action potential along a myelinated axon owing to voltage-gated Na+ channels being present only at the nodes of Ranvier
satellite cell
glial cell type in the PNS that provides support for neurons in the ganglia
scala tympani
portion of the cochlea that extends from the apex to the round window
scala vestibuli
portion of the cochlea that extends from the oval window to the apex
Schwann cell
glial cell type in the PNS that provides the myelin insulation for axons in nerves
saphenous nerve
systemic nerve of the lower anterior leg that is a branch from the femoral nerve
sciatic nerve
systemic nerve from the sacral plexus that is a combination of the tibial and fibular nerves and extends across the hip joint and gluteal region into the upper posterior leg
sciatica
painful condition resulting from inflammation or compression of the sciatic nerve or any of the spinal nerves that contribute to it
sclera
white of the eye
scrotum
external pouch of skin and muscle that houses the testes
second messenger
molecule that initiates a signaling cascade in response to hormone binding on a cell membrane receptor and activation of a G protein
secondary follicles
ovarian follicles with a primary oocyte and multiple layers of granulosa cells
secretin
hormone released in the small intestine to aid in digestion
secretory phase
phase of the menstrual cycle in which the endometrium secretes a nutrient-rich fluid in preparation for implantation of an embryo
segmentation
alternating contractions and relaxations of non-adjacent segments of the intestine that move food forward and backward, breaking it apart and mixing it with digestive juices
semen
ejaculatory fluid composed of sperm and secretions from the seminal vesicles, prostate, and bulbourethral glands
semicircular canals
structures within the inner ear responsible for transducing rotational movement information
seminal vesicle
gland that produces seminal fluid, which contributes to semen
seminiferous tubules
tube structures within the testes where spermatogenesis occurs
sensation
nervous system function that receives information from the environment and translates it into the electrical signals of nervous tissue
sensory modality
a particular system for interpreting and perceiving environmental stimuli by the nervous system
serosa
outermost layer of the alimentary canal wall present in regions within the abdominal cavity
Sertoli cells
cells that support germ cells through the process of spermatogenesis; a type of sustentacular cell
serous gland
group of cells within the serous membrane that secrete a lubricating substance onto the surface
sex chromosomes
pair of chromosomes involved in sex determination; in males, the XY chromosomes; in females, the XX chromosomes
shunt
circulatory shortcut that diverts the flow of blood from one region to another
simple columnar epithelium
tissue that consists of a single layer of column-like cells; promotes secretion and absorption in tissues and organs
simple cuboidal epithelium
tissue that consists of a single layer of cube-shaped cells; promotes secretion and absorption in ducts and tubules
simple squamous epithelium
tissue that consists of a single layer of flat scale-like cells; promotes diffusion and filtration across surface
sigmoid colon
end portion of the colon, which terminates at the rectum
sigmoid sinuses
dural sinuses that drain directly into the jugular veins
sister chromatid
one of a pair of identical chromosomes, formed during DNA replication
size exclusion
principle of selectively allowing ions through a channel on the basis of their relative size
small intestine
section of the alimentary canal where most digestion and absorption occurs
sodium bicarbonate
anion released into the small intestine to neutralize the pH of the food from the stomach
soft palate
posterior region of the bottom portion of the nasal cavity that consists of skeletal muscle
solution
homogeneous liquid mixture in which a solute is dissolved into molecules within a solvent
soma
in neurons, that portion of the cell that contains the nucleus; the cell body, as opposed to the cell processes (axons and dendrites)
somatic cell
all cells of the body excluding gamete cells
somatic nervous system (SNS)
functional division of the nervous system that is concerned with conscious perception, voluntary movement, and skeletal muscle reflexes
somite
one of the paired, repeating blocks of tissue located on either side of the notochord in the early embryo
somatosensation
general senses related to the body, usually thought of as the senses of touch, which would include pain, temperature, and proprioception
somatosensation
general sense associated with modalities lumped together as touch
spatial summation
combination of graded potentials across the neuronal cell membrane caused by signals from separate presynaptic elements that add up to initiate an action potential
special sense
any sensory system associated with a specific organ structure, namely smell, taste, sight, hearing, and balance
specific gravity
weight of a liquid compared to pure water, which has a specific gravity of 1.0; any solute added to water will increase its specific gravity
sperm
(also, spermatozoon) male gamete
spermatic cord
bundle of nerves and blood vessels that supplies the testes; contains ductus deferens
spermatid
immature sperm cells produced by meiosis II of secondary spermatocytes
spermatocyte
cell that results from the division of spermatogonium and undergoes meiosis I and meiosis II to form spermatids
spermatogenesis
formation of new sperm, occurs in the seminiferous tubules of the testes
spermatogonia
(singular = spermatogonium) diploid precursor cells that become sperm
spermiogenesis
transformation of spermatids to spermatozoa during spermatogenesis
spinal cord
organ of the central nervous system found within the vertebral cavity and connected with the periphery through spinal nerves; mediates reflex behaviors
spinal accessory nerve
eleventh cranial nerve; responsible for contraction of neck muscles
spinal nerve
one of 31 nerves connected to the spinal cord
spiral ganglion
location of neuronal cell bodies that transmit auditory information along the eighth cranial nerve
stage of exhaustion
stage three of the general adaptation syndrome; the body’s long-term response to stress mediated by the hormones of the adrenal cortex
stage of resistance
stage two of the general adaptation syndrome; the body’s continued response to stress after stage one diminishes
stapes
(also, stirrup) ossicle of the middle ear that is attached to the inner ear
steroid
(also, sterol) lipid compound composed of four hydrocarbon rings bonded to a variety of other atoms and molecules
stereocilia
array of apical membrane extensions in a hair cell that transduce movements when they are bent
stimulus
an event in the external or internal environment that registers as activity in a sensory neuron
stomach
alimentary canal organ that contributes to chemical and mechanical digestion of food from the esophagus before releasing it, as chyme, to the small intestine
straight sinus
dural sinus that drains blood from the deep center of the brain to collect with the other sinuses
stratified columnar epithelium
tissue that consists of two or more layers of column-like cells, contains glands and is found in some ducts
stratified cuboidal epithelium
tissue that consists of two or more layers of cube-shaped cells, found in some ducts
stratified squamous epithelium
tissue that consists of multiple layers of cells with the most apical being flat scale-like cells; protects surfaces from abrasion
striatum
the caudate and putamen collectively, as part of the basal nuclei, which receive input from the cerebral cortex
subarachnoid space
space between the arachnoid mater and pia mater that contains CSF and the fibrous connections of the arachnoid trabeculae
subcortical nucleus
all the nuclei beneath the cerebral cortex, including the basal nuclei and the basal forebrain
sublingual gland
one of a pair of major salivary glands located beneath the tongue
submandibular gland
one of a pair of major salivary glands located in the floor of the mouth
submodality
specific sense within a broader major sense such as sweet as a part of the sense of taste, or color as a part of vision
submucosa
layer of dense connective tissue in the alimentary canal wall that binds the overlying mucosa to the underlying muscularis
submucosal plexus
(plexus of Meissner) nerve supply that regulates activity of glands and smooth muscle
substantia nigra pars compacta
nuclei within the basal nuclei that release dopamine to modulate the function of the striatum; part of the motor pathway
substantia nigra pars reticulata
nuclei within the basal nuclei that serve as an output center of the nuclei; part of the motor pathway
substrate
reactant in an enzymatic reaction
subthalamus
nucleus within the basal nuclei that is part of the indirect pathway
sucrase
brush border enzyme that breaks down sucrose into glucose and fructose
sulcus
groove formed by convolutions in the surface of the cerebral cortex
summate
to add together, as in the cumulative change in postsynaptic potentials toward reaching threshold in the membrane, either across a span of the membrane or over a certain amount of time
superior oblique
extraocular muscle responsible for medial rotation of the eye
superior rectus
extraocular muscle responsible for looking up
superior sagittal sinus
dural sinus that runs along the top of the longitudinal fissure and drains blood from the majority of the outer cerebrum
superior colliculus
half of the midbrain tectum that is responsible for aligning visual, auditory, and somatosensory spatial perceptions
superior cervical ganglion
one of the paravertebral ganglia of the sympathetic system that projects to the head
superior mesenteric ganglion
one of the collateral ganglia of the sympathetic system that projects to the digestive system
suspension
liquid mixture in which particles distributed in the liquid settle out over time
suspensory ligaments
bands of connective tissue that suspend the breast onto the chest wall by attachment to the overlying dermis
synaptic cleft
small gap between cells in a chemical synapse where neurotransmitter diffuses from the presynaptic element to the postsynaptic element
synaptic end bulb
swelling at the end of an axon where neurotransmitter molecules are released onto a target cell across a synapse
synapse
narrow junction across which a chemical signal passes from neuron to the next, initiating a new electrical signal in the target cell
sympathetic chain ganglia
series of ganglia adjacent to the vertebral column that receive input from central sympathetic neurons
sympathetic chain ganglia
autonomic ganglia in a chain along the anterolateral aspect of the vertebral column that are responsible for contributing to homeostatic mechanisms of the autonomic nervous system
sympathetic division
division of the autonomic nervous system associated with the fight-or-flight response
syncytiotrophoblast
superficial cells of the trophoblast that fuse to form a multinucleated body that digests endometrial cells to firmly secure the blastocyst to the uterine wall
systemic nerve
nerve in the periphery distal to a nerve plexus or spinal nerve
target effector
organ, tissue, or gland that will respond to the control of an autonomic or somatic or endocrine signal
taste buds
structures within a papilla on the tongue that contain gustatory receptor cells
tectorial membrane
component of the organ of Corti that lays over the hair cells, into which the stereocilia are embedded
tectum
region of the midbrain, thought of as the roof of the cerebral aqueduct, which is subdivided into the inferior and superior colliculi
tegmentum
region of the midbrain, thought of as the floor of the cerebral aqueduct, which continues into the pons and medulla as the floor of the fourth ventricle
terminal ganglia
ganglia of the parasympathetic division of the autonomic system, which are located near or within the target effector, the latter also known as intramural ganglia
terminal ganglion
autonomic ganglia that are near or within the walls of organs that are responsible for contributing to homeostatic mechanisms of the autonomic nervous system
tertiary follicles
(also, antral follicles) ovarian follicles with a primary or secondary oocyte, multiple layers of granulosa cells, and a fully formed antrum
telophase
final stage of mitosis (and meiosis), preceding cytokinesis, characterized by the formation of two new daughter nuclei
temporal lobe
region of the cerebral cortex directly beneath the temporal bone of the cranium
temporal summation
combination of graded potentials at the same location on a neuron resulting in a strong signal from one input
tenia coli
one of three smooth muscle bands that make up the longitudinal muscle layer of the muscularis in all of the large intestine except the terminal end
terminal electron acceptor
oxygen, the recipient of the free hydrogen at the end of the electron transport chain
testes
(singular = testis) male gonads
testosterone
steroid hormone secreted by the male testes and important in the maturation of sperm cells, growth and development of the male reproductive system, and the development of male secondary sex characteristics
thalamus
major region of the diencephalon that is responsible for relaying information between the cerebrum and the hindbrain, spinal cord, and periphery
thalamus
region of the central nervous system that acts as a relay for sensory pathways
theca cells
estrogen-producing cells in a maturing ovarian follicle
thermoreceptor
type of sensory receptor capable of transducing temperature stimuli into neural action potentials
thermoreceptor
sensory receptor specialized for temperature stimuli
third ventricle
portion of the ventricular system that is in the region of the diencephalon
thoracolumbar system
alternate name for the sympathetic division of the autonomic nervous system that is based on the anatomical location of central neurons in the lateral horn of the thoracic and upper lumbar spinal cord
threshold
membrane voltage at which an action potential is initiated
thymosins
hormones produced and secreted by the thymus that play an important role in the development and differentiation of T cells
thymus
organ that is involved in the development and maturation of T-cells and is particularly active during infancy and childhood
thyroid gland
large endocrine gland responsible for the synthesis of thyroid hormones
thyroid-stimulating hormone (TSH)
anterior pituitary hormone that triggers secretion of thyroid hormones by the thyroid gland (also called thyrotropin)
thyroxine
(also, tetraiodothyronine, T4) amino acid–derived thyroid hormone that is more abundant but less potent than T3 and often converted to T3 by target cells
tibial nerve
systemic nerve of the posterior leg that begins as part of the sciatic nerve
tight junction
forms an impermeable barrier between cells
tongue
accessory digestive organ of the mouth, the bulk of which is composed of skeletal muscle
topographical
relating to positional information
tract
bundle of axons in the central nervous system having the same function and point of origin
trait
variation of an expressed characteristic
transamination
transfer of an amine group from one molecule to another as a way to turn nitrogen waste into ammonia so that it can enter the urea cycle
transduction
process of changing an environmental stimulus into the electrochemical signals of the nervous system
transitional epithelium
form of stratified epithelium found in the urinary tract, characterized by an apical layer of cells that change shape in response to the presence of urine
transverse colon
part of the colon between the ascending colon and the descending colon
transverse sinuses
dural sinuses that drain along either side of the occipital–cerebellar space
tricarboxylic acid cycle (TCA)
also called the Krebs cycle or the citric acid cycle; converts pyruvate into CO2 and high-energy FADH2, NADH, and ATP molecules
trigeminal ganglion
sensory ganglion that contributes sensory fibers to the trigeminal nerve
trigeminal nerve
fifth cranial nerve; responsible for cutaneous sensation of the face and contraction of the muscles of mastication
trigone
area at the base of the bladder marked by the two ureters in the posterior–lateral aspect and the urethral orifice in the anterior aspect oriented like points on a triangle
triglyceride
lipid compound composed of a glycerol molecule bonded with three fatty acid chains
triglycerides
lipids, or fats, consisting of three fatty acid chains attached to a glycerol backbone
triiodothyronine
(also, T3) amino acid–derived thyroid hormone that is less abundant but more potent than T4
trochlea
cartilaginous structure that acts like a pulley for the superior oblique muscle
trochlear nerve
fourth cranial nerve; responsible for contraction of one of the extraocular muscles
trophoblast
fluid-filled shell of squamous cells destined to become the chorionic villi, placenta, and associated fetal membranes
trypsin
pancreatic enzyme that activates chymotrypsin and digests protein
trypsinogen
proenzyme form of trypsin
tympanic membrane
ear drum
umami
taste submodality for sensitivity to the concentration of amino acids; also called the savory sense
umbilical cord
connection between the developing conceptus and the placenta; carries deoxygenated blood and wastes from the fetus and returns nutrients and oxygen from the mother
utricle
structure of the inner ear responsible for transducing linear acceleration in the horizontal plane
ulnar nerve
systemic nerve of the arm located close to the ulna, a bone of the forear
unipolar
shape of a neuron which has only one process that includes both the axon and dendrite
upper esophageal sphincter
skeletal muscle sphincter that regulates food movement from the pharynx to the esophagus
upper motor neuron
first neuron in the motor command pathway with its cell body in the cerebral cortex that synapses on the lower motor neuron in the spinal cord
upregulation
increase in the number of hormone receptors, typically in response to chronically reduced levels of a hormone
urea cycle
process that converts potentially toxic nitrogen waste into urea that can be eliminated through the kidneys
urethra
transports urine from the bladder to the outside environment
urinalysis
analysis of urine to diagnose disease
urochrome
heme-derived pigment that imparts the typical yellow color of urine
uterine tubes
(also, fallopian tubes or oviducts) ducts that facilitate transport of an ovulated oocyte to the uterus
uterus
muscular hollow organ in which a fertilized egg develops into a fetus
vagina
tunnel-like organ that provides access to the uterus for the insertion of semen and from the uterus for the birth of a baby
vagus nerve
tenth cranial nerve; responsible for the autonomic control of organs in the thoracic and upper abdominal cavities
Valsalva’s maneuver
voluntary contraction of the diaphragm and abdominal wall muscles and closing of the glottis, which increases intra-abdominal pressure and facilitates defecation
varicosity
structure of some autonomic connections that is not a typical synaptic end bulb, but a string of swellings along the length of a fiber that makes a network of connections with the target effector
vasa recta
branches of the efferent arterioles that parallel the course of the loops of Henle and are continuous with the peritubular capillaries; with the glomerulus, form a portal system
vascular tunic
middle layer of the eye primarily composed of connective tissue with a rich blood supply
ventral (anterior) nerve root
axons emerging from the anterior or lateral horns of the spinal cord
ventricle
central cavity within the brain where CSF is produced and circulates
ventricles
remnants of the hollow center of the neural tube that are spaces for cerebrospinal fluid to circulate through the brain
vernix caseosa
waxy, cheese-like substance that protects the delicate fetal skin until birth
vertebral arteries
arteries that ascend along either side of the vertebral column through the transverse foramina of the cervical vertebrae and enter the cranium through the foramen magnum
vestibular ganglion
location of neuronal cell bodies that transmit equilibrium information along the eighth cranial nerve
vestibule
in the ear, the portion of the inner ear responsible for the sense of equilibrium
vestibulocochlear nerve
eighth cranial nerve; responsible for the sensations of hearing and balance
villus
projection of the mucosa of the small intestine
visceral sense
sense associated with the internal organs
vision
special sense of sight based on transduction of light stimuli
visual acuity
property of vision related to the sharpness of focus, which varies in relation to retinal position
vitamins
organic compounds required by the body to perform biochemical reactions like metabolism and bone, cell, and tissue growth
vitreous humor
viscous fluid that fills the posterior chamber of the eye
voltage-gated channel
ion channel that opens because of a change in the charge distributed across the membrane where it is located
voluntary phase
initial phase of deglutition, in which the bolus moves from the mouth to the oropharynx
vulva
external female genitalia
white matter
regions of the nervous system containing mostly myelinated axons, making the tissue appear white because of the high lipid content of myelin
white rami communicantes
(singular = ramus communicans) myelinated structures that provide a short connection from a sympathetic chain ganglion to the spinal nerve that contains the preganglionic sympathetic fiber
X-linked
pattern of inheritance in which an allele is carried on the X chromosome of the 23rd pair
X-linked dominant
pattern of dominant inheritance that corresponds to a gene on the X chromosome of the 23rd pair
X-linked recessive
pattern of recessive inheritance that corresponds to a gene on the X chromosome of the 23rd pair
yolk sac
membrane associated with primitive circulation to the developing embryo; source of the first blood cells and germ cells and contributes to the umbilical cord structure
zona fasciculata
intermediate region of the adrenal cortex that produce hormones called glucocorticoids
zona glomerulosa
most superficial region of the adrenal cortex, which produces the hormones collectively referred to as mineralocorticoids
zona pellucida
thick, gel-like glycoprotein membrane that coats the oocyte and must be penetrated by sperm before fertilization can occur
zona reticularis
deepest region of the adrenal cortex, which produces the steroid sex hormones called androgens
zonule fibers
fibrous connections between the ciliary body and the lens
zygote
fertilized egg; a diploid cell resulting from the fertilization of haploid gametes from the male and female lines
α-dextrin
breakdown product of starch
α-dextrinase
brush border enzyme that acts on α-dextrins
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Learning Objectives

By the end of this section, you will be able to:
  • Identify and distinguish between the types of connective tissue: proper, supportive, and fluid
  • Explain the functions of connective tissues

As may be obvious from its name, one of the major functions of connective tissue is to connect tissues and organs. Unlike epithelial tissue, which is composed of cells closely packed with little or no extracellular space in between, connective tissue cells are dispersed in a matrix. The matrix usually includes a large amount of extracellular material produced by the connective tissue cells that are embedded within it. The matrix plays a major role in the functioning of this tissue. The major component of the matrix is a ground substance often crisscrossed by protein fibers. This ground substance is usually a fluid, but it can also be mineralized and solid, as in bones. Connective tissues come in a vast variety of forms, yet they typically have in common three characteristic components: cells, large amounts of amorphous ground substance, and protein fibers. The amount and structure of each component correlates with the function of the tissue, from the rigid ground substance in bones supporting the body to the inclusion of specialized cells; for example, a phagocytic cell that engulfs pathogens and also rids tissue of cellular debris.

Functions of Connective Tissues

Connective tissues perform many functions in the body, but most importantly, they support and connect other tissues; from the connective tissue sheath that surrounds muscle cells, to the tendons that attach muscles to bones, and to the skeleton that supports the positions of the body. Protection is another major function of connective tissue, in the form of fibrous capsules and bones that protect delicate organs and, of course, the skeletal system. Specialized cells in connective tissue defend the body from microorganisms that enter the body. Transport of fluid, nutrients, waste, and chemical messengers is ensured by specialized fluid connective tissues, such as blood and lymph. Adipose cells store surplus energy in the form of fat and contribute to the thermal insulation of the body.

Embryonic Connective Tissue

All connective tissues derive from the mesodermal layer of the embryo (see Chapter 4.1 Figure 2). The first connective tissue to develop in the embryo is mesenchyme, the stem cell line from which all connective tissues are later derived. Clusters of mesenchymal cells are scattered throughout adult tissue and supply the cells needed for replacement and repair after a connective tissue injury. A second type of embryonic connective tissue forms in the umbilical cord, called mucous connective tissue or Wharton’s jelly. This tissue is no longer present after birth, leaving only scattered mesenchymal cells throughout the body.

Classification of Connective Tissues

The three broad categories of connective tissue are classified according to the characteristics of their ground substance and the types of fibers found within the matrix (Table 1). Connective tissue proper includes loose connective tissue and dense connective tissue. Both tissues have a variety of cell types and protein fibers suspended in a viscous ground substance. Dense connective tissue is reinforced by bundles of fibers that provide tensile strength, elasticity, and protection. In loose connective tissue, the fibers are loosely organized, leaving large spaces in between. Supportive connective tissue—bone and cartilage—provide structure and strength to the body and protect soft tissues. A few distinct cell types and densely packed fibers in a matrix characterize these tissues. In bone, the matrix is rigid and described as calcified because of the deposited calcium salts. In fluid connective tissue, in other words, lymph and blood, various specialized cells circulate in a watery fluid containing salts, nutrients, and dissolved proteins.

Connective Tissue Examples (Table 1)
Connective tissue proper Supportive connective tissue Fluid connective tissue
Loose connective tissue
  • Areolar
  • Adipose
  • Reticular
 Cartilage
  • Hyaline
  • Fibrocartilage
  • Elastic
 Blood
Dense connective tissue
  • Regular elastic
  • Irregular elastic
 Bones
  • Compact bone
  • Cancellous bone
 Lymph

Connective Tissue Proper

Fibroblasts are present in all connective tissue proper (Figure 1). Fibrocytes, adipocytes, and mesenchymal cells are fixed cells, which means they remain within the connective tissue. Other cells move in and out of the connective tissue in response to chemical signals. Macrophages, mast cells, lymphocytes, plasma cells, and phagocytic cells are found in connective tissue proper but are actually part of the immune system protecting the body.

[caption id="" align="aligncenter" width="650"]The left image shows a diagram of connective tissue. As a whole, the connective tissue appears somewhat disorganized, with fibers and cells mixed together heterogeneously. There are many open spaces between the embedded elements, suggesting that the connective tissue is somewhat loosely packed. The thickest fibers are collagen fibers; the thinner fibers are elastic fibers. Both the collagen fibers and the elastic fibers crisscross randomly throughout the tissue. In addition, a net of reticular fibers appear in the upper part of the diagram. Two yellow and oval shaped adipocytes are embedded below the reticular fiber net, with a small dark nucleus squeezed into one corner of the cell. A mesenchymal cell is next to one of the adipocytes. The cell is rectangular and has four projections stemming from each corner of the cell. The projections appear to attach to the nearby collagen fibers. A fibroblast is located at the center of the diagram. The fibroblast appears similar to the mesenchymal cell, except that it is larger and has more projections. Finally, a white macrophage is in the lower right of the diagram. The macrophage is a white, oval shaped disc with a prominent nucleus. The right diagram is a micrograph of connective tissue. The tissue is mostly stained pink, however, the thick collagen fibers crisscrossing the tissue are white. Five adipocytes also appear white, except for their cell membrane and nucleus, which stained dark. A mesenchymal cell occupies the space between two adipocytes. It stains a very deep purple, but its shape is unclear in the micrograph. A fibrocyte is also visible as an oval shaped cell with a deep purple nucleus. Figure 1. Connective Tissue Proper. Fibroblasts produce this fibrous tissue. Connective tissue proper includes the fixed cells fibrocytes, adipocytes, and mesenchymal cells. LM × 400. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Cell Types

The most abundant cell in connective tissue proper is the fibroblast. Polysaccharides and proteins secreted by fibroblasts combine with extra-cellular fluids to produce a viscous ground substance that, with embedded fibrous proteins, forms the extra-cellular matrix. As you might expect, a fibrocyte, a less active form of fibroblast, is the second most common cell type in connective tissue proper.

Adipocytes are cells that store lipids as droplets that fill most of the cytoplasm. There are two basic types of adipocytes: white and brown. The brown adipocytes store lipids as many droplets, and have high metabolic activity. In contrast, white fat adipocytes store lipids as a single large drop and are metabolically less active. Their effectiveness at storing large amounts of fat is witnessed in obese individuals. The number and type of adipocytes depends on the tissue and location, and vary among individuals in the population. The mesenchymal cell is a multipotent adult stem cell. These cells can differentiate into any type of connective tissue cells needed for repair and healing of damaged tissue.

The macrophage cell is a large cell derived from a monocyte, a type of blood cell, which enters the connective tissue matrix from the blood vessels. The macrophage cells are an essential component of the immune system, which is the body’s defense against potential pathogens and degraded host cells. When stimulated, macrophages release cytokines, small proteins that act as chemical messengers. Cytokines recruit other cells of the immune system to infected sites and stimulate their activities. Roaming, or free, macrophages move rapidly by amoeboid movement, engulfing infectious agents and cellular debris. In contrast, fixed macrophages are permanent residents of their tissues.

The mast cell, found in connective tissue proper, has many cytoplasmic granules. These granules contain the chemical signals histamine and heparin. When irritated or damaged, mast cells release histamine, an inflammatory mediator, which causes vasodilation and increased blood flow at a site of injury or infection, along with itching, swelling, and redness you recognize as an allergic response. Like blood cells, mast cells are derived from hematopoietic stem cells and are part of the immune system.

Connective Tissue Fibers and Ground Substance

Three main types of fibers are secreted by fibroblasts: collagen fibers, elastic fibers, and reticular fibers. Collagen fiber is made from fibrous protein subunits linked together to form a long and straight fiber. Collagen fibers, while flexible, have great tensile strength, resist stretching, and give ligaments and tendons their characteristic resilience and strength. These fibers hold connective tissues together, even during the movement of the body.

Elastic fiber contains the protein elastin along with lesser amounts of other proteins and glycoproteins. The main property of elastin is that after being stretched or compressed, it will return to its original shape. Elastic fibers are prominent in elastic tissues found in skin and the elastic ligaments of the vertebral column.

Reticular fiber is also formed from the same protein subunits as collagen fibers; however, these fibers remain narrow and are arrayed in a branching network. They are found throughout the body, but are most abundant in the reticular tissue of soft organs, such as liver and spleen, where they anchor and provide structural support to the parenchyma (the functional cells, blood vessels, and nerves of the organ).

All of these fiber types are embedded in ground substance. Secreted by fibroblasts, ground substance is made of polysaccharides, specifically hyaluronic acid, and proteins. These combine to form a proteoglycan with a protein core and polysaccharide branches. The proteoglycan attracts and traps available moisture forming the clear, viscous, colorless matrix you now know as ground substance.

Loose Connective Tissue

Loose connective tissue is found between many organs where it acts both to absorb shock and bind tissues together. It allows water, salts, and various nutrients to diffuse through to adjacent or imbedded cells and tissues.

[caption id="" align="alignright" width="500"]Image A shows a collection of yellow adipocytes that do not have a consistent shape or size, however, most have the general appearance of a kernel of corn with a wide end that tapers to a point. Each adipocyte has a nucleus occupying a small area on one side of the cell. Nothing else is visible within the cells. Image B shows a micrograph of adipose tissue. Here, the adipocytes are stained purple. However, only their edges and their nuclei stain, giving the adipose tissue a honeycomb appearance. The adipocytes in the micrograph are large and round, but still show a diversity of shapes and sizes. The nucleus appears as a dark staining area very close to the cell membrane.Figure 2. Adipose Tissue. This is a loose connective tissue that consists of fat cells with little extracellular matrix. It stores fat for energy and provides insulation. LM × 800. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption] Adipose tissue consists mostly of fat storage cells, with little extracellular matrix (Figure 2). A large number of capillaries allow rapid storage and mobilization of lipid molecules. White adipose tissue is most abundant. It can appear yellow and owes its color to carotene and related pigments from plant food. White fat contributes mostly to lipid storage and can serve as insulation from cold temperatures and mechanical injuries. White adipose tissue can be found protecting the kidneys and cushioning the back of the eye. Brown adipose tissue is more common in infants, hence the term “baby fat.” In adults, there is a reduced amount of brown fat and it is found mainly in the neck and clavicular regions of the body. The many mitochondria in the cytoplasm of brown adipose tissue help explain its efficiency at metabolizing stored fat. Brown adipose tissue is thermogenic, meaning that as it breaks down fats, it releases metabolic heat, rather than producing adenosine triphosphate (ATP), a key molecule used in metabolism.  

Areolar tissue shows little specialization. It contains all the cell types and fibers previously described and is distributed in a random, web-like fashion. It fills the spaces between muscle fibers, surrounds blood and lymph vessels, and supports organs in the abdominal cavity. Areolar tissue underlies most epithelia and represents the connective tissue component of epithelial membranes, which are described further in a later section.

[caption id="" align="alignleft" width="500"]This figure shows reticular tissue alongside a micrograph. The diagram shows a series of small, oval cells embedded in a yellowish matrix. Thin reticular fibers spread and crisscross throughout the matrix. In the micrograph, the reticular fibers are thin, dark, and seem to travel between the many deeply stained cells.Figure 3. Reticular Tissue. This is a loose connective tissue made up of a network of reticular fibers that provides a supportive framework for soft organs. LM × 1600. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption] Reticular tissue is a mesh-like, supportive framework for soft organs such as lymphatic tissue, the spleen, and the liver (Figure 3). Reticular cells produce the reticular fibers that form the network onto which other cells attach. It derives its name from the Latin reticulus, which means “little net.”

Dense Connective Tissue

Dense connective tissue contains more collagen fibers than does loose connective tissue. As a consequence, it displays greater resistance to stretching. There are two major categories of dense connective tissue: regular and irregular. Dense regular connective tissue fibers are parallel to each other, enhancing tensile strength and resistance to stretching in the direction of the fiber orientations. Ligaments and tendons are made of dense regular connective tissue, but in ligaments not all fibers are parallel. Dense regular elastic tissue contains elastin fibers in addition to collagen fibers, which allows the ligament to return to its original length after stretching. The ligaments in the vocal folds and between the vertebrae in the vertebral column are elastic. [caption id="" align="alignright" width="500"]Part A shows a diagram of regular dense connective tissue alongside a micrograph. The tissue is composed of parallel, thread-like collagen fibers running vertically through the diagram. Between the vertical fibers, several dark, oval shaped fibroblast nuclei are visible. In the micrograph, the whitish collagen strands run horizontally. Several dark purple fibroblast nuclei are embedded in the lightly stained matrix. Part B shows a diagram of irregular dense connective tissue on the left and a micrograph on the right. In the diagram, the collagen fibers are arranged in bundles that curve and loop throughout the tissue. The fibers within a bundle run parallel to each other, but separate bundles crisscross throughout the tissue. Because of this, the irregular dense connective tissue appears less organized than the regular dense connective tissue. This is also evident in the micrograph, where the white collagen bundles radiate throughout the micrograph in all directions. The fibroblasts are visible as red stained cells with dark purple nuclei. Figure 4. Dense Connective Tissue. (a) Dense regular connective tissue consists of collagenous fibers packed into parallel bundles. (b) Dense irregular connective tissue consists of collagenous fibers interwoven into a mesh-like network. From top, LM × 1000, LM × 200. (Micrographs provided by the Regents of University of Michigan Medical School © 2012)[/caption] In dense irregular connective tissue, the direction of fibers is random. This arrangement gives the tissue greater strength in all directions and less strength in one particular direction. In some tissues, fibers crisscross and form a mesh. In other tissues, stretching in several directions is achieved by alternating layers where fibers run in the same orientation in each layer, and it is the layers themselves that are stacked at an angle. The dermis of the skin is an example of dense irregular connective tissue rich in collagen fibers. Dense irregular elastic tissues give arterial walls the strength and the ability to regain original shape after stretching (Figure 4).

Supportive Connective Tissues

Two major forms of supportive connective tissue, cartilage and bone, allow the body to maintain its posture and protect internal organs.

Cartilage

The distinctive appearance of cartilage is due to polysaccharides called chondroitin sulfates, which bind with ground substance proteins to form proteoglycans. Embedded within the cartilage matrix are chondrocytes, or cartilage cells, and the space they occupy are called lacunae (singular = lacuna). A layer of dense irregular connective tissue, the perichondrium, encapsulates the cartilage. Cartilaginous tissue is avascular, thus all nutrients need to diffuse through the matrix to reach the chondrocytes. This is a factor contributing to the very slow healing of cartilaginous tissues.

[caption id="" align="alignright" width="480"]Part A of this diagram is a drawing and a micrograph of hyaline cartilage. The cartilage contains chondrocytes encapsulated in lacunae. Several of the lacunae are joined into groups or small stacks and embedded in the surrounding matrix. The micrograph shows the lacunae as white rings surrounding the purple staining chondrocytes. Some occur as joined pairs while others are embedded singly within the pink staining matrix. Image B shows a diagram and a micrograph of fibrocartilage that contains many fine collagen fibers embedded in the matrix. The collagen fibers are roughly parallel to each but run through the matrix in a wavy fashion. There are also four round chondrocyte cells embedded within the matrix. In the micrograph, the matrix is shaded red and the collagen fibers are visible in white. The lacunae are clearly visible as a faint purple ring containing several dark purple chondrocytes. Part C shows a diagram and micrograph of elastic cartilage. In the diagram, fine elastic fibers are seen crisscrossing the matrix. Many of the elastic fibers branch off from each other, unlike the collagen fibers depicted in parts A and B. The lacunae are clearly visible as white rings containing stained chondrocytes. The fibers stain deeply in this micrograph and can been seen crisscrossing through the tissue. Figure 5. Types of Cartilage. Cartilage is a connective tissue consisting of collagenous fibers embedded in a firm matrix of chondroitin sulfates. (a) Hyaline cartilage provides support with some flexibility. The example is from dog tissue. (b) Fibrocartilage provides some compressibility and can absorb pressure. (c) Elastic cartilage provides firm but elastic support. From top, LM × 300, LM × 1200, LM × 1016. (Micrographs provided by the Regents of University of Michigan Medical School © 2012)[/caption] The three main types of cartilage tissue are hyaline cartilage, fibrocartilage, and elastic cartilage (Figure 5). Hyaline cartilage, the most common type of cartilage in the body, consists of short and dispersed collagen fibers and contains large amounts of proteoglycans. Under the microscope, tissue samples appear clear. The surface of hyaline cartilage is smooth. Both strong and flexible, it is found in the rib cage and nose and covers bones where they meet to form moveable joints. It makes up a template of the embryonic skeleton before bone formation. A plate of hyaline cartilage at the ends of bone allows continued growth until adulthood. Fibrocartilage is tough because it has thick bundles of collagen fibers dispersed through its matrix. The knee and jaw joints and the the intervertebral discs are examples of fibrocartilage. Elastic cartilage contains elastic fibers as well as collagen and proteoglycans. This tissue gives rigid support as well as elasticity. Tug gently at your ear lobes, and notice that the lobes return to their initial shape. The external ear contains elastic cartilage.

Bone

Bone is the hardest connective tissue. It provides protection to internal organs and supports the body. Bone’s rigid extracellular matrix contains mostly collagen fibers embedded in a mineralized ground substance containing hydroxyapatite, a form of calcium phosphate. Both components of the matrix, organic and inorganic, contribute to the unusual properties of bone. Without collagen, bones would be brittle and shatter easily. Without mineral crystals, bones would flex and provide little support. Osteocytes, bone cells like chondrocytes, are located within lacunae. The histology of transverse tissue from long bone shows a typical arrangement of osteocytes in concentric circles around a central canal. Bone is a highly vascularized tissue. Unlike cartilage, bone tissue can recover from injuries in a relatively short time. Cancellous bone ("trabecular bone" or "spongy bone") looks like a sponge under the microscope and contains empty spaces between trabeculae, or arches of bone proper. It is lighter than compact bone and found in the interior of some bones and at the end of long bones. Compact bone is solid and has greater structural strength.

Fluid Connective Tissue

Blood and lymph are fluid connective tissues. Cells circulate in a liquid extracellular matrix. The formed elements circulating in blood are all derived from hematopoietic stem cells located in bone marrow (Figure 6). Erythrocytes, red blood cells, transport oxygen and some carbon dioxide. Leukocytes, white blood cells, are responsible for defending against potentially harmful microorganisms or molecules. Platelets are cell fragments involved in blood clotting. Some white blood cells have the ability to cross the endothelial layer that lines blood vessels and enter adjacent tissues. Nutrients, salts, and wastes are dissolved in the liquid matrix and transported through the body.

[caption id="" align="alignright" width="420"]This micrograph of a blood smear shows a group of red blood cells and a single white blood cell. The red cells are small discs which have a slight depression at their centers with no nuclei present. The white blood cell is larger and more darkly stained and has a large, prominent nucleus that is also darkly stained. Figure 6. Blood: A Fluid Connective Tissue. Blood is a fluid connective tissue containing erythrocytes and various types of leukocytes that circulate in a liquid extracellular matrix. LM × 1600. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption] Lymph contains a liquid matrix and white blood cells. Lymphatic capillaries are extremely permeable, allowing larger molecules and excess fluid from interstitial spaces to enter the lymphatic vessels. Lymph drains into blood vessels, delivering molecules to the blood that could not otherwise directly enter the bloodstream. In this way, specialized lymphatic capillaries transport absorbed fats away from the intestine and deliver these molecules to the blood.

]]> 1999 1977 4 0 4.5 Nervous Tissue Mediates Perception and Response https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2004 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2004

Learning Objectives

By the end of this section, you will be able to:
  • Identify the classes of cells that make up nervous tissue
  • Discuss how nervous tissue mediates perception and response
[caption id="" align="alignright" width="480"]This figure shows a diagram of a neuron and a micrograph showing two neuron cells. The body of the neuron contains a single, purple nucleus. The cell is irregularly shaped, having many projections emerging from its surface. Six sets of dendrites project from the top, right, and bottom edges of the cell. The dendrites are yellow and branch many times after leaving the cell, taking on the appearance of tiny trees. The axon projects from the left edge of the cell. The axon is a long cable like structure that branches into several finger like projections at its end. This is where the neuron makes contact with other cells. A label also notes that the area where the axon emerges from the cell body contains microfibrils and microtubules. The micrograph is considerably less magnified than the diagram. The neurons stain darkly and their nuclei are clearly visible. Their irregular cell body is also visible, along with the beginning of the axons. Figure 1. The Neuron. The cell body of a neuron, also called the soma, contains the nucleus and mitochondria. The dendrites transfer the nerve impulse to the soma. The axon carries the action potential away to another excitable cell. LM × 1600. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Nervous tissue is characterized as being excitable and capable of sending and receiving electrochemical signals that provide the body with information. Two main classes of cells make up nervous tissue: the neuron and neuroglia (Figure 1). Neurons propagate information via electrochemical impulses, called action potentials, which are biochemically linked to the release of chemical signals. Neuroglia play an essential role in supporting neurons and modulating their information propagation.

 

Neurons display distinctive morphology, well suited to their role as conducting cells, with three main parts. The cell body includes most of the cytoplasm, the organelles, and the nucleus. Dendrites branch off the cell body and appear as thin extensions. A long “tail,” the axon, extends from the neuron body and can be wrapped in an insulating layer known as myelin, which is formed by accessory cells. The synapse is the gap between nerve cells, or between a nerve cell and its target, for example, a muscle or a gland, across which the impulse is transmitted by chemical compounds known as neurotransmitters. Neurons categorized as multipolar neurons have several dendrites and a single prominent axon. Bipolar neurons possess a single dendrite and axon with the cell body, while unipolar neurons have only a single process extending out from the cell body, which divides into a functional dendrite and into a functional axon. When a neuron is sufficiently stimulated, it generates an action potential that propagates down the axon towards the synapse. If enough neurotransmitters are released at the synapse to stimulate the next neuron or target, a response is generated.

[caption id="" align="alignleft" width="550"]Part A of this diagram shows various types of nerve cells. The largest cell is a neuron. The central body of the neuron contains a single nucleus. Six sets of dendrites project from the top, left and right, edges of the neuron. The dendrites are yellow and branch many times after leaving the cell, taking on the appearance of tiny trees. The axon projects from the bottom edge of the cell and is covered with purple sheaths labeled the myelin sheath. The sheath is not continuous, but instead is a series of equally spaced segments along the axon. Another cell, called an oligodendrocyte, is spider like in appearance, with its leg-like projections each connecting to a segment of the neuron’s myelin sheath. Above the neuron are three astrocytes. They are much smaller than the neuron and have no axons, and are also irregularly shaped cells with many dendrites projecting from the central body. Finally, a microglial cell is shown above the neuron. It is the smallest of the cells in this figure and is an elongated cell with many fine, tentacle-like projections. The projections are concentrated at the two ends of the cell, with the middle area lacking any projections. The micrograph of the neural tissue shows that this tissue is very heterogenous, with both large and small cells embedded in the matrix. Much of the space between the cells is occupied by threadlike nerve fibers. Figure 2. Nervous Tissue. Nervous tissue is made up of neurons and neuroglia. The cells of nervous tissue are specialized to transmit and receive impulses. LM × 872. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

The second class of neural cells comprises the neuroglia or glial cells, which have been characterized as having a simple support role. The word “glia” comes from the Greek word for glue. Recent research is shedding light on the more complex role of neuroglia in the function of the brain and nervous system. Astrocyte cells, named for their distinctive star shape, are abundant in the central nervous system. The astrocytes have many functions, including regulation of ion concentration in the intercellular space, uptake and/or breakdown of some neurotransmitters, and formation of the blood-brain barrier, the membrane that separates the circulatory system from the brain. Microglia protect the nervous system against infection but are not nervous tissue because they are related to macrophages. Oligodendrocyte cells produce myelin in the central nervous system (brain and spinal cord) while the Schwann cell produces myelin in the peripheral nervous system (Figure 2).

]]> 2004 1977 6 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2006 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2006 [caption id="" align="alignright" width="600"]Photo A shows a person getting a tattoo on the foot. Photo B shows a woman getting her eyebrows groomed. Photo C shows a person’s ear, with five earrings in the upper part and one in the lobe. Photo D shows a man and a boy shaving their faces. Figure 1. Your skin is a vital part of your life and appearance (a–d). Some people choose to embellish it with tattoos (a), makeup (b), and even piercings (c). (credit a: Steve Teo; credit b: “spaceodissey”/flickr; credit c: Mark/flickr; credit d: Lisa Schaffer)[/caption]

Chapter Objectives

After studying the chapter, you will be able to:
  • Describe the integumentary system and the role it plays in homeostasis
  • Describe the layers of the skin and the functions of each layer
  • Describe the accessory structures of the skin and the functions of each
  • Describe the changes that occur in the integumentary system during the aging process
  • Discuss several common diseases, disorders, and injuries that affect the integumentary system
  • Explain treatments for some common diseases, disorders, and injuries of the integumentary system
What do you think when you look at your skin in the mirror? Do you think about covering it with makeup, adding a tattoo, or maybe a body piercing? Or do you think about the fact that the skin belongs to one of the body’s most essential and dynamic systems: the integumentary system? The integumentary system refers to the skin and its accessory structures, and it is responsible for much more than simply lending to your outward appearance. In the adult human body, the skin makes up about 16 percent of body weight and covers an area of 1.5 to 2 m2. In fact, the skin and accessory structures are the largest organ system in the human body. As such, the skin protects your inner organs and it is in need of daily care and protection to maintain its health. This chapter will introduce the structure and functions of the integumentary system, as well as some of the diseases, disorders, and injuries that can affect this system.]]> 2006 2005 1 0 5.3 Functions of the Integumentary System https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2024 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2024

Learning Objectives

By the end of this section, you will be able to:
  • Describe the different functions of the skin and the structures that enable them
  • Explain how the skin helps maintain body temperature

The skin and accessory structures perform a variety of essential functions, such as protecting the body from invasion by microorganisms, chemicals, and other environmental factors; preventing dehydration; acting as a sensory organ; modulating body temperature and electrolyte balance; and synthesizing vitamin D. The underlying hypodermis has important roles in storing fats, forming a “cushion” over underlying structures, and providing insulation from cold temperatures.

 

Protection

The skin protects the rest of the body from the basic elements of nature such as wind, water, and UV sunlight. It acts as a protective barrier against water loss, due to the presence of layers of keratin and glycolipids in the stratum corneum. It also is the first line of defense against abrasive activity due to contact with grit, microbes, or harmful chemicals. Sweat excreted from sweat glands deters microbes from over-colonizing the skin surface by generating dermicidin, which has antibiotic properties.

 
[caption id="" align="alignright" width="380"]This micrograph shows a skin cross section at low magnification. The Meissner’s corpuscle is a large, oval-shaped structure located in the papillary layer of the dermis, under the lowest deepest layer of the epidermis. The corpuscle contains a dark staining oval within the outer, light staining oval. Several horizontal bars are arranged vertically within the inner oval. Also, several cells with dark purple nuclei can be seen scattered throughout the corpuscle. Figure 1. Light Micrograph of a Meissner Corpuscle. In this micrograph of a skin cross-section, you can see a Meissner corpuscle (arrow), a type of touch receptor located in a dermal papilla adjacent to the basement membrane and stratum basale of the overlying epidermis. LM × 100. (credit: “Wbensmith”/Wikimedia Commons)[/caption]

Sensory Function

The fact that you can feel an ant crawling on your skin, allowing you to flick it off before it bites, is because the skin, and especially the hairs projecting from hair follicles in the skin, can sense changes in the environment. The hair root plexus surrounding the base of the hair follicle senses a disturbance, and then transmits the information to the central nervous system (brain and spinal cord), which can then respond by activating the skeletal muscles of your eyes to see the ant and the skeletal muscles of the body to act against the ant.

The skin acts as a sense organ because the epidermis, dermis, and the hypodermis contain specialized sensory nerve structures that detect touch, surface temperature, and pain. These receptors are more concentrated on the tips of the fingers, which are most sensitive to touch, especially the Meissner corpuscle (tactile corpuscle) (Figure 1), which responds to light touch, and the Pacinian corpuscle (lamellated corpuscle), which responds to vibration. Merkel cells, seen scattered in the stratum basale, are also touch receptors. In addition to these specialized receptors, there are sensory nerves connected to each hair follicle, pain and temperature receptors scattered throughout the skin, and motor nerves innervate the arrector pili muscles and glands. This rich innervation helps us sense our environment and react accordingly.

[caption id="" align="alignleft" width="550"]Part A is a photo of a man skiing with several snow-covered trees in the background. Part B is a diagram with a right and left half. The left half is titled “ Heat is retained by the body,” while the right half is titled “Heat loss through radiation and convection.” Both show blood flowing from an artery through three capillary beds within the skin. The beds are arranged vertically, with the topmost bed located along the boundary of the dermis and epidermis. The bottommost bed is located deep in the hypodermis. The middle bed is evenly spaced between the topmost and bottommost beds. In each bed, oxygenated blood (red) enters the bed on the left and deoxygenated blood (blue) leaves the bed on the right. The left diagram shows a picture of snowflakes above the capillary beds, indicating that the weather is cold. Blood is only flowing through the deepest of the three capillary beds, as the upper beds are closed off to reduce heat loss from the outer layers of the skin. The right diagram shows a picture of the sun above the capillary beds, indicating that the weather is hot. Blood is flowing through all three capillary beds, allowing heat to radiate out of the blood, increasing heat loss. Part C is a photo of a man running through a forested trail on a summer day. Figure 2. Thermoregulation. During strenuous physical activities, such as skiing (a) or running (c), the dermal blood vessels dilate and sweat secretion increases (b). These mechanisms prevent the body from overheating. In contrast, the dermal blood vessels constrict to minimize heat loss in response to low temperatures (b). (credit a: “Trysil”/flickr; credit c: Ralph Daily)[/caption]

Thermoregulation

The integumentary system helps regulate body temperature through its tight association with the sympathetic nervous system, the division of the nervous system involved in our fight-or-flight responses. The sympathetic nervous system is continuously monitoring body temperature and initiating appropriate motor responses. Recall that sweat glands, accessory structures to the skin, secrete water, salt, and other substances to cool the body when it becomes warm. Even when the body does not appear to be noticeably sweating, approximately 500 mL of sweat (insensible perspiration) are secreted a day. If the body becomes excessively warm due to high temperatures, vigorous activity (Figure 2ac), or a combination of the two, sweat glands will be stimulated by the sympathetic nervous system to produce large amounts of sweat, as much as 0.7 to 1.5 L per hour for an active person. When the sweat evaporates from the skin surface, the body is cooled as body heat is dissipated.

In addition to sweating, arterioles in the dermis dilate so that excess heat carried by the blood can dissipate through the skin and into the surrounding environment (Figure 2b). This accounts for the skin redness that many people experience when exercising.

When body temperatures drop, the arterioles constrict to minimize heat loss, particularly in the ends of the digits and tip of the nose. This reduced circulation can result in the skin taking on a whitish hue. Although the temperature of the skin drops as a result, passive heat loss is prevented, and internal organs and structures remain warm. If the temperature of the skin drops too much (such as environmental temperatures below freezing), the conservation of body core heat can result in the skin actually freezing, a condition called frostbite.

 

Vitamin D Synthesis

The epidermal layer of human skin synthesizes vitamin D when exposed to UV radiation. In the presence of sunlight, a form of vitamin D3 called cholecalciferol is synthesized from a derivative of the steroid cholesterol in the skin. The liver converts cholecalciferol to calcidiol, which is then converted to calcitriol (the active chemical form of the vitamin) in the kidneys. Vitamin D is essential for normal absorption of calcium and phosphorous, which are required for healthy bones. The absence of sun exposure can lead to a lack of vitamin D in the body, leading to a condition called rickets, a painful condition in children where the bones are misshapen due to a lack of calcium, causing bowleggedness. Elderly individuals who suffer from vitamin D deficiency can develop a condition called osteomalacia, a softening of the bones. In present day society, vitamin D is added as a supplement to many foods, including milk and orange juice, compensating for the need for sun exposure.

In addition to its essential role in bone health, vitamin D is essential for general immunity against bacterial, viral, and fungal infections. Recent studies are also finding a link between insufficient vitamin D and cancer.

]]> 2024 2005 4 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2027 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2027 [caption id="" align="aligncenter" width="600"]This photo shows a boy looking at a museum exhibit that contains two fossilized crocodile skeletons embedded within a large boulder. The skull, spine and forelimbs of one of the crocodiles are visible. Figure 1. Child Looking at Bones. Bone is a living tissue. Unlike the bones of a fossil made inert by a process of mineralization, a child’s bones will continue to grow and develop while contributing to the support and function of other body systems. (credit: James Emery)[/caption]

Chapter Objectives

After studying this chapter, you will be able to:
  • List and describe the functions of bones
  • Describe the classes of bones
  • Discuss the process of bone formation and development
  • Explain how bone repairs itself after a fracture
  • Discuss the effect of exercise, nutrition, and hormones on bone tissue
  • Describe how an imbalance of calcium can affect bone tissue
Bones make good fossils. While the soft tissue of a once living organism will decay and fall away over time, bone tissue will, under the right conditions, undergo a process of mineralization, effectively turning the bone to stone. A well-preserved fossil skeleton can give us a good sense of the size and shape of an organism, just as your skeleton helps to define your size and shape. Unlike a fossil skeleton, however, your skeleton is a structure of living tissue that grows, repairs, and renews itself. The bones within it are dynamic and complex organs that serve a number of important functions, including some necessary to maintain homeostasis.]]> 2027 2025 1 0 6.1 The Functions of the Skeletal System https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2031 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2031

Learning Objectives

By the end of this section, you will be able to:
  • Define bone, cartilage, and the skeletal system
  • List and describe the functions of the skeletal system

Bone, or osseous tissue, is a hard, dense connective tissue that forms most of the adult skeleton, the support structure of the body. In the areas of the skeleton where bones move (for example, the ribcage and joints), cartilage, a semi-rigid form of connective tissue, provides flexibility and smooth surfaces for movement. The skeletal system is the body system composed of bones and cartilage and performs the following critical functions for the human body:

  • supports the body
  • facilitates movement
  • protects internal organs
  • produces blood cells
  • stores and releases minerals and fat
[caption id="" align="alignleft" width="290"]This photo shows a man exercising on a leg press machine at a gym. Figure 1. Bones Support Movement. Bones act as levers when muscles span a joint and contract. (credit: Benjamin J. DeLong)[/caption] [caption id="" align="alignright" width="250"]This illustration shows how the cranium protects and surrounds the brain. Only the outline of the cranium is visible, which is made transparent to show how the brain sits in the skull. There is a small amount of space between the brain and the cranium but the top and sides of the brain are completely protected by the cranial bones. The bottom of the brain extends below the cranial bones, with the base of the cerebellum seated just above the roof of the mouth. The medulla extends to the bottom of the skull where it meets with the spinal cord. Figure 2. Bones Protect Brain. The cranium completely surrounds and protects the brain from non-traumatic injury.[/caption]

Support, Movement, and Protection

The most apparent functions of the skeletal system are the gross functions—those visible by observation. Simply by looking at a person, you can see how the bones support, facilitate movement, and protect the human body.

Just as the steel beams of a building provide a scaffold to support its weight, the bones and cartilage of your skeletal system compose the scaffold that supports the rest of your body. Without the skeletal system, you would be a limp mass of organs, muscle, and skin.

Bones also facilitate movement by serving as points of attachment for your muscles. While some bones only serve as a support for the muscles, others also transmit the forces produced when your muscles contract. From a mechanical point of view, bones act as levers and joints serve as fulcrums (Figure 1). Unless a muscle spans a joint and contracts, a bone is not going to move. For information on the interaction of the skeletal and muscular systems, that is, the musculoskeletal system, seek additional content.
Bones also protect internal organs from injury by covering or surrounding them. For example, your ribs protect your lungs and heart, the bones of your vertebral column (spine) protect your spinal cord, and the bones of your cranium (skull) protect your brain (Figure 2).

Mineral Storage, Energy Storage, and Hematopoiesis

[caption id="" align="alignright" width="380"]This photo shows the head of the femur detached from the rest of the bone. The compact bone at the surface of the head has been removed to show the spongy bone beneath. Rather than being solid, like the compact bone, the spongy bone is mesh like with many open spaces, giving it the appearance of a sponge. A circle of yellow marrow is located at the exact center of the spongy bone. The red marrow surrounds the yellow marrow, occupying most of the interior space of the head. Figure 4. Head of Femur Showing Red and Yellow Marrow. The head of the femur contains both yellow and red marrow. Yellow marrow stores fat. Red marrow is responsible for hematopoiesis. (credit: modification of work by “stevenfruitsmaak”/Wikimedia Commons)[/caption] On a metabolic level, bone tissue performs several critical functions. For one, the bone matrix acts as a reservoir for a number of minerals important to the functioning of the body, especially calcium, and phosphorus. These minerals, incorporated into bone tissue, can be released back into the bloodstream to maintain levels needed to support physiological processes. Calcium ions, for example, are essential for muscle contractions and controlling the flow of other ions involved in the transmission of nerve impulses.

Bone also serves as a site for fat storage and blood cell production. The softer connective tissue that fills the interior of most bone is referred to as bone marrow (Figure 4). There are two types of bone marrow: yellow marrow and red marrow. Yellow marrow contains adipose tissue; the triglycerides stored in the adipocytes of the tissue can serve as a source of energy. Red marrow is where hematopoiesis—the production of blood cells—takes place. Red blood cells, white blood cells, and platelets are all produced in the red marrow.

]]> 2031 2025 2 0 6.2 Bone Classification https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2033 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2033

Learning Objectives

By the end of this section, you will be able to:
  • Classify bones according to their shapes
  • Describe the function of each category of bones

The 206 bones that compose the adult skeleton are divided into five categories based on their shapes (Figure 1). Their shapes and their functions are related such that each categorical shape of bone has a distinct function.

[caption id="" align="aligncenter" width="600"]This illustration shows an anterior view of a human skeleton with call outs of five bones. The first call out is the sternum, or breast bone, which lies along the midline of the thorax. The sternum is the bone to which the ribs connect at the front of the body. It is classified as a flat bone and appears somewhat like a tie, with an enlarged upper section and a thin, tapering, lower section. The next callout is the right femur, which is the thigh bone. The inferior end of the femur is broad where it connects to the knee while the superior edge is ball-shaped where it attaches to the hip socket. The femur is an example of a long bone. The next callout is of the patella or kneecap. It is a small, wedge-shaped bone that sits on the anterior side of the knee. The kneecap is an example of a sesamoid bone. The next callout is a dorsal view of the right foot. The lateral, intermediate and medial cuneiform bones are small, square-shaped bones of the top of the foot. These bones lie between the proximal edge of the toe bones and the inferior edge of the shin bones. The lateral cuneiform is proximal to the fourth toe while the medial cuneiform is proximal to the great toe. The intermediate cuneiform lies between the lateral and medial cuneiform. These bones are examples of short bones. The fifth callout shows a superior view of one of the lumbar vertebrae. The vertebra has a kidney-shaped body connected to a triangle of bone that projects above the body of the vertebra. Two spines project off of the triangle at approximately 45 degree angles. The vertebrae are examples of irregular bones. Figure 1. Classifications of Bones. Bones are classified according to their shape.[/caption]

Long Bones

A long bone is one that is cylindrical in shape, being longer than it is wide. Keep in mind, however, that the term describes the shape of a bone, not its size. Long bones are found in the arms (humerus, ulna, radius) and legs (femur, tibia, fibula), as well as in the fingers (metacarpals, phalanges) and toes (metatarsals, phalanges). Long bones function as levers; they move when muscles contract.

Short Bones

A short bone is one that is cube-like in shape, being approximately equal in length, width, and thickness. The only short bones in the human skeleton are in the carpals of the wrists and the tarsals of the ankles. Short bones provide stability and support as well as some limited motion.

Flat Bones

The term “flat bone” is somewhat of a misnomer because, although a flat bone is typically thin, it is also often curved. Examples include the cranial (skull) bones, the scapulae (shoulder blades), the sternum (breastbone), and the ribs. Flat bones serve as points of attachment for muscles and often protect internal organs.

Irregular Bones

An irregular bone is one that does not have any easily characterized shape and therefore does not fit any other classification. These bones tend to have more complex shapes, like the vertebrae that support the spinal cord and protect it from compressive forces. Many facial bones, particularly the ones containing sinuses, are classified as irregular bones.

Sesamoid Bones

A sesamoid bone is a small, round bone that, as the name suggests, is shaped like a sesame seed. These bones form in tendons (the sheaths of tissue that connect bones to muscles) where a great deal of pressure is generated in a joint. The sesamoid bones protect tendons by helping them overcome compressive forces. Sesamoid bones vary in number and placement from person to person but are typically found in tendons associated with the feet, hands, and knees. The patellae (singular = patella) are the only sesamoid bones found in common with every person. Table 1 reviews bone classifications with their associated features, functions, and examples.

Bone Classifications (Table 1)
Bone classification Features Function(s) Examples
Long Cylinder-like shape, longer than it is wide Leverage Femur, tibia, fibula, metatarsals, humerus, ulna, radius, metacarpals, phalanges
Short Cube-like shape, approximately equal in length, width, and thickness Provide stability, support, while allowing for some motion Carpals, tarsals
Flat Thin and curved Points of attachment for muscles; protectors of internal organs Sternum, ribs, scapulae, cranial bones
Irregular Complex shape Protect internal organs Vertebrae, facial bones
Sesamoid Small and round; embedded in tendons Protect tendons from compressive forces Patellae
]]> 2033 2025 3 0 6.4 Bone Formation and Development https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2048 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2048

Learning Objectives

By the end of this section, you will be able to:
  • Explain the function of cartilage
  • List the steps of intramembranous ossification
  • List the steps of endochondral ossification
  • Explain the growth activity at the epiphyseal plate
  • Compare and contrast the processes of modeling and remodeling

In the early stages of embryonic development, the embryo’s skeleton consists of fibrous membranes and hyaline cartilage. By the sixth or seventh week of embryonic life, the actual process of bone development, ossification (osteogenesis), begins. There are two osteogenic pathways—intramembranous ossification and endochondral ossification—but bone is the same regardless of the pathway that produces it.

Cartilage Templates

Bone is a replacement tissue; that is, it uses a model tissue on which to lay down its mineral matrix. For skeletal development, the most common template is cartilage. During fetal development, a framework is laid down that determines where bones will form. This framework is a flexible, semi-solid matrix produced by chondroblasts and consists of hyaluronic acid, chondroitin sulfate, collagen fibers, and water. As the matrix surrounds and isolates chondroblasts, they are called chondrocytes. Unlike most connective tissues, cartilage is avascular, meaning that it has no blood vessels supplying nutrients and removing metabolic wastes. All of these functions are carried on by diffusion through the matrix. This is why damaged cartilage does not repair itself as readily as most tissues do.

Throughout fetal development and into childhood growth and development, bone forms on the cartilaginous matrix. By the time a fetus is born, most of the cartilage has been replaced with bone. Some additional cartilage will be replaced throughout childhood, and some cartilage remains in the adult skeleton.

[caption id="" align="alignright" width="550"]Image A shows seven osteoblasts, cells with small, finger like projections. They are surrounded by granules of osteoid. Both the cells and the osteoid are contained within a blue, circular, ossification center that is surrounded by a “socket” of dark, string-like collagen fibers and gray mesenchymal cells. The cells are generally amorphous, similar in appearance to an amoeba. In image B, the ossification center is no longer surrounded by a ring of osteoblasts. The osteoblasts have secreted bone into the ossification center, creating a new bone matrix. There are also five osteocytes embedded in the new bone matrix. The osteocytes are thin, oval-shaped cells with many fingerlike projections. Osteoid particles are still embedded in the bony matrix in image B. In image C, the ring of osteoblasts surrounding the ossification center has separated, forming an upper and lower layer of osteoblasts sandwiched between the two layers of mesenchyme cells. A label indicates that the mesenchyme cells and the surrounding collagen fibers form the periosteum. The osteoblasts secrete spongy bone into the space between the two osteoblast rows. Therefore, the accumulating spongy bone pushes the upper and lower rows of osteoblasts away from each other. In this image, most of the spongy bone has been secreted by the osteoblasts, as the trabeculae are visible. In addition, an artery has already broken through the periosteum and invaded the spongy bone. Image D looks similar to image C, except that the rows of osteoblasts are now secreting layers of compact bone between the spongy bone and the periosteum. The artery has now branched and spread throughout the spongy bone. A label indicates that the cavities between the trabeculae now contain red bone marrow. Figure 1. Intramembranous Ossification. Intramembranous ossification follows four steps. (a) Mesenchymal cells group into clusters, and ossification centers form. (b) Secreted osteoid traps osteoblasts, which then become osteocytes. (c) Trabecular matrix and periosteum form. (d) Compact bone develops superficial to the trabecular bone, and crowded blood vessels condense into red marrow.[/caption]

Intramembranous Ossification

During intramembranous ossification, compact and spongy bone develops directly from sheets of mesenchymal (undifferentiated) connective tissue. The flat bones of the face, most of the cranial bones, and the clavicles (collarbones) are formed via intramembranous ossification.

The process begins when mesenchymal cells in the embryonic skeleton gather together and begin to differentiate into specialized cells (Figure 1a). Some of these cells will differentiate into capillaries, while others will become osteogenic cells and then osteoblasts. Although they will ultimately be spread out by the formation of bone tissue, early osteoblasts appear in a cluster called an ossification center.

The osteoblasts secrete osteoid, uncalcified matrix, which calcifies (hardens) within a few days as mineral salts are deposited on it, thereby entrapping the osteoblasts within. Once entrapped, the osteoblasts become osteocytes (Figure 1b). As osteoblasts transform into osteocytes, osteogenic cells in the surrounding connective tissue differentiate into new osteoblasts.

Osteoid (unmineralized bone matrix) secreted around the capillaries results in a trabecular matrix, while osteoblasts on the surface of the spongy bone become the periosteum (Figure 1c). The periosteum then creates a protective layer of compact bone superficial to the trabecular bone. The trabecular bone crowds nearby blood vessels, which eventually condense into red marrow (Figure 1d).

Intramembranous ossification begins in utero during fetal development and continues on into adolescence. At birth, the skull and clavicles are not fully ossified nor are the sutures of the skull closed. This allows the skull and shoulders to deform during passage through the birth canal. The last bones to ossify via intramembranous ossification are the flat bones of the face, which reach their adult size at the end of the adolescent growth spurt.

[caption id="" align="alignleft" width="600"]Image A shows a small piece of hyaline cartilage that looks like a bone but without the characteristic enlarged ends. The hyaline cartilage is surrounded by a thin perichondrium. In image B, the hyaline cartilage has increased in size and the ends have begun to bulge outwards. A group of dark granules form at the center of the cartilage. This is labeled the calcified matrix, as opposed to the rest of the cartilage, which is uncalcified matrix. In image C, the hyaline cartilage has again increased in size and spongy bone has formed at the calcified matrix. This is now called the primary ossification center. A nutrient artery has invaded the ossification center and is growing through the cavities of the new spongy bone. In image D, the cartilage now looks like a bone, as it has greatly increased in size and each end has two bulges. Only the proximal half of the bone is shown in all of the remaining images. In image D, spongy bone has completely developed in the medullary cavity, which is surrounded, on both sides, by compact bone. Now, the calcified matrix is located at the border between the proximal metaphysis and the proximal epiphysis. The epiphysis is still composed of uncalcified matrix. In image E, arteries and veins have now invaded the epiphysis, forming a calcified matrix at its center. This is called a secondary ossification center. In image F, the interior of the epiphysis is now completely calcified into bone. The outer edge of the epiphysis remains as cartilage, forming the articular cartilage at the joint. In addition, the border between the epiphysis and the metaphysis remains uncalcified, forming the epiphyseal plate. Figure 2. Endochondral Ossification. Endochondral ossification follows five steps. (a) Mesenchymal cells differentiate into chondrocytes. (b) The cartilage model of the future bony skeleton and the perichondrium form. (c) Capillaries penetrate cartilage. Perichondrium transforms into periosteum. Periosteal collar develops. Primary ossification center develops. (d) Cartilage and chondrocytes continue to grow at ends of the bone. (e) Secondary ossification centers develop. (f) Cartilage remains at epiphyseal (growth) plate and at joint surface as articular cartilage.[/caption]

Endochondral Ossification

In endochondral ossification, bone develops by replacing hyaline cartilage. Cartilage does not become bone. Instead, cartilage serves as a template to be completely replaced by new bone. Endochondral ossification takes much longer than intramembranous ossification. Bones at the base of the skull and long bones form via endochondral ossification.

In a long bone, for example, at about 6 to 8 weeks after conception, some of the mesenchymal cells differentiate into chondrocytes (cartilage cells) that form the cartilaginous skeletal precursor of the bones (Figure 2a). Soon after, the perichondrium, a membrane that covers the cartilage, appears Figure 2b).

As more matrix is produced, the chondrocytes in the center of the cartilaginous model grow in size. As the matrix calcifies, nutrients can no longer reach the chondrocytes. This results in their death and the disintegration of the surrounding cartilage. Blood vessels invade the resulting spaces, not only enlarging the cavities but also carrying osteogenic cells with them, many of which will become osteoblasts. These enlarging spaces eventually combine to become the medullary cavity.

As the cartilage grows, capillaries penetrate it. This penetration initiates the transformation of the perichondrium into the bone-producing periosteum. Here, the osteoblasts form a periosteal collar of compact bone around the cartilage of the diaphysis. By the second or third month of fetal life, bone cell development and ossification ramps up and creates the primary ossification center, a region deep in the periosteal collar where ossification begins (Figure 2c).

While these deep changes are occurring, chondrocytes and cartilage continue to grow at the ends of the bone (the future epiphyses), which increases the bone’s length at the same time bone is replacing cartilage in the diaphyses. By the time the fetal skeleton is fully formed, cartilage only remains at the joint surface as articular cartilage and between the diaphysis and epiphysis as the epiphyseal plate, the latter of which is responsible for the longitudinal growth of bones. After birth, this same sequence of events (matrix mineralization, death of chondrocytes, invasion of blood vessels from the periosteum, and seeding with osteogenic cells that become osteoblasts) occurs in the epiphyseal regions, and each of these centers of activity is referred to as a secondary ossification center (Figure 2e).

How Bones Grow in Length

The epiphyseal plate is the area of growth in a long bone. It is a layer of hyaline cartilage where ossification occurs in immature bones. On the epiphyseal side of the epiphyseal plate, cartilage is formed. On the diaphyseal side, cartilage is ossified, and the diaphysis grows in length. The epiphyseal plate is composed of four zones of cells and activity (Figure 3). The reserve zone is the region closest to the epiphyseal end of the plate and contains small chondrocytes within the matrix. These chondrocytes do not participate in bone growth but secure the epiphyseal plate to the osseous tissue of the epiphysis.

[caption id="" align="alignright" width="350"]This illustration shows the zones bordering the epiphyseal plate of the epiphysis. The topmost layer of the epiphysis is the reserve zone, which is colored blue because it is made of cartilage. Two arteries are shown travelling through this zone to supply nutrients to the second zone: the proliferative zone. Here, five chondrocytes are undergoing mitosis. They continually divide, producing five long rows of chondrocytes. The next zone is the zone of maturation and hypertrophy. Here, lipids, glycogen and alkaline phosphatase accumulate, causing the cartilaginous matrix to calcify. This zone consists of five rows of ten chondrocytes which are increasing in size as one moves down a row. The next zone is the calcified matrix. Here, the chondrocytes have hardened and die as the matrix around them has calcified. The bottommost row is the zone of ossification. This zone is actually part of the metaphysis. Arteries from the metaphysis branch through the newly-formed trabeculae in this zone. The newly deposited bone tissue at the top of the zone of ossification is called the primary spongiosa. The older bone at the bottom of the zone of ossification is labeled the secondary spongiosa. Figure 3. Longitudinal Bone Growth. The epiphyseal plate is responsible for longitudinal bone growth.[/caption] The proliferative zone is the next layer toward the diaphysis and contains stacks of slightly larger chondrocytes. It makes new chondrocytes (via mitosis) to replace those that die at the diaphyseal end of the plate. Chondrocytes in the next layer, the zone of maturation and hypertrophy, are older and larger than those in the proliferative zone. The more mature cells are situated closer to the diaphyseal end of the plate. The longitudinal growth of bone is a result of cellular division in the proliferative zone and the maturation of cells in the zone of maturation and hypertrophy. 

Most of the chondrocytes in the zone of calcified matrix, the zone closest to the diaphysis, are dead because the matrix around them has calcified. Capillaries and osteoblasts from the diaphysis penetrate this zone, and the osteoblasts secrete bone tissue on the remaining calcified cartilage. Thus, the zone of calcified matrix connects the epiphyseal plate to the diaphysis. A bone grows in length when osseous tissue is added to the diaphysis.

Bones continue to grow in length until early adulthood. The rate of growth is controlled by hormones, which will be discussed later. When the chondrocytes in the epiphyseal plate cease their proliferation and bone replaces the cartilage, longitudinal growth stops. All that remains of the epiphyseal plate is the epiphyseal line (Figure 4).
[caption id="" align="alignleft" width="380"]This illustration shows anterior views of a right and left femur. The left femur possesses a growth plate at the border of its distal metaphysis and distal epiphysis. The proximal epiphysis has two growth plates. The first is located below the head of the femur while the second is located below the trochanter, which is the bump on the lateral side of the femur. The right femur has the same plates as the left femur. However, the left femur represents a mature long bone. Here, growth is completed and the epiphyseal plate has degraded to a thin, faint, epiphyseal line. Figure 4. Progression from Epiphyseal Plate to Epiphyseal Line. As a bone matures, the epiphyseal plate progresses to an epiphyseal line. (a) Epiphyseal plates are visible in a growing bone. (b) Epiphyseal lines are the remnants of epiphyseal plates in a mature bone.[/caption]

How Bones Grow in Diameter

While bones are increasing in length, they are also increasing in diameter; growth in diameter can continue even after longitudinal growth ceases. This is called appositional growth. Osteoclasts resorb old bone that lines the medullary cavity, while osteoblasts, via intramembranous ossification, produce new bone tissue beneath the periosteum. The erosion of old bone along the medullary cavity and the deposition of new bone beneath the periosteum not only increase the diameter of the diaphysis but also increase the diameter of the medullary cavity. This process is called modeling.

Bone Remodeling

The process in which matrix is resorbed on one surface of a bone and deposited on another is known as bone modeling. Modeling primarily takes place during a bone’s growth. However, in adult life, bone undergoes remodeling, in which resorption of old or damaged bone takes place on the same surface where osteoblasts lay new bone to replace that which is resorbed. Injury, exercise, and other activities lead to remodeling. Those influences are discussed later in the chapter, but even without injury or exercise, about 5 to 10 percent of the skeleton is remodeled annually just by destroying old bone and renewing it with fresh bone.

]]> 2048 2025 5 0 6.5 Fractures: Bone Repair https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2051 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2051

Learning Objectives

By the end of this section, you will be able to:
  • Differentiate among the different types of fractures
  • Describe the steps involved in bone repair

A fracture is a broken bone. It will heal whether or not a physician resets it in its anatomical position. If the bone is not reset correctly, the healing process will keep the bone in its deformed position.

When a broken bone is manipulated and set into its natural position without surgery, the procedure is called a closed reduction. Open reduction requires surgery to expose the fracture and reset the bone. While some fractures can be minor, others are quite severe and result in grave complications. For example, a fractured diaphysis of the femur has the potential to release fat globules into the bloodstream. These can become lodged in the capillary beds of the lungs, leading to respiratory distress and if not treated quickly, death.

Types of Fractures

Fractures are classified by their complexity, location, and other features (Figure 1). Table 4 outlines common types of fractures. Some fractures may be described using more than one term because it may have the features of more than one type (e.g., an open transverse fracture). [caption id="" align="aligncenter" width="395"]In this illustration, each type of fracture is shown on the right femur from an anterior view. In the closed fracture, the femur is broken in the middle of the shaft with the upper and lower halves of the bone completely separated. However, the two halves of the bones are still aligned in that the broken edges are still facing each other. In an open fracture, the femur is broken in the middle of the shaft with the upper and lower halves of the bone completely separated. Unlike the closed fracture, in the open fracture, the two bone halves are misaligned. The lower half is turned laterally and it has protruded through the skin of the thigh. The broken ends no longer line up with each other. In a transverse fracture, the bone has a crack entirely through its width, however, the broken ends are not separated. The crack is perpendicular to the long axis of the bone. Arrows indicate that this is usually caused by compression of the bone in a superior-inferior direction. A spiral fracture travels diagonally through the diameter of the bone. In a comminuted fracture, the bone has several connecting cracks at its middle. It is possible that the bone could splinter into several small pieces at the site of the comminuted fracture. In an impacted fracture, the crack zig zags throughout the width of the bone like a lightning bolt. An arrow indicates that these are usually caused by an impact that pushes the femur up into the body. A greenstick fracture is a small crack that does not extend through the entire width of the bone. The oblique fracture shown here is travelling diagonally through the shaft of the femur at about a thirty degree angle. Figure 1. Types of Fractures. Compare healthy bone with different types of fractures: (a) closed fracture, (b) open fracture, (c) transverse fracture, (d) spiral fracture, (e) comminuted fracture, (f) impacted fracture, (g) greenstick fracture, and (h) oblique fracture.[/caption]
Types of Fractures (Table 4)
Type of fracture Description
Transverse Occurs straight across the long axis of the bone
Oblique Occurs at an angle that is not 90 degrees
Spiral Bone segments are pulled apart as a result of a twisting motion
Comminuted Several breaks result in many small pieces between two large segments
Impacted One fragment is driven into the other, usually as a result of compression
Greenstick A partial fracture in which only one side of the bone is broken
Open (or compound) A fracture in which at least one end of the broken bone tears through the skin; carries a high risk of infection
Closed (or simple) A fracture in which the skin remains intact

Bone Repair

When a bone breaks, blood flows from any vessel torn by the fracture. These vessels could be in the periosteum, osteons, and/or medullary cavity. The blood begins to clot, and about six to eight hours after the fracture, the clotting blood has formed a fracture hematoma (Figure 2a). The disruption of blood flow to the bone results in the death of bone cells around the fracture.

[caption id="" align="aligncenter" width="485"]This illustration shows a left to right progression of bone repair. The break is shown in the leftmost image, where the femur has an oblique, closed fracture in the middle of its shaft. The next image magnifies the break, showing that blood has filled the area between the broken bones. Blood has also filled in around the lateral and medial sides of the break. The influx of blood causes the broken area to swell, creating a hematoma. In the next image, the hematoma has been replaced with an external callus between the two broken ends. Within the internal callus, the blood vessels have reconnected and some spongy bone has regenerated in the gap between the two bone halves. In the next image, spongy bone has completely regenerated, connecting the two broken ends, referred to as the bony callus. The external callus still remains on the lateral and medial sides of the break, as the compact bone has not yet regenerated. In the final image, the compact bone has fully regenerated, encapsulating the bony callus and completely reconnecting the two bone halves. The bone has a slight bulge at the location of the healed fracture, which is clearly shown in the final image, which shows a zoomed out image of the completely healed femur. Figure 2. Stages in Fracture Repair. The healing of a bone fracture follows a series of progressive steps: (a) A fracture hematoma forms. (b) Internal and external calli form. (c) Cartilage of the calli is replaced by trabecular bone. (d) Remodeling occurs.[/caption]

Within about 48 hours after the fracture, chondrocytes from the endosteum have created an internal callus (plural = calli) by secreting a fibrocartilaginous matrix between the two ends of the broken bone, while the periosteal chondrocytes and osteoblasts create an external callus of hyaline cartilage and bone, respectively, around the outside of the break (Figure 2b). This stabilizes the fracture.

Over the next several weeks, osteoclasts resorb the dead bone; osteogenic cells become active, divide, and differentiate into osteoblasts. The cartilage in the calli is replaced by trabecular bone via endochondral ossification (Figure 2c).

Eventually, the internal and external calli unite, compact bone replaces spongy bone at the outer margins of the fracture, and healing is complete. A slight swelling may remain on the outer surface of the bone, but quite often, that region undergoes remodeling (Figure 2d), and no external evidence of the fracture remains...
]]> 2051 2025 6 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2054 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2054

Chapter Objectives

After this chapter, you will be able to:
  • Describe the functions of the skeletal system and define its two major subdivisions
  • Identify the bones and bony structures of the skull, the cranial suture lines, the cranial fossae, and the openings in the skull
  • Discuss the vertebral column and regional variations in its bony components and curvatures
  • Describe the components of the thoracic cage
  • Discuss the embryonic development of the axial skeleton
[caption id="" align="aligncenter" width="250"]This image shows a side view of the human skull. The major parts of the cell are labeled. Figure 1. Lateral View of the Human Skull.[/caption] The skeletal system forms the rigid internal framework of the body. It consists of the bones, cartilages, and ligaments. Bones support the weight of the body, allow for body movements, and protect internal organs. Cartilage provides flexible strength and support for body structures such as the thoracic cage, the external ear, and the trachea and larynx. At joints of the body, cartilage can also unite adjacent bones or provide cushioning between them. Ligaments are the strong connective tissue bands that hold the bones at a moveable joint together and serve to prevent excessive movements of the joint that would result in injury. Providing movement of the skeleton are the muscles of the body, which are firmly attached to the skeleton via connective tissue structures called tendons. As muscles contract, they pull on the bones to produce movements of the body. Thus, without a skeleton, you would not be able to stand, run, or even feed yourself! Each bone of the body serves a particular function, and therefore bones vary in size, shape, and strength based on these functions. For example, the bones of the lower back and lower limb are thick and strong to support your body weight. Similarly, the size of a bony landmark that serves as a muscle attachment site on an individual bone is related to the strength of this muscle. Muscles can apply very strong pulling forces to the bones of the skeleton. To resist these forces, bones have enlarged bony landmarks at sites where powerful muscles attach. This means that not only the size of a bone, but also its shape, is related to its function. For this reason, the identification of bony landmarks is important during your study of the skeletal system. Bones are also dynamic organs that can modify their strength and thickness in response to changes in muscle strength or body weight. Thus, muscle attachment sites on bones will thicken if you begin a workout program that increases muscle strength. Similarly, the walls of weight-bearing bones will thicken if you gain body weight or begin pounding the pavement as part of a new running regimen. In contrast, a reduction in muscle strength or body weight will cause bones to become thinner. This may happen during a prolonged hospital stay, following limb immobilization in a cast, or going into the weightlessness of outer space. Even a change in diet, such as eating only soft food due to the loss of teeth, will result in a noticeable decrease in the size and thickness of the jaw bones.]]> 2054 2052 1 0 7.1 Divisions of the Skeletal System https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2056 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2056

Learning Objectives

By the end of this section, you will be able to:
  • Discuss the functions of the skeletal system
  • Distinguish between the axial skeleton and appendicular skeleton
  • Define the axial skeleton and its components
  • Define the appendicular skeleton and its components

The skeletal system includes all of the bones, cartilages, and ligaments of the body that support and give shape to the body and body structures. The skeleton consists of the bones of the body. For adults, there are 206 bones in the skeleton. Younger individuals have higher numbers of bones because some bones fuse together during childhood and adolescence to form an adult bone. The primary functions of the skeleton are to provide a rigid, internal structure that can support the weight of the body against the force of gravity, and to provide a structure upon which muscles can act to produce movements of the body. The lower portion of the skeleton is specialized for stability during walking or running. In contrast, the upper skeleton has greater mobility and ranges of motion, features that allow you to lift and carry objects or turn your head and trunk.

In addition to providing for support and movements of the body, the skeleton has protective and storage functions. It protects the internal organs, including the brain, spinal cord, heart, lungs, and pelvic organs. The bones of the skeleton serve as the primary storage site for important minerals such as calcium and phosphate. The bone marrow found within bones stores fat and houses the blood-cell producing tissue of the body.

The skeleton is subdivided into two major divisions—the axial and appendicular.

The Axial Skeleton

The skeleton is subdivided into two major divisions—the axial and appendicular. The axial skeleton forms the vertical, central axis of the body and includes all bones of the head, neck, chest, and back (Figure 1). It serves to protect the brain, spinal cord, heart, and lungs. It also serves as the attachment site for muscles that move the head, neck, and back, and for muscles that act across the shoulder and hip joints to move their corresponding limbs.

The axial skeleton of the adult consists of 80 bones, including the skull, the vertebral column, and the thoracic cage. The skull is formed by 22 bones. Also associated with the head are an additional seven bones, including the hyoid bone and the ear ossicles (three small bones found in each middle ear). The vertebral column consists of 24 bones, each called a vertebra, plus the sacrum and coccyx. The thoracic cage includes the 12 pairs of ribs, and the sternum, the flattened bone of the anterior chest.

[caption id="" align="aligncenter" width="500"]This diagram shows the human skeleton and identifies the major bones. The left panel shows the anterior view (from the front) and the right panel shows the posterior view (from the back). Figure 1. Axial and Appendicular Skeleton. The axial skeleton supports the head, neck, back, and chest and thus forms the vertical axis of the body. It consists of the skull, vertebral column (including the sacrum and coccyx), and the thoracic cage, formed by the ribs and sternum. The appendicular skeleton is made up of all bones of the upper and lower limbs.[/caption]

The Appendicular Skeleton

The appendicular skeleton includes all bones of the upper and lower limbs, plus the bones that attach each limb to the axial skeleton. There are 126 bones in the appendicular skeleton of an adult. The bones of the appendicular skeleton are covered in a separate chapter.

]]> 2056 2052 2 0 7.2 The Skull https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2065 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2065

Summary

The cranium (skull) is the skeletal structure of the head that supports the face and protects the brain. It is subdivided into the facial bones and the brain case, or cranial vault (Figure 1). The facial bones underlie the facial structures, form the nasal cavity, enclose the eyeballs, and support the teeth of the upper and lower jaws. The rounded brain case surrounds and protects the brain and houses the middle and inner ear structures.

In the adult, the skull consists of 22 individual bones, 21 of which are immobile and united into a single unit. The 22nd bone is the mandible (lower jaw), which is the only moveable bone of the skull.

[caption id="" align="aligncenter" width="380"]In this image, the lateral view of the human skull is shown and the brain case and facial bones are labeled. Figure 1. Parts of the Skull. The skull consists of the rounded brain case that houses the brain and the facial bones that form the upper and lower jaws, nose, orbits, and other facial structures.[/caption]

Anterior View of Skull

The anterior skull consists of the facial bones and provides the bony support for the eyes and structures of the face. This view of the skull is dominated by the openings of the orbits and the nasal cavity. Also seen are the upper and lower jaws, with their respective teeth (Figure 2).

The orbit is the bony socket that houses the eyeball and muscles that move the eyeball or open the upper eyelid. The upper margin of the anterior orbit is the supraorbital margin. Located near the midpoint of the supraorbital margin is a small opening called the supraorbital foramen. This provides for passage of a sensory nerve to the skin of the forehead. Below the orbit is the infraorbital foramen, which is the point of emergence for a sensory nerve that supplies the anterior face below the orbit.

[caption id="" align="aligncenter" width="480"]This image shows the anterior view (from the front) of the human skull. The major bones on the skull are labeled. Figure 2. Anterior View of Skull. An anterior view of the skull shows the bones that form the forehead, orbits (eye sockets), nasal cavity, nasal septum, and upper and lower jaws.[/caption]

Inside the nasal area of the skull, the nasal cavity is divided into halves by the nasal septum. The upper portion of the nasal septum is formed by the perpendicular plate of the ethmoid bone and the lower portion is the vomer bone. Each side of the nasal cavity is triangular in shape, with a broad inferior space that narrows superiorly. When looking into the nasal cavity from the front of the skull, two bony plates are seen projecting from each lateral wall. The larger of these is the inferior nasal concha, an independent bone of the skull. Located just above the inferior concha is the middle nasal concha, which is part of the ethmoid bone. A third bony plate, also part of the ethmoid bone, is the superior nasal concha. It is much smaller and out of sight, above the middle concha. The superior nasal concha is located just lateral to the perpendicular plate, in the upper nasal cavity.

Lateral View of Skull

A view of the lateral skull is dominated by the large, rounded brain case above and the upper and lower jaws with their teeth below (Figure 3). Separating these areas is the bridge of bone called the zygomatic arch. The zygomatic arch is the bony arch on the side of skull that spans from the area of the cheek to just above the ear canal. It is formed by the junction of two bony processes: a short anterior component, the temporal process of the zygomatic bone (the cheekbone) and a longer posterior portion, the zygomatic process of the temporal bone, extending forward from the temporal bone. Thus the temporal process (anteriorly) and the zygomatic process (posteriorly) join together, like the two ends of a drawbridge, to form the zygomatic arch. One of the major muscles that pulls the mandible upward during biting and chewing arises from the zygomatic arch.

On the lateral side of the brain case, above the level of the zygomatic arch, is a shallow space called the temporal fossa. Below the level of the zygomatic arch and deep to the vertical portion of the mandible is another space called the infratemporal fossa. Both the temporal fossa and infratemporal fossa contain muscles that act on the mandible during chewing.

[caption id="" align="aligncenter" width="480"]This image shows the lateral view of the human skull and identifies the major parts. Figure 3. Lateral View of Skull. The lateral skull shows the large rounded brain case, zygomatic arch, and the upper and lower jaws. The zygomatic arch is formed jointly by the zygomatic process of the temporal bone and the temporal process of the zygomatic bone. The shallow space above the zygomatic arch is the temporal fossa. The space inferior to the zygomatic arch and deep to the posterior mandible is the infratemporal fossa.[/caption]

Bones of the Brain Case

The brain case contains and protects the brain. The interior space that is almost completely occupied by the brain is called the cranial cavity. This cavity is bounded superiorly by the rounded top of the skull, which is called the calvaria (skullcap), and the lateral and posterior sides of the skull. The bones that form the top and sides of the brain case are usually referred to as the “flat” bones of the skull.

The floor of the brain case is referred to as the base of the skull. This is a complex area that varies in depth and has numerous openings for the passage of cranial nerves, blood vessels, and the spinal cord. Inside the skull, the base is subdivided into three large spaces, called the anterior cranial fossa, middle cranial fossa, and posterior cranial fossa (fossa = “trench or ditch”) (Figure 4). From anterior to posterior, the fossae increase in depth. The shape and depth of each fossa corresponds to the shape and size of the brain region that each houses. The boundaries and openings of the cranial fossae (singular = fossa) will be described in a later section.

The brain case consists of eight bones. These include the paired parietal and temporal bones, plus the unpaired frontal, occipital, sphenoid, and ethmoid bones.

Parietal Bone

The parietal bone forms most of the upper lateral side of the skull (see Figure 3). These are paired bones, with the right and left parietal bones joining together at the top of the skull. Each parietal bone is also bounded anteriorly by the frontal bone, inferiorly by the temporal bone, and posteriorly by the occipital bone.

Temporal Bone

The temporal bone forms the lower lateral side of the skull (see Figure 3). Common wisdom has it that the temporal bone (temporal = “time”) is so named because this area of the head (the temple) is where hair typically first turns gray, indicating the passage of time.

The temporal bone is subdivided into several regions (Figure 5). The flattened, upper portion is the squamous portion of the temporal bone. Below this area and projecting anteriorly is the zygomatic process of the temporal bone, which forms the posterior portion of the zygomatic arch. Posteriorly is the mastoid portion of the temporal bone. Projecting inferiorly from this region is a large prominence, the mastoid process, which serves as a muscle attachment site. The mastoid process can easily be felt on the side of the head just behind your earlobe. On the interior of the skull, the petrous portion of each temporal bone forms the prominent, diagonally oriented petrous ridge in the floor of the cranial cavity. Located inside each petrous ridge are small cavities that house the structures of the middle and inner ears.

[caption id="" align="aligncenter" width="450"]This image shows the location of the temporal bone. A small image of the skull on the top left shows the temporal bone highlighted in pink and a magnified view of this region then highlights the important parts of the temporal bone. Figure 5. Temporal Bone. A lateral view of the isolated temporal bone shows the squamous, mastoid, and zygomatic portions of the temporal bone.[/caption]

Frontal Bone

The frontal bone is the single bone that forms the forehead. At its anterior midline, between the eyebrows, there is a slight depression called the glabella (see Figure 3). The frontal bone also forms the supraorbital margin of the orbit. Near the middle of this margin, is the supraorbital foramen, the opening that provides passage for a sensory nerve to the forehead. The frontal bone is thickened just above each supraorbital margin, forming rounded brow ridges. These are located just behind your eyebrows and vary in size among individuals, although they are generally larger in males. Inside the cranial cavity, the frontal bone extends posteriorly. This flattened region forms both the roof of the orbit below and the floor of the anterior cranial cavity above (see Figure 6b).

Occipital Bone

The occipital bone is the single bone that forms the posterior skull and posterior base of the cranial cavity (Figure 7; see also Figure 6). On its outside surface, at the posterior midline, is a small protrusion called the external occipital protuberance, which serves as an attachment site for a ligament of the posterior neck. Lateral to either side of this bump is a superior nuchal line (nuchal = “nape” or “posterior neck”). The nuchal lines represent the most superior point at which muscles of the neck attach to the skull, with only the scalp covering the skull above these lines. On the base of the skull, the occipital bone contains the large opening of the foramen magnum, which allows for passage of the spinal cord as it exits the skull. On either side of the foramen magnum is an oval-shaped occipital condyle. These condyles form joints with the first cervical vertebra and thus support the skull on top of the vertebral column.

[caption id="" align="aligncenter" width="420"]This figure shows the posterior view of the skull and the major parts are labeled. Figure 7. Posterior View of Skull. This view of the posterior skull shows attachment sites for muscles and joints that support the skull.[/caption]

Sutures of the Skull

A suture is an immobile joint between adjacent bones of the skull. The narrow gap between the bones is filled with dense, fibrous connective tissue that unites the bones. The long sutures located between the bones of the brain case are not straight, but instead follow irregular, tightly twisting paths. These twisting lines serve to tightly interlock the adjacent bones, thus adding strength to the skull for brain protection.

The two suture lines seen on the top of the skull are the coronal and sagittal sutures. The coronal suture runs from side to side across the skull, within the coronal plane of section (see Figure 3). It joins the frontal bone to the right and left parietal bones. The sagittal suture extends posteriorly from the coronal suture, running along the midline at the top of the skull in the sagittal plane of section (see Figure 7). It unites the right and left parietal bones. On the posterior skull, the sagittal suture terminates by joining the lambdoid suture. The lambdoid suture extends downward and laterally to either side away from its junction with the sagittal suture. The lambdoid suture joins the occipital bone to the right and left parietal and temporal bones. This suture is named for its upside-down "V" shape, which resembles the capital letter version of the Greek letter lambda (Λ). The squamous suture is located on the lateral skull. It unites the squamous portion of the temporal bone with the parietal bone (see Figure 3). At the intersection of four bones is the pterion, a small, capital-H-shaped suture line region that unites the frontal bone, parietal bone, squamous portion of the temporal bone, and greater wing of the sphenoid bone. It is the weakest part of the skull. The pterion is located approximately two finger widths above the zygomatic arch and a thumb’s width posterior to the upward portion of the zygomatic bone.

Facial Bones of the Skull

The facial bones of the skull form the upper and lower jaws, the nose, nasal cavity and nasal septum, and the orbit. The facial bones include 14 bones, with six paired bones and two unpaired bones. The paired bones are the maxilla, palatine, zygomatic, nasal, lacrimal, and inferior nasal conchae bones. The unpaired bones are the vomer and mandible bones. Although classified with the brain-case bones, the ethmoid bone also contributes to the nasal septum and the walls of the nasal cavity and orbit.

Maxillary Bone

The maxillary bone, often referred to simply as the maxilla (plural = maxillae), is one of a pair that together form the upper jaw, much of the hard palate, the medial floor of the orbit, and the lateral base of the nose (see Figure 2). The curved, inferior margin of the maxillary bone that forms the upper jaw and contains the upper teeth is the alveolar process of the maxilla (Figure 12). Each tooth is anchored into a deep socket called an alveolus. On the anterior maxilla, just below the orbit, is the infraorbital foramen. This is the point of exit for a sensory nerve that supplies the nose, upper lip, and anterior cheek. On the inferior skull, the palatine process from each maxillary bone can be seen joining together at the midline to form the anterior three-quarters of the hard palate (see Figure 6a). The hard palate is the bony plate that forms the roof of the mouth and floor of the nasal cavity, separating the oral and nasal cavities.

[caption id="" align="aligncenter" width="420"]This image shows the location and structure of the maxilla. A small image of the skull on the top left shows the maxilla in ochre yellow. A magnified view shows the detailed structure of the maxilla. Figure 12. Maxillary Bone. The maxillary bone forms the upper jaw and supports the upper teeth. Each maxilla also forms the lateral floor of each orbit and the majority of the hard palate.[/caption]

Mandible

The mandible forms the lower jaw and is the only moveable bone of the skull. At the time of birth, the mandible consists of paired right and left bones, but these fuse together during the first year to form the single U-shaped mandible of the adult skull. Each side of the mandible consists of a horizontal body and posteriorly, a vertically oriented ramus of the mandible (ramus = “branch”). The outside margin of the mandible, where the body and ramus come together is called the angle of the mandible (Figure 13).

The ramus on each side of the mandible has two upward-going bony projections. The more anterior projection is the flattened coronoid process of the mandible, which provides attachment for one of the biting muscles. The posterior projection is the condylar process of the mandible, which is topped by the oval-shaped condyle. The condyle of the mandible articulates (joins) with the mandibular fossa and articular tubercle of the temporal bone. Together these articulations form the temporomandibular joint, which allows for opening and closing of the mouth (see Figure 3). The broad U-shaped curve located between the coronoid and condylar processes is the mandibular notch.

[caption id="" align="aligncenter" width="420"]This image shows the structure of the mandible. On the top left, a lateral view of the skull shows the location of the mandible in purple. A magnified image shows the right lateral view of the mandible with the major parts labeled. Figure 13. Isolated Mandible. The mandible is the only moveable bone of the skull.[/caption]

Hyoid Bone

The hyoid bone is an independent bone that does not contact any other bone and thus is not part of the skull (Figure 17). It is a small U-shaped bone located in the upper neck near the level of the inferior mandible, with the tips of the “U” pointing posteriorly. The hyoid serves as the base for the tongue above, and is attached to the larynx below and the pharynx posteriorly. The hyoid is held in position by a series of small muscles that attach to it either from above or below. These muscles act to move the hyoid up/down or forward/back. Movements of the hyoid are coordinated with movements of the tongue, larynx, and pharynx during swallowing and speaking.

[caption id="" align="aligncenter" width="300"]In this image, the location and structure of the hyoid bone are shown. The top panel shows a person’s face and neck, with the hyoid bone highlighted in grey. The middle panel shows the anterior view and the bottom panel shows the right anterior view. Figure 17. Hyoid Bone. The hyoid bone is located in the upper neck and does not join with any other bone. It provides attachments for muscles that act on the tongue, larynx, and pharynx.[/caption]

]]> 2065 2052 3 0 7.3 The Vertebral Column https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2076 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2076

Learning Objectives

By the end of this section, you will be able to:
  • Describe each region of the vertebral column and the number of bones in each region
  • Discuss the curves of the vertebral column and how these change after birth
  • Describe a typical vertebra and determine the distinguishing characteristics for vertebrae in each vertebral region and features of the sacrum and the coccyx
  • Define the structure of an intervertebral disc
  • Determine the location of the ligaments that provide support for the vertebral column

The vertebral column is also known as the spinal column or spine (Figure 1). It consists of a sequence of vertebrae (singular = vertebra), each of which is separated and united by an intervertebral disc. Together, the vertebrae and intervertebral discs form the vertebral column. It is a flexible column that supports the head, neck, and body and allows for their movements. It also protects the spinal cord, which passes down the back through openings in the vertebrae.

[caption id="" align="aligncenter" width="480"]This image shows the structure of the vertebral column. The left panel shows the front view of the vertebral column and the right panel shows the side view of the vertebral column. Figure 1. Vertebral Column. The adult vertebral column consists of 24 vertebrae, plus the sacrum and coccyx. The vertebrae are divided into three regions: cervical C1–C7 vertebrae, thoracic T1–T12 vertebrae, and lumbar L1–L5 vertebrae. The vertebral column is curved, with two primary curvatures (thoracic and sacrococcygeal curves) and two secondary curvatures (cervical and lumbar curves).[/caption]

Regions of the Vertebral Column

The vertebral column originally develops as a series of 33 vertebrae, but this number is eventually reduced to 24 vertebrae, plus the sacrum and coccyx. The vertebral column is subdivided into five regions, with the vertebrae in each area named for that region and numbered in descending order. In the neck, there are seven cervical vertebrae, each designated with the letter “C” followed by its number. Superiorly, the C1 vertebra articulates (forms a joint) with the occipital condyles of the skull. Inferiorly, C1 articulates with the C2 vertebra, and so on. Below these are the 12 thoracic vertebrae, designated T1–T12. The lower back contains the L1–L5 lumbar vertebrae. The single sacrum, which is also part of the pelvis, is formed by the fusion of five sacral vertebrae. Similarly, the coccyx, or tailbone, results from the fusion of four small coccygeal vertebrae. However, the sacral and coccygeal fusions do not start until age 20 and are not completed until middle age.

An interesting anatomical fact is that almost all mammals have seven cervical vertebrae, regardless of body size. This means that there are large variations in the size of cervical vertebrae, ranging from the very small cervical vertebrae of a shrew to the greatly elongated vertebrae in the neck of a giraffe. In a full-grown giraffe, each cervical vertebra is 11 inches tall.

Curvatures of the Vertebral Column

The adult vertebral column does not form a straight line, but instead has four curvatures along its length (see Figure 1). These curves increase the vertebral column’s strength, flexibility, and ability to absorb shock. When the load on the spine is increased, by carrying a heavy backpack for example, the curvatures increase in depth (become more curved) to accommodate the extra weight. They then spring back when the weight is removed. The four adult curvatures are classified as either primary or secondary curvatures. Primary curves are retained from the original fetal curvature, while secondary curvatures develop after birth.

During fetal development, the body is flexed anteriorly into the fetal position, giving the entire vertebral column a single curvature that is concave anteriorly. In the adult, this fetal curvature is retained in two regions of the vertebral column as the thoracic curve, which involves the thoracic vertebrae, and the sacrococcygeal curve, formed by the sacrum and coccyx. Each of these is thus called a primary curve because they are retained from the original fetal curvature of the vertebral column.

A secondary curve develops gradually after birth as the child learns to sit upright, stand, and walk. Secondary curves are concave posteriorly, opposite in direction to the original fetal curvature. The cervical curve of the neck region develops as the infant begins to hold their head upright when sitting. Later, as the child begins to stand and then to walk, the lumbar curve of the lower back develops. In adults, the lumbar curve is generally deeper in females.

Disorders associated with the curvature of the spine include kyphosis (an excessive posterior curvature of the thoracic region), lordosis (an excessive anterior curvature of the lumbar region), and scoliosis (an abnormal, lateral curvature, accompanied by twisting of the vertebral column).

General Structure of a Vertebra

Within the different regions of the vertebral column, vertebrae vary in size and shape, but they all follow a similar structural pattern. A typical vertebra will consist of a body, a vertebral arch, and seven processes (Figure 4).

The body is the anterior portion of each vertebra and is the part that supports the body weight. Because of this, the vertebral bodies progressively increase in size and thickness going down the vertebral column. The bodies of adjacent vertebrae are separated and strongly united by an intervertebral disc.

The vertebral arch forms the posterior portion of each vertebra. It consists of four parts, the right and left pedicles and the right and left laminae. Each pedicle forms one of the lateral sides of the vertebral arch. The pedicles are anchored to the posterior side of the vertebral body. Each lamina forms part of the posterior roof of the vertebral arch. The large opening between the vertebral arch and body is the vertebral foramen, which contains the spinal cord. In the intact vertebral column, the vertebral foramina of all of the vertebrae align to form the vertebral (spinal) canal, which serves as the bony protection and passageway for the spinal cord down the back. When the vertebrae are aligned together in the vertebral column, notches in the margins of the pedicles of adjacent vertebrae together form an intervertebral foramen, the opening through which a spinal nerve exits from the vertebral column (Figure 5).

Seven processes arise from the vertebral arch. Each paired transverse process projects laterally and arises from the junction point between the pedicle and lamina. The single spinous process (vertebral spine) projects posteriorly at the midline of the back. The vertebral spines can easily be felt as a series of bumps just under the skin down the middle of the back. The transverse and spinous processes serve as important muscle attachment sites. A superior articular process extends or faces upward, and an inferior articular process faces or projects downward on each side of a vertebrae. The paired superior articular processes of one vertebra join with the corresponding paired inferior articular processes from the next higher vertebra. These junctions form slightly moveable joints between the adjacent vertebrae. The shape and orientation of the articular processes vary in different regions of the vertebral column and play a major role in determining the type and range of motion available in each region.

[caption id="" align="aligncenter" width="550"]This image shows the detailed structure of each vertebra. The left panel shows the superior view of the vertebra and the right panel shows the left posterolateral view. Figure 4. Parts of a Typical Vertebra. A typical vertebra consists of a body and a vertebral arch. The arch is formed by the paired pedicles and paired laminae. Arising from the vertebral arch are the transverse, spinous, superior articular, and inferior articular processes. The vertebral foramen provides for passage of the spinal cord. Each spinal nerve exits through an intervertebral foramen, located between adjacent vertebrae. Intervertebral discs unite the bodies of adjacent vertebrae.[/caption]
[caption id="" align="aligncenter" width="480"]This image shows the structure of the intervertebral disk. The left panel shows the lateral view and the right panel shows the superior view. Figure 5. Intervertebral Disc. The bodies of adjacent vertebrae are separated and united by an intervertebral disc, which provides padding and allows for movements between adjacent vertebrae. The disc consists of a fibrous outer layer called the anulus fibrosus and a gel-like center called the nucleus pulposus. The intervertebral foramen is the opening formed between adjacent vertebrae for the exit of a spinal nerve.[/caption]

Regional Modifications of Vertebrae

In addition to the general characteristics of a typical vertebra described above, vertebrae also display characteristic size and structural features that vary between the different vertebral column regions. Thus, cervical vertebrae are smaller than lumbar vertebrae due to differences in the proportion of body weight that each supports. Thoracic vertebrae have sites for rib attachment, and the vertebrae that give rise to the sacrum and coccyx have fused together into single bones.

Cervical Vertebrae

Typical cervical vertebrae, such as C4 or C5, have several characteristic features that differentiate them from thoracic or lumbar vertebrae (Figure 6). Cervical vertebrae have a small body, reflecting the fact that they carry the least amount of body weight. Cervical vertebrae usually have a bifid (Y-shaped) spinous process. The spinous processes of the C3–C6 vertebrae are short, but the spine of C7 is much longer. You can find these vertebrae by running your finger down the midline of the posterior neck until you encounter the prominent C7 spine located at the base of the neck. The transverse processes of the cervical vertebrae are sharply curved (U-shaped) to allow for passage of the cervical spinal nerves. Each transverse process also has an opening called the transverse foramen. An important artery that supplies the brain ascends up the neck by passing through these openings. The superior and inferior articular processes of the cervical vertebrae are flattened and largely face upward or downward, respectively.

The first and second cervical vertebrae are further modified, giving each a distinctive appearance. The first cervical (C1) vertebra is also called the atlas, because this is the vertebra that supports the skull on top of the vertebral column (in Greek mythology, Atlas was the god who supported the heavens on his shoulders). The C1 vertebra does not have a body or spinous process. Instead, it is ring-shaped, consisting of an anterior arch and a posterior arch. The transverse processes of the atlas are longer and extend more laterally than do the transverse processes of any other cervical vertebrae. The superior articular processes face upward and are deeply curved for articulation with the occipital condyles on the base of the skull. The inferior articular processes are flat and face downward to join with the superior articular processes of the C2 vertebra.

The second cervical (C2) vertebra is called the axis, because it serves as the axis for rotation when turning the head toward the right or left. The axis resembles typical cervical vertebrae in most respects, but is easily distinguished by the dens (odontoid process), a bony projection that extends upward from the vertebral body. The dens joins with the inner aspect of the anterior arch of the atlas, where it is held in place by transverse ligament.

[caption id="" align="aligncenter" width="550"]This figure shows the structure of the cervical vertebrae. The left panel shows the location of the cervical vertebrae in green along the vertebral column. The middle panel shows the structure of a typical cervical vertebra and the right panel shows the superior and anterior view of the axis. Figure 6. Cervical Vertebrae. A typical cervical vertebra has a small body, a bifid spinous process, transverse processes that have a transverse foramen and are curved for spinal nerve passage. The atlas (C1 vertebra) does not have a body or spinous process. It consists of an anterior and a posterior arch and elongated transverse processes. The axis (C2 vertebra) has the upward projecting dens, which articulates with the anterior arch of the atlas.[/caption]

Thoracic Vertebrae

The bodies of the thoracic vertebrae are larger than those of cervical vertebrae (Figure 7). The characteristic feature for a typical midthoracic vertebra is the spinous process, which is long and has a pronounced downward angle that causes it to overlap the next inferior vertebra. The superior articular processes of thoracic vertebrae face anteriorly and the inferior processes face posteriorly. These orientations are important determinants for the type and range of movements available to the thoracic region of the vertebral column.

Thoracic vertebrae have several additional articulation sites, each of which is called a facet, where a rib is attached. Most thoracic vertebrae have two facets located on the lateral sides of the body, each of which is called a costal facet (costal = “rib”). These are for articulation with the head (end) of a rib. An additional facet is located on the transverse process for articulation with the tubercle of a rib.

[caption id="" align="aligncenter" width="450"]This figure shows the structure of the thoracic vertebra. The left panel shows the vertebral column with the thoracic vertebrae highlighted in pink. The right panel shows the detailed structure of a single thoracic vertebra. Figure 7. Thoracic Vertebrae. A typical thoracic vertebra is distinguished by the spinous process, which is long and projects downward to overlap the next inferior vertebra. It also has articulation sites (facets) on the vertebral body and a transverse process for rib attachment.[/caption]
[caption id="" align="aligncenter" width="380"]This diagram shows how the thoracic vertebra connects to the angle of the rib. The major parts of the vertebra and the processes connecting the vertebra to the rib are labeled. Figure 8. Rib Articulation in Thoracic Vertebrae. Thoracic vertebrae have superior and inferior articular facets on the vertebral body for articulation with the head of a rib, and a transverse process facet for articulation with the rib tubercle.[/caption]

Lumbar Vertebrae

Lumbar vertebrae carry the greatest amount of body weight and are thus characterized by the large size and thickness of the vertebral body (Figure 9). They have short transverse processes and a short, blunt spinous process that projects posteriorly. The articular processes are large, with the superior process facing backward and the inferior facing forward.

[caption id="" align="aligncenter" width="350"]This image shows the location and structure of the lumbar vertebrae. The left panel shows the location of the lumbar vertebrae (highlighted in green) along the vertebral column. The right panel shows the inferior articular process and the major parts are labeled. Figure 9. Lumbar Vertebrae. Lumbar vertebrae are characterized by having a large, thick body and a short, rounded spinous process.[/caption]

Sacrum and Coccyx

The sacrum is a triangular-shaped bone that is thick and wide across its superior base where it is weight bearing and then tapers down to an inferior, non-weight bearing apex (Figure 10). It is formed by the fusion of five sacral vertebrae, a process that does not begin until after the age of 20. On the anterior surface of the older adult sacrum, the lines of vertebral fusion can be seen as four transverse ridges. On the posterior surface, running down the midline, is the median sacral crest, a bumpy ridge that is the remnant of the fused spinous processes (median = “midline”; while medial = “toward, but not necessarily at, the midline”). Similarly, the fused transverse processes of the sacral vertebrae form the lateral sacral crest.

The sacral promontory is the anterior lip of the superior base of the sacrum. Lateral to this is the roughened auricular surface, which joins with the ilium portion of the hipbone to form the immobile sacroiliac joints of the pelvis. Passing inferiorly through the sacrum is a bony tunnel called the sacral canal, which terminates at the sacral hiatus near the inferior tip of the sacrum. The anterior and posterior surfaces of the sacrum have a series of paired openings called sacral foramina (singular = foramen) that connect to the sacral canal. Each of these openings is called a posterior (dorsal) sacral foramen or anterior (ventral) sacral foramen. These openings allow for the anterior and posterior branches of the sacral spinal nerves to exit the sacrum. The superior articular process of the sacrum, one of which is found on either side of the superior opening of the sacral canal, articulates with the inferior articular processes from the L5 vertebra.

The coccyx, or tailbone, is derived from the fusion of four very small coccygeal vertebrae (see Figure 10). It articulates with the inferior tip of the sacrum. It is not weight bearing in the standing position, but may receive some body weight when sitting.

[caption id="" align="aligncenter" width="550"]This figure shows the structure of the sacrum and coccyx. The left panel shows the vertebral column with the sacrum and coccyx highlighted in pink. The middle panel shows the anterior view and the right panel shows the posterior view of the sacrum and coccyx. Figure 10. Sacrum and Coccyx. The sacrum is formed from the fusion of five sacral vertebrae, whose lines of fusion are indicated by the transverse ridges. The fused spinous processes form the median sacral crest, while the lateral sacral crest arises from the fused transverse processes. The coccyx is formed by the fusion of four small coccygeal vertebrae.[/caption]

Intervertebral Discs and Ligaments of the Vertebral Column

The bodies of adjacent vertebrae are strongly anchored to each other by an intervertebral disc. This structure provides padding between the bones during weight bearing, and because it can change shape, also allows for movement between the vertebrae. Although the total amount of movement available between any two adjacent vertebrae is small, when these movements are summed together along the entire length of the vertebral column, large body movements can be produced. Ligaments that extend along the length of the vertebral column also contribute to its overall support and stability.

Intervertebral Disc

An intervertebral disc is a fibrocartilaginous pad that fills the gap between adjacent vertebral bodies (see Figure 5). Each disc is anchored to the bodies of its adjacent vertebrae, thus strongly uniting these. The discs also provide padding between vertebrae during weight bearing. Because of this, intervertebral discs are thin in the cervical region and thickest in the lumbar region, which carries the most body weight. In total, the intervertebral discs account for approximately 25 percent of your body height between the top of the pelvis and the base of the skull. Intervertebral discs are also flexible and can change shape to allow for movements of the vertebral column.

Each intervertebral disc consists of two parts. The anulus fibrosus is the tough, fibrous outer layer of the disc. It forms a circle (anulus = “ring” or “circle”) and is firmly anchored to the outer margins of the adjacent vertebral bodies. Inside is the nucleus pulposus, consisting of a softer, more gel-like material. It has a high water content that serves to resist compression and thus is important for weight bearing. With increasing age, the water content of the nucleus pulposus gradually declines. This causes the disc to become thinner, decreasing total body height somewhat, and reduces the flexibility and range of motion of the disc, making bending more difficult.

[caption id="" align="alignright" width="120"]QR Code representing a URL Watch this animation to see what it means to “slip” a disk. Watch this second animation to see one possible treatment for a herniated disc, removing and replacing the damaged disc with an artificial one that allows for movement between the adjacent certebrae.[/caption] The gel-like nature of the nucleus pulposus also allows the intervertebral disc to change shape as one vertebra rocks side to side or forward and back in relation to its neighbors during movements of the vertebral column. Thus, bending forward causes compression of the anterior portion of the disc but expansion of the posterior disc. If the posterior anulus fibrosus is weakened due to injury or increasing age, the pressure exerted on the disc when bending forward and lifting a heavy object can cause the nucleus pulposus to protrude posteriorly through the anulus fibrosus, resulting in a herniated disc (“ruptured” or “slipped” disc) (Figure 11). The posterior bulging of the nucleus pulposus can cause compression of a spinal nerve at the point where it exits through the intervertebral foramen, with resulting pain and/or muscle weakness in those body regions supplied by that nerve. The most common sites for disc herniation are the L4/L5 or L5/S1 intervertebral discs, which can cause sciatica, a widespread pain that radiates from the lower back down the thigh and into the leg. Similar injuries of the C5/C6 or C6/C7 intervertebral discs, following forcible hyperflexion of the neck from a collision accident or football injury, can produce pain in the neck, shoulder, and upper limb.
[caption id="" align="aligncenter" width="450"]This figure shows a herniated disk. The left panel shows the superior view highlighting how the herniated disk compresses the nerve. The right panel shows a photograph of a herniated disk. Figure 11. Herniated Intervertebral Disc. Weakening of the anulus fibrosus can result in herniation (protrusion) of the nucleus pulposus and compression of a spinal nerve, resulting in pain and/or muscle weakness in the body regions supplied by that nerve.[/caption]
]]> 2076 2052 4 0 7.4 The Thoracic Cage https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2078 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2078

Learning Objectives

By the end of this section, you will be able to:
  • Discuss the components that make up the thoracic cage
  • Identify the parts of the sternum and define the sternal angle
  • Discuss the parts of a rib and rib classifications

The thoracic cage (rib cage) forms the thorax (chest) portion of the body. It consists of the 12 pairs of ribs with their costal cartilages and the sternum (Figure 1). The ribs are anchored posteriorly to the 12 thoracic vertebrae (T1–T12). The thoracic cage protects the heart and lungs.

[caption id="" align="aligncenter" width="550"]This figure shows the skeletal structure of the rib cage. The left panel shows the anterior view of the sternum and the right panel shows the anterior panel of the sternum including the entire rib cage. Figure 1. Thoracic Cage. The thoracic cage is formed by the (a) sternum and (b) 12 pairs of ribs with their costal cartilages. The ribs are anchored posteriorly to the 12 thoracic vertebrae. The sternum consists of the manubrium, body, and xiphoid process. The ribs are classified as true ribs (1–7) and false ribs (8–12). The last two pairs of false ribs are also known as floating ribs (11–12).[/caption]

Sternum

The sternum is the elongated bony structure that anchors the anterior thoracic cage. It consists of three parts: the manubrium, body, and xiphoid process. The manubrium is the wider, superior portion of the sternum. The top of the manubrium has a shallow, U-shaped border called the jugular (suprasternal) notch. This can be easily felt at the anterior base of the neck, between the medial ends of the clavicles. The clavicular notch is the shallow depression located on either side at the superior-lateral margins of the manubrium. This is the site of the sternoclavicular joint, between the sternum and clavicle. The first ribs also attach to the manubrium.

The elongated, central portion of the sternum is the body. The manubrium and body join together at the sternal angle, so called because the junction between these two components is not flat, but forms a slight bend. The second rib attaches to the sternum at the sternal angle. Since the first rib is hidden behind the clavicle, the second rib is the highest rib that can be identified by palpation. Thus, the sternal angle and second rib are important landmarks for the identification and counting of the lower ribs. Ribs 3–7 attach to the sternal body.

The inferior tip of the sternum is the xiphoid process. This small structure is cartilaginous early in life, but gradually becomes ossified starting during middle age.

Ribs

Each rib is a curved, flattened bone that contributes to the wall of the thorax. The ribs articulate posteriorly with the T1–T12 thoracic vertebrae, and most attach anteriorly via their costal cartilages to the sternum. There are 12 pairs of ribs. The ribs are numbered 1–12 in accordance with the thoracic vertebrae.

Parts of a Typical Rib

The posterior end of a typical rib is called the head of the rib (see Chapter 7.3 Figure 8). This region articulates primarily with the costal facet located on the body of the same numbered thoracic vertebra and to a lesser degree, with the costal facet located on the body of the next higher vertebra. Lateral to the head is the narrowed neck of the rib. A small bump on the posterior rib surface is the tubercle of the rib, which articulates with the facet located on the transverse process of the same numbered vertebra. The remainder of the rib is the body of the rib (shaft). Just lateral to the tubercle is the angle of the rib, the point at which the rib has its greatest degree of curvature. The angles of the ribs form the most posterior extent of the thoracic cage. In the anatomical position, the angles align with the medial border of the scapula. A shallow costal groove for the passage of blood vessels and a nerve is found along the inferior margin of each rib.

Rib Classifications

The bony ribs do not extend anteriorly completely around to the sternum. Instead, each rib ends in a costal cartilage. These cartilages are made of hyaline cartilage and can extend for several inches. Most ribs are then attached, either directly or indirectly, to the sternum via their costal cartilage (see Figure 1). The ribs are classified into three groups based on their relationship to the sternum.

Ribs 1–7 are classified as true ribs (vertebrosternal ribs). The costal cartilage from each of these ribs attaches directly to the sternum. Ribs 8–12 are called false ribs (vertebrochondral ribs). The costal cartilages from these ribs do not attach directly to the sternum. For ribs 8–10, the costal cartilages are attached to the cartilage of the next higher rib. Thus, the cartilage of rib 10 attaches to the cartilage of rib 9, rib 9 then attaches to rib 8, and rib 8 is attached to rib 7. The last two false ribs (11–12) are also called floating ribs (vertebral ribs). These are short ribs that do not attach to the sternum at all. Instead, their small costal cartilages terminate within the musculature of the lateral abdominal wall.1.

]]> 2078 2052 5 0 7.5 Embryonic Development of the Axial Skeleton https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2081 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2081

Learning Objectives

By the end of this section, you will be able to:
  • Discuss the two types of embryonic bone development within the skull
  • Describe the development of the vertebral column and thoracic cage

The axial skeleton begins to form during early embryonic development. However, growth, remodeling, and ossification (bone formation) continue for several decades after birth before the adult skeleton is fully formed. Knowledge of the developmental processes that give rise to the skeleton is important for understanding the abnormalities that may arise in skeletal structures.

Development of the Skull

The bones of the skull arise from mesenchyme during embryonic development in two different ways. The first mechanism produces the bones that form the top and sides of the brain case. This involves the local accumulation of mesenchymal cells at the site of the future bone. These cells then differentiate directly into bone producing cells, which form the skull bones through the process of intramembranous ossification. As the brain case bones grow in the fetal skull, they remain separated from each other by large areas of dense connective tissue, each of which is called a fontanelle (Figure 1). The fontanelles are the soft spots on an infant’s head. They are important during birth because these areas allow the skull to change shape as it squeezes through the birth canal. After birth, the fontanelles allow for continued growth and expansion of the skull as the brain enlarges. The largest fontanelle is located on the anterior head, at the junction of the frontal and parietal bones. The fontanelles decrease in size and disappear by age 2. However, the skull bones remained separated from each other at the sutures, which contain dense fibrous connective tissue that unites the adjacent bones. The connective tissue of the sutures allows for continued growth of the skull bones as the brain enlarges during childhood growth.

The second mechanism for bone development in the skull produces the facial bones and floor of the brain case. This also begins with the localized accumulation of mesenchymal cells. However, these cells differentiate into cartilage cells, which produce a hyaline cartilage model of the future bone. As this cartilage model grows, it is gradually converted into bone through the process of endochondral ossification. This is a slow process and the cartilage is not completely converted to bone until the skull achieves its full adult size.

At birth, the brain case and orbits of the skull are disproportionally large compared to the bones of the jaws and lower face. This reflects the relative underdevelopment of the maxilla and mandible, which lack teeth, and the small sizes of the paranasal sinuses and nasal cavity. During early childhood, the mastoid process enlarges, the two halves of the mandible and frontal bone fuse together to form single bones, and the paranasal sinuses enlarge. The jaws also expand as the teeth begin to appear. These changes all contribute to the rapid growth and enlargement of the face during childhood.

[caption id="" align="aligncenter" width="550"]This diagram shows the image of a newborn human skull. The major parts of the skull are labeled. The left panel shows the superior view (from the top) and the right side shows the lateral view (from the side). Figure 1. Newborn Skull. The bones of the newborn skull are not fully ossified and are separated by large areas called fontanelles, which are filled with fibrous connective tissue. The fontanelles allow for continued growth of the skull after birth. At the time of birth, the facial bones are small and underdeveloped, and the mastoid process has not yet formed.[/caption]

Development of the Vertebral Column and Thoracic cage

Development of the vertebrae begins with the accumulation of mesenchyme cells from each sclerotome around the notochord. These cells differentiate into a hyaline cartilage model for each vertebra, which then grow and eventually ossify into bone through the process of endochondral ossification. As the developing vertebrae grow, the notochord largely disappears. However, small areas of notochord tissue persist between the adjacent vertebrae and this contributes to the formation of each intervertebral disc.

[caption id="" align="alignright" width="120"]QR Code representing a URL View this video to review the two processes that give rise to the bones of the skull and body.[/caption] The ribs and sternum also develop from mesenchyme. The ribs initially develop as part of the cartilage model for each vertebra, but in the thorax region, the rib portion separates from the vertebra by the eighth week. The cartilage model of the rib then ossifies, except for the anterior portion, which remains as the costal cartilage. The sternum initially forms as paired hyaline cartilage models on either side of the anterior midline, beginning during the fifth week of development. The cartilage models of the ribs become attached to the lateral sides of the developing sternum. Eventually, the two halves of the cartilaginous sternum fuse together along the midline and then ossify into bone. The manubrium and body of the sternum are converted into bone first, with the xiphoid process remaining as cartilage until late in life.
]]> 2081 2052 6 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2084 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2084 [caption id="" align="aligncenter" width="500"]This photograph shows a dancer striking a pose. Figure 1. Dancer. The appendicular skeleton consists of the upper and lower limb bones, the bones of the hands and feet, and the bones that anchor the limbs to the axial skeleton. (credit: Melissa Dooley/flickr)[/caption]

Chapter Objectives

After this chapter, you will be able to:
  • Discuss the bones of the pectoral and pelvic girdles, and describe how these unite the limbs with the axial skeleton
  • Describe the bones of the upper limb, including the bones of the arm, forearm, wrist, and hand
  • Identify the features of the pelvis and explain how these differ between the adult male and female pelvis
  • Describe the bones of the lower limb, including the bones of the thigh, leg, ankle, and foot
  • Describe the embryonic formation and growth of the limb bones
Your skeleton provides the internal supporting structure of the body. The adult axial skeleton consists of 80 bones that form the head and body trunk. Attached to this are the limbs, whose 126 bones constitute the appendicular skeleton. These bones are divided into two groups: the bones that are located within the limbs themselves, and the girdle bones that attach the limbs to the axial skeleton. The bones of the shoulder region form the pectoral girdle, which anchors the upper limb to the thoracic cage of the axial skeleton. The lower limb is attached to the vertebral column by the pelvic girdle. Because of our upright stance, different functional demands are placed upon the upper and lower limbs. Thus, the bones of the lower limbs are adapted for weight-bearing support and stability, as well as for body locomotion via walking or running. In contrast, our upper limbs are not required for these functions. Instead, our upper limbs are highly mobile and can be utilized for a wide variety of activities. The large range of upper limb movements, coupled with the ability to easily manipulate objects with our hands and opposable thumbs, has allowed humans to construct the modern world in which we live.]]> 2084 2082 1 0 8.1 The Pectoral Girdle https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2088 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2088

Learning Objectives

By the end of this section, you will be able to:
  • Describe the bones that form the pectoral girdle
  • List the functions of the pectoral girdle

The appendicular skeleton includes all of the limb bones, plus the bones that unite each limb with the axial skeleton (Figure 1). The bones that attach each upper limb to the axial skeleton form the pectoral girdle (shoulder girdle). This consists of two bones, the scapula and clavicle (Figure 2). The clavicle (collarbone) is an S-shaped bone located on the anterior side of the shoulder. It is attached on its medial end to the sternum of the thoracic cage, which is part of the axial skeleton. The lateral end of the clavicle articulates (joins) with the scapula just above the shoulder joint. You can easily palpate, or feel with your fingers, the entire length of your clavicle.

[caption id="" align="aligncenter" width="600"]This figure shows the human skeleton. The left panel shows the anterior view, and the right panel shows the posterior view. Figure 1. Axial and Appendicular Skeletons. The axial skeleton forms the central axis of the body and consists of the skull, vertebral column, and thoracic cage. The appendicular skeleton consists of the pectoral and pelvic girdles, the limb bones, and the bones of the hands and feet.[/caption]
[caption id="" align="aligncenter" width="450"]This figure shows the rib change. The top left panel shows the anterior view, and the top right panel shows the posterior view. The bottom panel shows two bones. Figure 2. Pectoral Girdle. The pectoral girdle consists of the clavicle and the scapula, which serve to attach the upper limb to the sternum of the axial skeleton.[/caption]
The scapula (shoulder blade) lies on the posterior aspect of the shoulder. It is supported by the clavicle, which also articulates with the humerus (arm bone) to form the shoulder joint. The scapula is a flat, triangular-shaped bone with a prominent ridge running across its posterior surface. This ridge extends out laterally, where it forms the bony tip of the shoulder and joins with the lateral end of the clavicle. By following along the clavicle, you can palpate out to the bony tip of the shoulder, and from there, you can move back across your posterior shoulder to follow the ridge of the scapula. Move your shoulder around and feel how the clavicle and scapula move together as a unit. Both of these bones serve as important attachment sites for muscles that aid with movements of the shoulder and arm.

The right and left pectoral girdles are not joined to each other, allowing each to operate independently. In addition, the clavicle of each pectoral girdle is anchored to the axial skeleton by a single, highly mobile joint. This allows for the extensive mobility of the entire pectoral girdle, which in turn enhances movements of the shoulder and upper limb.

Clavicle

The clavicle is the only long bone that lies in a horizontal position in the body (see Figure 2). The clavicle has several important functions. First, anchored by muscles from above, it serves as a strut that extends laterally to support the scapula. This in turn holds the shoulder joint superiorly and laterally from the body trunk, allowing for maximal freedom of motion for the upper limb. The clavicle also transmits forces acting on the upper limb to the sternum and axial skeleton. Finally, it serves to protect the underlying nerves and blood vessels as they pass between the trunk of the body and the upper limb.

The clavicle has three regions: the medial end, the lateral end, and the shaft. The medial end, known as the sternal end of the clavicle, has a triangular shape and articulates with the manubrium portion of the sternum. This forms the sternoclavicular joint, which is the only bony articulation between the pectoral girdle of the upper limb and the axial skeleton. This joint allows considerable mobility, enabling the clavicle and scapula to move in upward/downward and anterior/posterior directions during shoulder movements. The sternoclavicular joint is indirectly supported by the costoclavicular ligament (costo- = “rib”), which spans the sternal end of the clavicle and the underlying first rib. The lateral or acromial end of the clavicle articulates with the acromion of the scapula, the portion of the scapula that forms the bony tip of the shoulder. There are some sex differences in the morphology of the clavicle. In women, the clavicle tends to be shorter, thinner, and less curved. In men, the clavicle is heavier and longer, and has a greater curvature and rougher surfaces where muscles attach, features that are more pronounced in manual workers.

The clavicle is the most commonly fractured bone in the body. Such breaks often occur because of the force exerted on the clavicle when a person falls onto his or her outstretched arms, or when the lateral shoulder receives a strong blow. Because the sternoclavicular joint is strong and rarely dislocated, excessive force results in the breaking of the clavicle, usually between the middle and lateral portions of the bone. If the fracture is complete, the shoulder and lateral clavicle fragment will drop due to the weight of the upper limb, causing the person to support the sagging limb with their other hand. Muscles acting across the shoulder will also pull the shoulder and lateral clavicle anteriorly and medially, causing the clavicle fragments to override. The clavicle overlies many important blood vessels and nerves for the upper limb, but fortunately, due to the anterior displacement of a broken clavicle, these structures are rarely affected when the clavicle is fractured.

Scapula

The scapula is also part of the pectoral girdle and thus plays an important role in anchoring the upper limb to the body. The scapula is located on the posterior side of the shoulder. It is surrounded by muscles on both its anterior (deep) and posterior (superficial) sides, and thus does not articulate with the ribs of the thoracic cage.

The scapula has several important landmarks (Figure 3). The three margins or borders of the scapula, named for their positions within the body, are the superior border of the scapula, the medial border of the scapula, and the lateral border of the scapula. The suprascapular notch is located lateral to the midpoint of the superior border. The corners of the triangular scapula, at either end of the medial border, are the superior angle of the scapula, located between the medial and superior borders, and the inferior angle of the scapula, located between the medial and lateral borders. The inferior angle is the most inferior portion of the scapula, and is particularly important because it serves as the attachment point for several powerful muscles involved in shoulder and upper limb movements. The remaining corner of the scapula, between the superior and lateral borders, is the location of the glenoid cavity (glenoid fossa). This shallow depression articulates with the humerus bone of the arm to form the glenohumeral joint (shoulder joint). The small bony bumps located immediately above and below the glenoid cavity are the supraglenoid tubercle and the infraglenoid tubercle, respectively. These provide attachments for muscles of the arm.

[caption id="" align="aligncenter" width="450"]This diagram shows the anterior and posterior view of the scapula. Figure 3. Scapula. The isolated scapula is shown here from its anterior (deep) side and its posterior (superficial) side.[/caption]

The scapula also has two prominent projections. Toward the lateral end of the superior border, between the suprascapular notch and glenoid cavity, is the hook-like coracoid process (coracoid = “shaped like a crow’s beak”). This process projects anteriorly and curves laterally. At the shoulder, the coracoid process is located inferior to the lateral end of the clavicle. It is anchored to the clavicle by a strong ligament, and serves as the attachment site for muscles of the anterior chest and arm. On the posterior aspect, the spine of the scapula is a long and prominent ridge that runs across its upper portion. Extending laterally from the spine is a flattened and expanded region called the acromion or acromial process. The acromion forms the bony tip of the superior shoulder region and articulates with the lateral end of the clavicle, forming the acromioclavicular joint (see Figure 2). Together, the clavicle, acromion, and spine of the scapula form a V-shaped bony line that provides for the attachment of neck and back muscles that act on the shoulder, as well as muscles that pass across the shoulder joint to act on the arm.

The scapula has three depressions, each of which is called a fossa (plural = fossae). Two of these are found on the posterior scapula, above and below the scapular spine. Superior to the spine is the narrow supraspinous fossa, and inferior to the spine is the broad infraspinous fossa. The anterior (deep) surface of the scapula forms the broad subscapular fossa. All of these fossae provide large surface areas for the attachment of muscles that cross the shoulder joint to act on the humerus.

The acromioclavicular joint transmits forces from the upper limb to the clavicle. The ligaments around this joint are relatively weak. A hard fall onto the elbow or outstretched hand can stretch or tear the acromioclavicular ligaments, resulting in a moderate injury to the joint. However, the primary support for the acromioclavicular joint comes from a very strong ligament called the coracoclavicular ligament (see Figure 2). This connective tissue band anchors the coracoid process of the scapula to the inferior surface of the acromial end of the clavicle and thus provides important indirect support for the acromioclavicular joint. Following a strong blow to the lateral shoulder, such as when a hockey player is driven into the boards, a complete dislocation of the acromioclavicular joint can result. In this case, the acromion is thrust under the acromial end of the clavicle, resulting in ruptures of both the acromioclavicular and coracoclavicular ligaments. The scapula then separates from the clavicle, with the weight of the upper limb pulling the shoulder downward. This dislocation injury of the acromioclavicular joint is known as a “shoulder separation” and is common in contact sports such as hockey, football, or martial arts.

]]> 2088 2082 2 0 8.2 Bones of the Upper Limb https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2095 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2095

Learning Objectives

By the end of this section, you will be able to:
  • Identify the divisions of the upper limb and describe the bones in each region
  • List the bones and bony landmarks that articulate at each joint of the upper limb

The upper limb is divided into three regions. These consist of the arm, located between the shoulder and elbow joints; the forearm, which is between the elbow and wrist joints; and the hand, which is located distal to the wrist. There are 30 bones in each upper limb (see Chapter 8.1 Figure 1). The humerus is the single bone of the upper arm, and the ulna (medially) and the radius (laterally) are the paired bones of the forearm. The base of the hand contains eight bones, each called a carpal bone, and the palm of the hand is formed by five bones, each called a metacarpal bone. The fingers and thumb contain a total of 14 bones, each of which is a phalanx bone of the hand.

Humerus

The humerus is the single bone of the upper arm region (Figure 1). At its proximal end is the head of the humerus. This is the large, round, smooth region that faces medially. The head articulates with the glenoid cavity of the scapula to form the glenohumeral (shoulder) joint. The margin of the smooth area of the head is the anatomical neck of the humerus. Located on the lateral side of the proximal humerus is an expanded bony area called the greater tubercle. The smaller lesser tubercle of the humerus is found on the anterior aspect of the humerus. Both the greater and lesser tubercles serve as attachment sites for muscles that act across the shoulder joint. Passing between the greater and lesser tubercles is the narrow intertubercular groove (sulcus), which is also known as the bicipital groove because it provides passage for a tendon of the biceps brachii muscle. The surgical neck is located at the base of the expanded, proximal end of the humerus, where it joins the narrow shaft of the humerus. The surgical neck is a common site of arm fractures. The deltoid tuberosity is a roughened, V-shaped region located on the lateral side in the middle of the humerus shaft. As its name indicates, it is the site of attachment for the deltoid muscle.
[caption id="" align="aligncenter" width="350"]This diagram shows the bones of the upper arm and the elbow joint. The left panel shows the anterior view, and the right panel shows the posterior view. Figure 1. Humerus and Elbow Joint. The humerus is the single bone of the upper arm region. It articulates with the radius and ulna bones of the forearm to form the elbow joint.[/caption]

Distally, the humerus becomes flattened. The prominent bony projection on the medial side is the medial epicondyle of the humerus. The much smaller lateral epicondyle of the humerus is found on the lateral side of the distal humerus. The roughened ridge of bone above the lateral epicondyle is the lateral supracondylar ridge. All of these areas are attachment points for muscles that act on the forearm, wrist, and hand. The powerful grasping muscles of the anterior forearm arise from the medial epicondyle, which is thus larger and more robust than the lateral epicondyle that gives rise to the weaker posterior forearm muscles.

The distal end of the humerus has two articulation areas, which join the ulna and radius bones of the forearm to form the elbow joint. The more medial of these areas is the trochlea, a spindle- or pulley-shaped region (trochlea = “pulley”), which articulates with the ulna bone. Immediately lateral to the trochlea is the capitulum (“small head”), a knob-like structure located on the anterior surface of the distal humerus. The capitulum articulates with the radius bone of the forearm. Just above these bony areas are two small depressions. These spaces accommodate the forearm bones when the elbow is fully bent (flexed). Superior to the trochlea is the coronoid fossa, which receives the coronoid process of the ulna, and above the capitulum is the radial fossa, which receives the head of the radius when the elbow is flexed. Similarly, the posterior humerus has the olecranon fossa, a larger depression that receives the olecranon process of the ulna when the forearm is fully extended.

Ulna

The ulna is the medial bone of the forearm. It runs parallel to the radius, which is the lateral bone of the forearm (Figure 2). The proximal end of the ulna resembles a crescent wrench with its large, C-shaped trochlear notch. This region articulates with the trochlea of the humerus as part of the elbow joint. The inferior margin of the trochlear notch is formed by a prominent lip of bone called the coronoid process of the ulna. Just below this on the anterior ulna is a roughened area called the ulnar tuberosity. To the lateral side and slightly inferior to the trochlear notch is a small, smooth area called the radial notch of the ulna. This area is the site of articulation between the proximal radius and the ulna, forming the proximal radioulnar joint. The posterior and superior portions of the proximal ulna make up the olecranon process, which forms the bony tip of the elbow.
[caption id="" align="aligncenter" width="350"]This figure shows the bones of the lower arm. Figure 2. Ulna and Radius. The ulna is located on the medial side of the forearm, and the radius is on the lateral side. These bones are attached to each other by an interosseous membrane.[/caption]

More distal is the shaft of the ulna. The lateral side of the shaft forms a ridge called the interosseous border of the ulna. This is the line of attachment for the interosseous membrane of the forearm, a sheet of dense connective tissue that unites the ulna and radius bones. The small, rounded area that forms the distal end is the head of the ulna. Projecting from the posterior side of the ulnar head is the styloid process of the ulna, a short bony projection. This serves as an attachment point for a connective tissue structure that unites the distal ends of the ulna and radius.

In the anatomical position, with the elbow fully extended and the palms facing forward, the arm and forearm do not form a straight line. Instead, the forearm deviates laterally by 5–15 degrees from the line of the arm. This deviation is called the carrying angle. It allows the forearm and hand to swing freely or to carry an object without hitting the hip. The carrying angle is larger in females to accommodate their wider pelvis.

Radius

The radius runs parallel to the ulna, on the lateral (thumb) side of the forearm (see Figure 2). The head of the radius is a disc-shaped structure that forms the proximal end. The small depression on the surface of the head articulates with the capitulum of the humerus as part of the elbow joint, whereas the smooth, outer margin of the head articulates with the radial notch of the ulna at the proximal radioulnar joint. The neck of the radius is the narrowed region immediately below the expanded head. Inferior to this point on the medial side is the radial tuberosity, an oval-shaped, bony protuberance that serves as a muscle attachment point. The shaft of the radius is slightly curved and has a small ridge along its medial side. This ridge forms the interosseous border of the radius, which, like the similar border of the ulna, is the line of attachment for the interosseous membrane that unites the two forearm bones. The distal end of the radius has a smooth surface for articulation with two carpal bones to form the radiocarpal joint or wrist joint (Figure 3 and Figure 4). On the medial side of the distal radius is the ulnar notch of the radius. This shallow depression articulates with the head of the ulna, which together form the distal radioulnar joint. The lateral end of the radius has a pointed projection called the styloid process of the radius. This provides attachment for ligaments that support the lateral side of the wrist joint. Compared to the styloid process of the ulna, the styloid process of the radius projects more distally, thereby limiting the range of movement for lateral deviations of the hand at the wrist joint.

Carpal Bones

The wrist and base of the hand are formed by a series of eight small carpal bones (see Figure 3). The carpal bones are arranged in two rows, forming a proximal row of four carpal bones and a distal row of four carpal bones. The bones in the proximal row, running from the lateral (thumb) side to the medial side, are the scaphoid (“boat-shaped”), lunate (“moon-shaped”), triquetrum (“three-cornered”), and pisiform (“pea-shaped”) bones. The small, rounded pisiform bone articulates with the anterior surface of the triquetrum bone. The pisiform thus projects anteriorly, where it forms the bony bump that can be felt at the medial base of your hand. The distal bones (lateral to medial) are the trapezium (“table”), trapezoid (“resembles a table”), capitate (“head-shaped”), and hamate (“hooked bone”) bones. The hamate bone is characterized by a prominent bony extension on its anterior side called the hook of the hamate bone.

A helpful mnemonic for remembering the arrangement of the carpal bones is “So Long To Pinky, Here Comes The Thumb.” This mnemonic starts on the lateral side and names the proximal bones from lateral to medial (scaphoid, lunate, triquetrum, pisiform), then makes a U-turn to name the distal bones from medial to lateral (hamate, capitate, trapezoid, trapezium). Thus, it starts and finishes on the lateral side.

[caption id="" align="aligncenter" width="480"]This figure shows the bones in the hand and wrist joints. The left panel shows the anterior view, and the right panel shows the posterior view. Figure 3. Bones of the Wrist and Hand. The eight carpal bones form the base of the hand. These are arranged into proximal and distal rows of four bones each. The metacarpal bones form the palm of the hand. The thumb and fingers consist of the phalanx bones.[/caption]

The carpal bones form the base of the hand. This can be seen in the radiograph (X-ray image) of the hand that shows the relationships of the hand bones to the skin creases of the hand (see Figure 4). Within the carpal bones, the four proximal bones are united to each other by ligaments to form a unit. Only three of these bones, the scaphoid, lunate, and triquetrum, contribute to the radiocarpal joint. The scaphoid and lunate bones articulate directly with the distal end of the radius, whereas the triquetrum bone articulates with a fibrocartilaginous pad that spans the radius and styloid process of the ulna. The distal end of the ulna thus does not directly articulate with any of the carpal bones.

The four distal carpal bones are also held together as a group by ligaments. The proximal and distal rows of carpal bones articulate with each other to form the midcarpal joint (see Figure 4). Together, the radiocarpal and midcarpal joints are responsible for all movements of the hand at the wrist. The distal carpal bones also articulate with the metacarpal bones of the hand.

[caption id="" align="aligncenter" width="480"]This image shows a radiograph of a human hand. Figure 4. Bones of the Hand. This radiograph shows the position of the bones within the hand. Note the carpal bones that form the base of the hand. (credit: modification of work by Trace Meek)[/caption]

In the articulated hand, the carpal bones form a U-shaped grouping. A strong ligament called the flexor retinaculum spans the top of this U-shaped area to maintain this grouping of the carpal bones. The flexor retinaculum is attached laterally to the trapezium and scaphoid bones, and medially to the hamate and pisiform bones. Together, the carpal bones and the flexor retinaculum form a passageway called the carpal tunnel, with the carpal bones forming the walls and floor, and the flexor retinaculum forming the roof of this space (Figure 5). The tendons of nine muscles of the anterior forearm and an important nerve pass through this narrow tunnel to enter the hand. Overuse of the muscle tendons or wrist injury can produce inflammation and swelling within this space. This produces compression of the nerve, resulting in carpal tunnel syndrome, which is characterized by pain or numbness, and muscle weakness in those areas of the hand supplied by this nerve.

[caption id="" align="aligncenter" width="350"]This figure shows a hand and a cross-section image of the nerves at the wrist. Figure 5. Carpal Tunnel. The carpal tunnel is the passageway by which nine muscle tendons and a major nerve enter the hand from the anterior forearm. The walls and floor of the carpal tunnel are formed by the U-shaped grouping of the carpal bones, and the roof is formed by the flexor retinaculum, a strong ligament that anteriorly unites the bones.[/caption]

Metacarpal Bones

The palm of the hand contains five elongated metacarpal bones. These bones lie between the carpal bones of the wrist and the bones of the fingers and thumb (see Figure 3). The proximal end of each metacarpal bone articulates with one of the distal carpal bones. Each of these articulations is a carpometacarpal joint (see Figure 4). The expanded distal end of each metacarpal bone articulates at the metacarpophalangeal joint with the proximal phalanx bone of the thumb or one of the fingers. The distal end also forms the knuckles of the hand, at the base of the fingers. The metacarpal bones are numbered 1–5, beginning at the thumb.

The first metacarpal bone, at the base of the thumb, is separated from the other metacarpal bones. This allows it a freedom of motion that is independent of the other metacarpal bones, which is very important for thumb mobility. The remaining metacarpal bones are united together to form the palm of the hand. The second and third metacarpal bones are firmly anchored in place and are immobile. However, the fourth and fifth metacarpal bones have limited anterior-posterior mobility, a motion that is greater for the fifth bone. This mobility is important during power gripping with the hand (Figure 6). The anterior movement of these bones, particularly the fifth metacarpal bone, increases the strength of contact for the medial hand during gripping actions.
[caption id="" align="aligncenter" width="480"]The left panel of this figure shows a hand gripping a motorcycle handle loosely, and the right panel shows a hand gripping a motorcycle handle tightly. Figure 6. Hand During Gripping. During tight gripping—compare (b) to (a)—the fourth and, particularly, the fifth metatarsal bones are pulled anteriorly. This increases the contact between the object and the medial side of the hand, thus improving the firmness of the grip.[/caption]

Phalanx Bones

The fingers and thumb contain 14 bones, each of which is called a phalanx bone (plural = phalanges), named after the ancient Greek phalanx (a rectangular block of soldiers). The thumb (pollex) is digit number 1 and has two phalanges, a proximal phalanx, and a distal phalanx bone (see Figure 3). Digits 2 (index finger) through 5 (little finger) have three phalanges each, called the proximal, middle, and distal phalanx bones. An interphalangeal joint is one of the articulations between adjacent phalanges of the digits (see Figure 4).
]]> 2095 2082 3 0 8.3 The Pelvic Girdle and Pelvis https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2099 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2099

Learning Objectives

By the end of this section, you will be able to:
  • Define the pelvic girdle and describe the bones and ligaments of the pelvis
  • Explain the three regions of the hip bone and identify their bony landmarks
  • Describe the openings of the pelvis and the boundaries of the greater and lesser pelvis
The pelvic girdle (hip girdle) is formed by a single bone, the hip bone or coxal bone (coxal = “hip”), which serves as the attachment point for each lower limb. Each hip bone, in turn, is firmly joined to the axial skeleton via its attachment to the sacrum of the vertebral column. The right and left hip bones also converge anteriorly to attach to each other. The bony pelvis is the entire structure formed by the two hip bones, the sacrum, and, attached inferiorly to the sacrum, the coccyx (Figure 1).

Unlike the bones of the pectoral girdle, which are highly mobile to enhance the range of upper limb movements, the bones of the pelvis are strongly united to each other to form a largely immobile, weight-bearing structure. This is important for stability because it enables the weight of the body to be easily transferred laterally from the vertebral column, through the pelvic girdle and hip joints, and into either lower limb whenever the other limb is not bearing weight. Thus, the immobility of the pelvis provides a strong foundation for the upper body as it rests on top of the mobile lower limbs.

[caption id="" align="aligncenter" width="480"]This figure shows the bone of the pelvis. Figure 1. Pelvis. The pelvic girdle is formed by a single hip bone. The hip bone attaches the lower limb to the axial skeleton through its articulation with the sacrum. The right and left hip bones, plus the sacrum and the coccyx, together form the pelvis.[/caption]

Hip Bone

The hip bone, or coxal bone, forms the pelvic girdle portion of the pelvis. The paired hip bones are the large, curved bones that form the lateral and anterior aspects of the pelvis. Each adult hip bone is formed by three separate bones that fuse together during the late teenage years. These bony components are the ilium, ischium, and pubis (Figure 2). These names are retained and used to define the three regions of the adult hip bone.

[caption id="" align="aligncenter" width="480"]This figure shows the right hip bone. The left panel shows the lateral view, and the right panel shows the medial view. Figure 2. The Hip Bone. The adult hip bone consists of three regions. The ilium forms the large, fan-shaped superior portion, the ischium forms the posteroinferior portion, and the pubis forms the anteromedial portion.[/caption]

The ilium is the fan-like, superior region that forms the largest part of the hip bone. It is firmly united to the sacrum at the largely immobile sacroiliac joint (see Figure 1). The ischium forms the posteroinferior region of each hip bone. It supports the body when sitting. The pubis forms the anterior portion of the hip bone. The pubis curves medially, where it joins to the pubis of the opposite hip bone at a specialized joint called the pubic symphysis.

Ilium

When you place your hands on your waist, you can feel the arching, superior margin of the ilium along your waistline (see Figure 2). This curved, superior margin of the ilium is the iliac crest. The rounded, anterior termination of the iliac crest is the anterior superior iliac spine. This important bony landmark can be felt at your anterolateral hip. Inferior to the anterior superior iliac spine is a rounded protuberance called the anterior inferior iliac spine. Both of these iliac spines serve as attachment points for muscles of the thigh. Posteriorly, the iliac crest curves downward to terminate as the posterior superior iliac spine. Muscles and ligaments surround but do not cover this bony landmark, thus sometimes producing a depression seen as a “dimple” located on the lower back. More inferiorly is the posterior inferior iliac spine. This is located at the inferior end of a large, roughened area called the auricular surface of the ilium. The auricular surface articulates with the auricular surface of the sacrum to form the sacroiliac joint. Both the posterior superior and posterior inferior iliac spines serve as attachment points for the muscles and very strong ligaments that support the sacroiliac joint.

The shallow depression located on the anteromedial (internal) surface of the upper ilium is called the iliac fossa. The inferior margin of this space is formed by the arcuate line of the ilium, the ridge formed by the pronounced change in curvature between the upper and lower portions of the ilium. The large, inverted U-shaped indentation located on the posterior margin of the lower ilium is called the greater sciatic notch.

Ischium

The ischium forms the posterolateral portion of the hip bone (see Figure 2). The large, roughened area of the inferior ischium is the ischial tuberosity. This serves as the attachment for the posterior thigh muscles and also carries the weight of the body when sitting. You can feel the ischial tuberosity if you wiggle your pelvis against the seat of a chair. Projecting superiorly and anteriorly from the ischial tuberosity is a narrow segment of bone called the ischial ramus. The slightly curved posterior margin of the ischium above the ischial tuberosity is the lesser sciatic notch. The bony projection separating the lesser sciatic notch and greater sciatic notch is the ischial spine.

Pubis

The pubis forms the anterior portion of the hip bone (see Figure 2). The enlarged medial portion of the pubis is the pubic body. Located superiorly on the pubic body is a small bump called the pubic tubercle. The superior pubic ramus is the segment of bone that passes laterally from the pubic body to join the ilium. The narrow ridge running along the superior margin of the superior pubic ramus is the pectineal line of the pubis.

The pubic body is joined to the pubic body of the opposite hip bone by the pubic symphysis. Extending downward and laterally from the body is the inferior pubic ramus. The pubic arch is the bony structure formed by the pubic symphysis, and the bodies and inferior pubic rami of the adjacent pubic bones. The inferior pubic ramus extends downward to join the ischial ramus. Together, these form the single ischiopubic ramus, which extends from the pubic body to the ischial tuberosity. The inverted V-shape formed as the ischiopubic rami from both sides come together at the pubic symphysis is called the subpubic angle.

Pelvis

The pelvis consists of four bones: the right and left hip bones, the sacrum, and the coccyx (see Figure 1). The pelvis has several important functions. Its primary role is to support the weight of the upper body when sitting and to transfer this weight to the lower limbs when standing. It serves as an attachment point for trunk and lower limb muscles, and also protects the internal pelvic organs. When standing in the anatomical position, the pelvis is tilted anteriorly. In this position, the anterior superior iliac spines and the pubic tubercles lie in the same vertical plane, and the anterior (internal) surface of the sacrum faces forward and downward.

The three areas of each hip bone, the ilium, pubis, and ischium, converge centrally to form a deep, cup-shaped cavity called the acetabulum. This is located on the lateral side of the hip bone and is part of the hip joint. The large opening in the anteroinferior hip bone between the ischium and pubis is the obturator foramen. This space is largely filled in by a layer of connective tissue and serves for the attachment of muscles on both its internal and external surfaces.

The space enclosed by the bony pelvis is divided into two regions (Figure 4). The broad, superior region, defined laterally by the large, fan-like portion of the upper hip bone, is called the greater pelvis (greater pelvic cavity; false pelvis). This broad area is occupied by portions of the small and large intestines, and because it is more closely associated with the abdominal cavity, it is sometimes referred to as the false pelvis. More inferiorly, the narrow, rounded space of the lesser pelvis (lesser pelvic cavity; true pelvis) contains the bladder and other pelvic organs, and thus is also known as the true pelvis. The pelvic brim (also known as the pelvic inlet) forms the superior margin of the lesser pelvis, separating it from the greater pelvis. The pelvic brim is defined by a line formed by the upper margin of the pubic symphysis anteriorly, and the pectineal line of the pubis, the arcuate line of the ilium, and the sacral promontory (the anterior margin of the superior sacrum) posteriorly. The inferior limit of the lesser pelvic cavity is called the pelvic outlet. This large opening is defined by the inferior margin of the pubic symphysis anteriorly, and the ischiopubic ramus, the ischial tuberosity, the sacrotuberous ligament, and the inferior tip of the coccyx posteriorly. Because of the anterior tilt of the pelvis, the lesser pelvis is also angled, giving it an anterosuperior (pelvic inlet) to posteroinferior (pelvic outlet) orientation.

[caption id="" align="aligncenter" width="480"]This figure shows the structure of the female pelvic girdle on the left and the male pelvic girdle on the right. Figure 4. Male and Female Pelvis. The female pelvis is adapted for childbirth and is broader, with a larger subpubic angle, a rounder pelvic brim, and a wider and more shallow lesser pelvic cavity than the male pelvis.[/caption]

Comparison of the Female and Male Pelvis

The differences between the adult female and male pelvis relate to function and body size. In general, the bones of the male pelvis are thicker and heavier, adapted for support of the male’s heavier physical build and stronger muscles. The greater sciatic notch of the male hip bone is narrower and deeper than the broader notch of females. Because the female pelvis is adapted for childbirth, it is wider than the male pelvis, as evidenced by the distance between the anterior superior iliac spines (see Figure 4). The ischial tuberosities of females are also farther apart, which increases the size of the pelvic outlet. Because of this increased pelvic width, the subpubic angle is larger in females (greater than 80 degrees) than it is in males (less than 70 degrees). The female sacrum is wider, shorter, and less curved, and the sacral promontory projects less into the pelvic cavity, thus giving the female pelvic inlet (pelvic brim) a more rounded or oval shape compared to males. The lesser pelvic cavity of females is also wider and more shallow than the narrower, deeper, and tapering lesser pelvis of males. Because of the obvious differences between female and male hip bones, this is the one bone of the body that allows for the most accurate sex determination. Table 1 provides an overview of the general differences between the female and male pelvis.

Overview of Differences between the Female and Male Pelvis (Table 1)
Female pelvis Male pelvis
Pelvic weight Bones of the pelvis are lighter and thinner Bones of the pelvis are thicker and heavier
Pelvic inlet shape Pelvic inlet has a round or oval shape Pelvic inlet is heart-shaped
Lesser pelvic cavity shape Lesser pelvic cavity is shorter and wider Lesser pelvic cavity is longer and narrower
Subpubic angle Subpubic angle is greater than 80 degrees Subpubic angle is less than 70 degrees
Pelvic outlet shape Pelvic outlet is rounded and larger Pelvic outlet is smaller
]]> 2099 2082 4 0 8.4 Bones of the Lower Limb https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2105 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2105

Learning Objectives

By the end of this section, you will be able to:
  • Identify the divisions of the lower limb and describe the bones of each region
  • Describe the bones and bony landmarks that articulate at each joint of the lower limb
Like the upper limb, the lower limb is divided into three regions. The thigh is that portion of the lower limb located between the hip joint and knee joint. The leg is specifically the region between the knee joint and the ankle joint. Distal to the ankle is the foot. The lower limb contains 30 bones. These are the femur, patella, tibia, fibula, tarsal bones, metatarsal bones, and phalanges (see Chapter 8.1 Figure 1). The femur is the single bone of the thigh. The patella is the kneecap and articulates with the distal femur. The tibia is the larger, weight-bearing bone located on the medial side of the leg, and the fibula is the thin bone of the lateral leg. The bones of the foot are divided into three groups. The posterior portion of the foot is formed by a group of seven bones, each of which is known as a tarsal bone, whereas the mid-foot contains five elongated bones, each of which is a metatarsal bone. The toes contain 14 small bones, each of which is a phalanx bone of the foot.

Femur

The femur, or thigh bone, is the single bone of the thigh region (Figure 1). It is the longest and strongest bone of the body, and accounts for approximately one-quarter of a person’s total height. The rounded, proximal end is the head of the femur, which articulates with the acetabulum of the hip bone to form the hip joint. The fovea capitis is a minor indentation on the medial side of the femoral head that serves as the site of attachment for the ligament of the head of the femur. This ligament spans the femur and acetabulum, but is weak and provides little support for the hip joint. It does, however, carry an important artery that supplies the head of the femur.

[caption id="" align="aligncenter" width="380"]This diagram shows the bones of the femur and the patella. The left panel shows the anterior view, and the right panel shows the posterior view. Figure 1. Femur and Patella. The femur is the single bone of the thigh region. It articulates superiorly with the hip bone at the hip joint, and inferiorly with the tibia at the knee joint. The patella only articulates with the distal end of the femur.[/caption]

The narrowed region below the head is the neck of the femur. This is a common area for fractures of the femur. The greater trochanter is the large, upward, bony projection located above the base of the neck. Multiple muscles that act across the hip joint attach to the greater trochanter, which, because of its projection from the femur, gives additional leverage to these muscles. The greater trochanter can be felt just under the skin on the lateral side of your upper thigh. The lesser trochanter is a small, bony prominence that lies on the medial aspect of the femur, just below the neck. A single, powerful muscle attaches to the lesser trochanter. Running between the greater and lesser trochanters on the anterior side of the femur is the roughened intertrochanteric line. The trochanters are also connected on the posterior side of the femur by the larger intertrochanteric crest.

The elongated shaft of the femur has a slight anterior bowing or curvature. At its proximal end, the posterior shaft has the gluteal tuberosity, a roughened area extending inferiorly from the greater trochanter. More inferiorly, the gluteal tuberosity becomes continuous with the linea aspera (“rough line”). This is the roughened ridge that passes distally along the posterior side of the mid-femur. Multiple muscles of the hip and thigh regions make long, thin attachments to the femur along the linea aspera.

The distal end of the femur has medial and lateral bony expansions. On the lateral side, the smooth portion that covers the distal and posterior aspects of the lateral expansion is the lateral condyle of the femur. The roughened area on the outer, lateral side of the condyle is the lateral epicondyle of the femur. Similarly, the smooth region of the distal and posterior medial femur is the medial condyle of the femur, and the irregular outer, medial side of this is the medial epicondyle of the femur. The lateral and medial condyles articulate with the tibia to form the knee joint. The epicondyles provide attachment for muscles and supporting ligaments of the knee. The adductor tubercle is a small bump located at the superior margin of the medial epicondyle. Posteriorly, the medial and lateral condyles are separated by a deep depression called the intercondylar fossa. Anteriorly, the smooth surfaces of the condyles join together to form a wide groove called the patellar surface, which provides for articulation with the patella bone. The combination of the medial and lateral condyles with the patellar surface gives the distal end of the femur a horseshoe (U) shape.

Patella

The patella (kneecap) is largest sesamoid bone of the body (see Figure 1). A sesamoid bone is a bone that is incorporated into the tendon of a muscle where that tendon crosses a joint. The sesamoid bone articulates with the underlying bones to prevent damage to the muscle tendon due to rubbing against the bones during movements of the joint. The patella is found in the tendon of the quadriceps femoris muscle, the large muscle of the anterior thigh that passes across the anterior knee to attach to the tibia. The patella articulates with the patellar surface of the femur and thus prevents rubbing of the muscle tendon against the distal femur. The patella also lifts the tendon away from the knee joint, which increases the leverage power of the quadriceps femoris muscle as it acts across the knee. The patella does not articulate with the tibia.

[caption id="" align="aligncenter" width="225"]This figure shows the anterior view of the femur. Figure 2. The Q-Angle. The Q-angle is a measure of the amount of lateral deviation of the femur from the vertical line of the tibia. Adult females have a larger Q-angle due to their wider pelvis than adult males.[/caption]

Tibia

The tibia (shin bone) is the medial bone of the leg and is larger than the fibula, with which it is paired (Figure 3). The tibia is the main weight-bearing bone of the lower leg and the second longest bone of the body, after the femur. The medial side of the tibia is located immediately under the skin, allowing it to be easily palpated down the entire length of the medial leg.

[caption id="" align="aligncenter" width="400"]This image shows the structure of the tibia and the fibula. The left panel shows the anterior view, and the right panel shows the posterior view. Figure 3. Tibia and Fibula. The tibia is the larger, weight-bearing bone located on the medial side of the leg. The fibula is the slender bone of the lateral side of the leg and does not bear weight.[/caption]

The proximal end of the tibia is greatly expanded. The two sides of this expansion form the medial condyle of the tibia and the lateral condyle of the tibia. The tibia does not have epicondyles. The top surface of each condyle is smooth and flattened. These areas articulate with the medial and lateral condyles of the femur to form the knee joint. Between the articulating surfaces of the tibial condyles is the intercondylar eminence, an irregular, elevated area that serves as the inferior attachment point for two supporting ligaments of the knee.

The tibial tuberosity is an elevated area on the anterior side of the tibia, near its proximal end. It is the final site of attachment for the muscle tendon associated with the patella. More inferiorly, the shaft of the tibia becomes triangular in shape. The anterior apex of this triangle forms the anterior border of the tibia, which begins at the tibial tuberosity and runs inferiorly along the length of the tibia. Both the anterior border and the medial side of the triangular shaft are located immediately under the skin and can be easily palpated along the entire length of the tibia. A small ridge running down the lateral side of the tibial shaft is the interosseous border of the tibia. This is for the attachment of the interosseous membrane of the leg, the sheet of dense connective tissue that unites the tibia and fibula bones. Located on the posterior side of the tibia is the soleal line, a diagonally running, roughened ridge that begins below the base of the lateral condyle, and runs down and medially across the proximal third of the posterior tibia. Muscles of the posterior leg attach to this line.

The large expansion found on the medial side of the distal tibia is the medial malleolus (“little hammer”). This forms the large bony bump found on the medial side of the ankle region. Both the smooth surface on the inside of the medial malleolus and the smooth area at the distal end of the tibia articulate with the talus bone of the foot as part of the ankle joint. On the lateral side of the distal tibia is a wide groove called the fibular notch. This area articulates with the distal end of the fibula, forming the distal tibiofibular joint.

Fibula

The fibula is the slender bone located on the lateral side of the leg (see Figure 3). The fibula does not bear weight. It serves primarily for muscle attachments and thus is largely surrounded by muscles. Only the proximal and distal ends of the fibula can be palpated.

The head of the fibula is the small, knob-like, proximal end of the fibula. It articulates with the inferior aspect of the lateral tibial condyle, forming the proximal tibiofibular joint. The thin shaft of the fibula has the interosseous border of the fibula, a narrow ridge running down its medial side for the attachment of the interosseous membrane that spans the fibula and tibia. The distal end of the fibula forms the lateral malleolus, which forms the easily palpated bony bump on the lateral side of the ankle. The deep (medial) side of the lateral malleolus articulates with the talus bone of the foot as part of the ankle joint. The distal fibula also articulates with the fibular notch of the tibia.

Tarsal Bones

The posterior half of the foot is formed by seven tarsal bones (Figure 4). The most superior bone is the talus. This has a relatively square-shaped, upper surface that articulates with the tibia and fibula to form the ankle joint. Three areas of articulation form the ankle joint: The superomedial surface of the talus bone articulates with the medial malleolus of the tibia, the top of the talus articulates with the distal end of the tibia, and the lateral side of the talus articulates with the lateral malleolus of the fibula. Inferiorly, the talus articulates with the calcaneus (heel bone), the largest bone of the foot, which forms the heel. Body weight is transferred from the tibia to the talus to the calcaneus, which rests on the ground. The medial calcaneus has a prominent bony extension called the sustentaculum tali (“support for the talus”) that supports the medial side of the talus bone.

[caption id="" align="aligncenter" width="600"]This figure shows the bones of the foot. The left panel shows the superior view, the top right panel shows the medial view, and the bottom right panel shows the lateral view. Figure 4. Bones of the Foot. The bones of the foot are divided into three groups. The posterior foot is formed by the seven tarsal bones. The mid-foot has the five metatarsal bones. The toes contain the phalanges.[/caption]
[caption id="" align="alignright" width="120"]QR Code representing a URL Use this tutorial to review the bones of the foot.[/caption] The cuboid bone articulates with the anterior end of the calcaneus bone. The cuboid has a deep groove running across its inferior surface, which provides passage for a muscle tendon. The talus bone articulates anteriorly with the navicular bone, which in turn articulates anteriorly with the three cuneiform (“wedge-shaped”) bones. These bones are the medial cuneiform, the intermediate cuneiform, and the lateral cuneiform. Each of these bones has a broad superior surface and a narrow inferior surface, which together produce the transverse (medial-lateral) curvature of the foot. The navicular and lateral cuneiform bones also articulate with the medial side of the cuboid bone.

Metatarsal Bones

The anterior half of the foot is formed by the five metatarsal bones, which are located between the tarsal bones of the posterior foot and the phalanges of the toes (see Figure 4). These elongated bones are numbered 1–5, starting with the medial side of the foot. The first metatarsal bone is shorter and thicker than the others. The second metatarsal is the longest. The base of the metatarsal bone is the proximal end of each metatarsal bone. These articulate with the cuboid or cuneiform bones. The base of the fifth metatarsal has a large, lateral expansion that provides for muscle attachments. This expanded base of the fifth metatarsal can be felt as a bony bump at the midpoint along the lateral border of the foot. The expanded distal end of each metatarsal is the head of the metatarsal bone. Each metatarsal bone articulates with the proximal phalanx of a toe to form a metatarsophalangeal joint. The heads of the metatarsal bones also rest on the ground and form the ball (anterior end) of the foot.

Phalanges

The toes contain a total of 14 phalanx bones (phalanges), arranged in a similar manner as the phalanges of the fingers (see Figure 4). The toes are numbered 1–5, starting with the big toe (hallux). The big toe has two phalanx bones, the proximal and distal phalanges. The remaining toes all have proximal, middle, and distal phalanges. A joint between adjacent phalanx bones is called an interphalangeal joint.

]]> 2105 2082 5 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2110 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2110 [caption id="" align="aligncenter" width="600"]This picture shows a girl kayaking in the ocean. Figure 1. Girl Kayaking. Without joints, body movements would be impossible. (credit: Graham Richardson/flickr.com)[/caption]

Chapter Objectives

After this chapter, you will be able to:
  • Discuss both functional and structural classifications for body joints
  • Describe the characteristic features for fibrous, cartilaginous, and synovial joints and give examples of each
  • Define and identify the different body movements
  • Discuss the structure of specific body joints and the movements allowed by each
  • Explain the development of body joints
The adult human body has 206 bones, and with the exception of the hyoid bone in the neck, each bone is connected to at least one other bone. Joints are the location where bones come together. Many joints allow for movement between the bones. At these joints, the articulating surfaces of the adjacent bones can move smoothly against each other. However, the bones of other joints may be joined to each other by connective tissue or cartilage. These joints are designed for stability and provide for little or no movement. Importantly, joint stability and movement are related to each other. This means that stable joints allow for little or no mobility between the adjacent bones. Conversely, joints that provide the most movement between bones are the least stable. Understanding the relationship between joint structure and function will help to explain why particular types of joints are found in certain areas of the body. The articulating surfaces of bones at stable types of joints, with little or no mobility, are strongly united to each other. For example, most of the joints of the skull are held together by fibrous connective tissue and do not allow for movement between the adjacent bones. This lack of mobility is important, because the skull bones serve to protect the brain. Similarly, other joints united by fibrous connective tissue allow for very little movement, which provides stability and weight-bearing support for the body. For example, the tibia and fibula of the leg are tightly united to give stability to the body when standing. At other joints, the bones are held together by cartilage, which permits limited movements between the bones. Thus, the joints of the vertebral column only allow for small movements between adjacent vertebrae, but when added together, these movements provide the flexibility that allows your body to twist, or bend to the front, back, or side. In contrast, at joints that allow for wide ranges of motion, the articulating surfaces of the bones are not directly united to each other. Instead, these surfaces are enclosed within a space filled with lubricating fluid, which allows the bones to move smoothly against each other. These joints provide greater mobility, but since the bones are free to move in relation to each other, the joint is less stable. Most of the joints between the bones of the appendicular skeleton are this freely moveable type of joint. These joints allow the muscles of the body to pull on a bone and thereby produce movement of that body region. Your ability to kick a soccer ball, pick up a fork, and dance the tango depend on mobility at these types of joints.]]> 2110 2108 1 0 9.3 Cartilaginous Joints https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2119 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2119

Learning Objectives

By the end of this section, you will be able to:
  • Describe the structural features of cartilaginous joints
  • Distinguish between a synchondrosis and symphysis
  • Give an example of each type of cartilaginous joint
As the name indicates, at a cartilaginous joint, the adjacent bones are united by cartilage, a tough but flexible type of connective tissue. These types of joints lack a joint cavity and involve bones that are joined together by either hyaline cartilage or fibrocartilage (Figure 1). There are two types of cartilaginous joints. A synchondrosis is a cartilaginous joint where the bones are joined by hyaline cartilage. Also classified as a synchondrosis are places where bone is united to a cartilage structure, such as between the anterior end of a rib and the costal cartilage of the thoracic cage. The second type of cartilaginous joint is a symphysis, where the bones are joined by fibrocartilage.
[caption id="" align="aligncenter" width="520"]This figure shows the cartilaginous joints. The left panel shows a hyaline cartilage joint, and the right panel shows the fibrocartilaginous joint of the pubic symphisis. Figure 1. Cartiliginous Joints. At cartilaginous joints, bones are united by hyaline cartilage to form a synchondrosis or by fibrocartilage to form a symphysis. (a) The hyaline cartilage of the epiphyseal plate (growth plate) forms a synchondrosis that unites the shaft (diaphysis) and end (epiphysis) of a long bone and allows the bone to grow in length. (b) The pubic portions of the right and left hip bones of the pelvis are joined together by fibrocartilage, forming the pubic symphysis.[/caption]

Synchondrosis

A synchondrosis (“joined by cartilage”) is a cartilaginous joint where bones are joined together by hyaline cartilage, or where bone is united to hyaline cartilage. A synchondrosis may be temporary or permanent. A temporary synchondrosis is the epiphyseal plate (growth plate) of a growing long bone. The epiphyseal plate is the region of growing hyaline cartilage that unites the diaphysis (shaft) of the bone to the epiphysis (end of the bone). Bone lengthening involves growth of the epiphyseal plate cartilage and its replacement by bone, which adds to the diaphysis. For many years during childhood growth, the rates of cartilage growth and bone formation are equal and thus the epiphyseal plate does not change in overall thickness as the bone lengthens. During the late teens and early 20s, growth of the cartilage slows and eventually stops. The epiphyseal plate is then completely replaced by bone, and the diaphysis and epiphysis portions of the bone fuse together to form a single adult bone. This fusion of the diaphysis and epiphysis is a synostosis. Once this occurs, bone lengthening ceases. For this reason, the epiphyseal plate is considered to be a temporary synchondrosis. Because cartilage is softer than bone tissue, injury to a growing long bone can damage the epiphyseal plate cartilage, thus stopping bone growth and preventing additional bone lengthening.

Growing layers of cartilage also form synchondroses that join together the ilium, ischium, and pubic portions of the hip bone during childhood and adolescence. When body growth stops, the cartilage disappears and is replaced by bone, forming synostoses and fusing the bony components together into the single hip bone of the adult. Similarly, synostoses unite the sacral vertebrae that fuse together to form the adult sacrum.

Visit this website to view a radiograph (X-ray image) of a child’s hand and wrist. The growing bones of child have an epiphyseal plate that forms a synchondrosis between the shaft and end of a long bone. Being less dense than bone, the area of epiphyseal cartilage is seen on this radiograph as the dark epiphyseal gaps located near the ends of the long bones, including the radius, ulna, metacarpal, and phalanx bones. Which of the bones in this image do not show an epiphyseal plate (epiphyseal gap)?

Examples of permanent synchondroses are found in the thoracic cage. One example is the first sternocostal joint, where the first rib is anchored to the manubrium by its costal cartilage. (The articulations of the remaining costal cartilages to the sternum are all synovial joints.) Additional synchondroses are formed where the anterior end of the other 11 ribs is joined to its costal cartilage. Unlike the temporary synchondroses of the epiphyseal plate, these permanent synchondroses retain their hyaline cartilage and thus do not ossify with age. Due to the lack of movement between the bone and cartilage, both temporary and permanent synchondroses are functionally classified as a synarthrosis.

Symphysis

A cartilaginous joint where the bones are joined by fibrocartilage is called a symphysis (“growing together”). Fibrocartilage is very strong because it contains numerous bundles of thick collagen fibers, thus giving it a much greater ability to resist pulling and bending forces when compared with hyaline cartilage. This gives symphyses the ability to strongly unite the adjacent bones, but can still allow for limited movement to occur. Thus, a symphysis is functionally classified as an amphiarthrosis.

The gap separating the bones at a symphysis may be narrow or wide. Examples in which the gap between the bones is narrow include the pubic symphysis and the manubriosternal joint. At the pubic symphysis, the pubic portions of the right and left hip bones of the pelvis are joined together by fibrocartilage across a narrow gap. Similarly, at the manubriosternal joint, fibrocartilage unites the manubrium and body portions of the sternum.

The intervertebral symphysis is a wide symphysis located between the bodies of adjacent vertebrae of the vertebral column. Here a thick pad of fibrocartilage called an intervertebral disc strongly unites the adjacent vertebrae by filling the gap between them. The width of the intervertebral symphysis is important because it allows for small movements between the adjacent vertebrae. In addition, the thick intervertebral disc provides cushioning between the vertebrae, which is important when carrying heavy objects or during high-impact activities such as running or jumping.

]]> 2119 2108 4 0 9.4 Synovial Joints https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2126 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2126

Learning Objectives

By the end of this section, you will be able to:
  • Describe the structural features of a synovial joint
  • Discuss the function of additional structures associated with synovial joints
  • List the six types of synovial joints and give an example of each
Synovial joints are the most common type of joint in the body (Figure 1). A key structural characteristic for a synovial joint that is not seen at fibrous or cartilaginous joints is the presence of a joint cavity. This fluid-filled space is the site at which the articulating surfaces of the bones contact each other. Also unlike fibrous or cartilaginous joints, the articulating bone surfaces at a synovial joint are not directly connected to each other with fibrous connective tissue or cartilage. This gives the bones of a synovial joint the ability to move smoothly against each other, allowing for increased joint mobility.
[caption id="" align="aligncenter" width="380"]This figure shows a synovial joint. The cavity between two bones contains the synovial fluid which lubricates the two joints. Figure 1. Synovial Joints. Synovial joints allow for smooth movements between the adjacent bones. The joint is surrounded by an articular capsule that defines a joint cavity filled with synovial fluid. The articulating surfaces of the bones are covered by a thin layer of articular cartilage. Ligaments support the joint by holding the bones together and resisting excess or abnormal joint motions.[/caption]

Structural Features of Synovial Joints

Synovial joints are characterized by the presence of a joint cavity. The walls of this space are formed by the articular capsule, a fibrous connective tissue structure that is attached to each bone just outside the area of the bone’s articulating surface. The bones of the joint articulate with each other within the joint cavity.

Friction between the bones at a synovial joint is prevented by the presence of the articular cartilage, a thin layer of hyaline cartilage that covers the entire articulating surface of each bone. However, unlike at a cartilaginous joint, the articular cartilages of each bone are not continuous with each other. Instead, the articular cartilage acts like a Teflon® coating over the bone surface, allowing the articulating bones to move smoothly against each other without damaging the underlying bone tissue. Lining the inner surface of the articular capsule is a thin synovial membrane. The cells of this membrane secrete synovial fluid (synovia = “a thick fluid”), a thick, slimy fluid that provides lubrication to further reduce friction between the bones of the joint. This fluid also provides nourishment to the articular cartilage, which does not contain blood vessels. The ability of the bones to move smoothly against each other within the joint cavity, and the freedom of joint movement this provides, means that each synovial joint is functionally classified as a diarthrosis.

Outside of their articulating surfaces, the bones are connected together by ligaments, which are strong bands of fibrous connective tissue. These strengthen and support the joint by anchoring the bones together and preventing their separation. Ligaments allow for normal movements at a joint, but limit the range of these motions, thus preventing excessive or abnormal joint movements. Ligaments are classified based on their relationship to the fibrous articular capsule. An extrinsic ligament is located outside of the articular capsule, an intrinsic ligament is fused to or incorporated into the wall of the articular capsule, and an intracapsular ligament is located inside of the articular capsule.

At many synovial joints, additional support is provided by the muscles and their tendons that act across the joint. A tendon is the dense connective tissue structure that attaches a muscle to bone. As forces acting on a joint increase, the body will automatically increase the overall strength of contraction of the muscles crossing that joint, thus allowing the muscle and its tendon to serve as a “dynamic ligament” to resist forces and support the joint. This type of indirect support by muscles is very important at the shoulder joint, for example, where the ligaments are relatively weak.

Additional Structures Associated with Synovial Joints

A few synovial joints of the body have a fibrocartilage structure located between the articulating bones. This is called an articular disc, which is generally small and oval-shaped, or a meniscus, which is larger and C-shaped. These structures can serve several functions, depending on the specific joint. In some places, an articular disc may act to strongly unite the bones of the joint to each other. Examples of this include the articular discs found at the sternoclavicular joint or between the distal ends of the radius and ulna bones. At other synovial joints, the disc can provide shock absorption and cushioning between the bones, which is the function of each meniscus within the knee joint. Finally, an articular disc can serve to smooth the movements between the articulating bones, as seen at the temporomandibular joint. Some synovial joints also have a fat pad, which can serve as a cushion between the bones.

Additional structures located outside of a synovial joint serve to prevent friction between the bones of the joint and the overlying muscle tendons or skin. A bursa (plural = bursae) is a thin connective tissue sac filled with lubricating liquid. They are located in regions where skin, ligaments, muscles, or muscle tendons can rub against each other, usually near a body joint (Figure 2). Bursae reduce friction by separating the adjacent structures, preventing them from rubbing directly against each other. Bursae are classified by their location. A subcutaneous bursa is located between the skin and an underlying bone. It allows skin to move smoothly over the bone. Examples include the prepatellar bursa located over the kneecap and the olecranon bursa at the tip of the elbow. A submuscular bursa is found between a muscle and an underlying bone, or between adjacent muscles. These prevent rubbing of the muscle during movements. A large submuscular bursa, the trochanteric bursa, is found at the lateral hip, between the greater trochanter of the femur and the overlying gluteus maximus muscle. A subtendinous bursa is found between a tendon and a bone. Examples include the subacromial bursa that protects the tendon of shoulder muscle as it passes under the acromion of the scapula, and the suprapatellar bursa that separates the tendon of the large anterior thigh muscle from the distal femur just above the knee.

[caption id="" align="aligncenter" width="380"]This diagram shows the location of the bursae which are fluid filled sacs in a bone joint. The major parts of the joint are labeled. Figure 2. Bursae. Bursae are fluid-filled sacs that serve to prevent friction between skin, muscle, or tendon and an underlying bone. Three major bursae and a fat pad are part of the complex joint that unites the femur and tibia of the leg.[/caption]

A tendon sheath is similar in structure to a bursa, but smaller. It is a connective tissue sac that surrounds a muscle tendon at places where the tendon crosses a joint. It contains a lubricating fluid that allows for smooth motions of the tendon during muscle contraction and joint movements.

Bursitis is the inflammation of a bursa near a joint. This will cause pain, swelling, or tenderness of the bursa and surrounding area, and may also result in joint stiffness. Bursitis is most commonly associated with the bursae found at or near the shoulder, hip, knee, or elbow joints. At the shoulder, subacromial bursitis may occur in the bursa that separates the acromion of the scapula from the tendon of a shoulder muscle as it passes deep to the acromion. In the hip region, trochanteric bursitis can occur in the bursa that overlies the greater trochanter of the femur, just below the lateral side of the hip. Ischial bursitis occurs in the bursa that separates the skin from the ischial tuberosity of the pelvis, the bony structure that is weight bearing when sitting. At the knee, inflammation and swelling of the bursa located between the skin and patella bone is prepatellar bursitis (“housemaid’s knee”), a condition more commonly seen today in roofers or floor and carpet installers who do not use knee pads. At the elbow, olecranon bursitis is inflammation of the bursa between the skin and olecranon process of the ulna. The olecranon forms the bony tip of the elbow, and bursitis here is also known as “student’s elbow.”

Bursitis can be either acute (lasting only a few days) or chronic. It can arise from muscle overuse, trauma, excessive or prolonged pressure on the skin, rheumatoid arthritis, gout, or infection of the joint. Repeated acute episodes of bursitis can result in a chronic condition. Treatments for the disorder include antibiotics if the bursitis is caused by an infection, or anti-inflammatory agents, such as nonsteroidal anti-inflammatory drugs (NSAIDs) or corticosteroids if the bursitis is due to trauma or overuse. Chronic bursitis may require that fluid be drained, but additional surgery is usually not required.

Types of Synovial Joints

Synovial joints are subdivided based on the shapes of the articulating surfaces of the bones that form each joint. The six types of synovial joints are pivot, hinge, condyloid, saddle, plane, and ball-and socket-joints (Figure 3).
[caption id="" align="aligncenter" width="600"]This composite image shows the different types of synovial joints in the body. In the center of the figure is a skeleton, and call outs from each joint show their names and locations. Figure 3. Types of Synovial Joints. The six types of synovial joints allow the body to move in a variety of ways. (a) Pivot joints allow for rotation around an axis, such as between the first and second cervical vertebrae, which allows for side-to-side rotation of the head. (b) The hinge joint of the elbow works like a door hinge. (c) The articulation between the trapezium carpal bone and the first metacarpal bone at the base of the thumb is a saddle joint. (d) Plane joints, such as those between the tarsal bones of the foot, allow for limited gliding movements between bones. (e) The radiocarpal joint of the wrist is a condyloid joint. (f) The hip and shoulder joints are the only ball-and-socket joints of the body.[/caption]

Pivot Joint

At a pivot joint, a rounded portion of a bone is enclosed within a ring formed partially by the articulation with another bone and partially by a ligament (see Figure 3a). The bone rotates within this ring. Since the rotation is around a single axis, pivot joints are functionally classified as a uniaxial diarthrosis type of joint. An example of a pivot joint is the atlantoaxial joint, found between the C1 (atlas) and C2 (axis) vertebrae. Here, the upward projecting dens of the axis articulates with the inner aspect of the atlas, where it is held in place by a ligament. Rotation at this joint allows you to turn your head from side to side. A second pivot joint is found at the proximal radioulnar joint. Here, the head of the radius is largely encircled by a ligament that holds it in place as it articulates with the radial notch of the ulna. Rotation of the radius allows for forearm movements.

Hinge Joint

In a hinge joint, the convex end of one bone articulates with the concave end of the adjoining bone (see Figure 3b). This type of joint allows only for bending and straightening motions along a single axis, and thus hinge joints are functionally classified as uniaxial joints. A good example is the elbow joint, with the articulation between the trochlea of the humerus and the trochlear notch of the ulna. Other hinge joints of the body include the knee, ankle, and interphalangeal joints between the phalanx bones of the fingers and toes.

Condyloid Joint

At a condyloid joint (ellipsoid joint), the shallow depression at the end of one bone articulates with a rounded structure from an adjacent bone or bones (see Figure 3e). The knuckle (metacarpophalangeal) joints of the hand between the distal end of a metacarpal bone and the proximal phalanx bone are condyloid joints. Another example is the radiocarpal joint of the wrist, between the shallow depression at the distal end of the radius bone and the rounded scaphoid, lunate, and triquetrum carpal bones. In this case, the articulation area has a more oval (elliptical) shape. Functionally, condyloid joints are biaxial joints that allow for two planes of movement. One movement involves the bending and straightening of the fingers or the anterior-posterior movements of the hand. The second movement is a side-to-side movement, which allows you to spread your fingers apart and bring them together, or to move your hand in a medial-going or lateral-going direction.

Saddle Joint

At a saddle joint, both of the articulating surfaces for the bones have a saddle shape, which is concave in one direction and convex in the other (see Figure 3c). This allows the two bones to fit together like a rider sitting on a saddle. Saddle joints are functionally classified as biaxial joints. The primary example is the first carpometacarpal joint, between the trapezium (a carpal bone) and the first metacarpal bone at the base of the thumb. This joint provides the thumb the ability to move away from the palm of the hand along two planes. Thus, the thumb can move within the same plane as the palm of the hand, or it can jut out anteriorly, perpendicular to the palm. This movement of the first carpometacarpal joint is what gives humans their distinctive “opposable” thumbs. The sternoclavicular joint is also classified as a saddle joint.

Plane Joint

At a plane joint (gliding joint), the articulating surfaces of the bones are flat or slightly curved and of approximately the same size, which allows the bones to slide against each other (see Figure 3d). The motion at this type of joint is usually small and tightly constrained by surrounding ligaments. Based only on their shape, plane joints can allow multiple movements, including rotation. Thus plane joints can be functionally classified as a multiaxial joint. However, not all of these movements are available to every plane joint due to limitations placed on it by ligaments or neighboring bones. Thus, depending upon the specific joint of the body, a plane joint may exhibit only a single type of movement or several movements. Plane joints are found between the carpal bones (intercarpal joints) of the wrist or tarsal bones (intertarsal joints) of the foot, between the clavicle and acromion of the scapula (acromioclavicular joint), and between the superior and inferior articular processes of adjacent vertebrae (zygapophysial joints).

Ball-and-Socket Joint

The joint with the greatest range of motion is the ball-and-socket joint. At these joints, the rounded head of one bone (the ball) fits into the concave articulation (the socket) of the adjacent bone (see Figure 3f). The hip joint and the glenohumeral (shoulder) joint are the only ball-and-socket joints of the body. At the hip joint, the head of the femur articulates with the acetabulum of the hip bone, and at the shoulder joint, the head of the humerus articulates with the glenoid cavity of the scapula.

[caption id="" align="alignright" width="120"]QR Code representing a URL Watch this video to see an animation of synovial joints in action.[/caption]

Ball-and-socket joints are classified functionally as multiaxial joints. The femur and the humerus are able to move in both anterior-posterior and medial-lateral directions and they can also rotate around their long axis. The shallow socket formed by the glenoid cavity allows the shoulder joint an extensive range of motion. In contrast, the deep socket of the acetabulum and the strong supporting ligaments of the hip joint serve to constrain movements of the femur, reflecting the need for stability and weight-bearing ability at the hip.

Arthritis is a common disorder of synovial joints that involves inflammation of the joint. This often results in significant joint pain, along with swelling, stiffness, and reduced joint mobility. There are more than 100 different forms of arthritis. Arthritis may arise from aging, damage to the articular cartilage, autoimmune diseases, bacterial or viral infections, or unknown (probably genetic) causes.

The most common type of arthritis is osteoarthritis, which is associated with aging and “wear and tear” of the articular cartilage (Figure 4). Risk factors that may lead to osteoarthritis later in life include injury to a joint; jobs that involve physical labor; sports with running, twisting, or throwing actions; and being overweight. These factors put stress on the articular cartilage that covers the surfaces of bones at synovial joints, causing the cartilage to gradually become thinner. As the articular cartilage layer wears down, more pressure is placed on the bones. The joint responds by increasing production of the lubricating synovial fluid, but this can lead to swelling of the joint cavity, causing pain and joint stiffness as the articular capsule is stretched. The bone tissue underlying the damaged articular cartilage also responds by thickening, producing irregularities and causing the articulating surface of the bone to become rough or bumpy. Joint movement then results in pain and inflammation. In its early stages, symptoms of osteoarthritis may be reduced by mild activity that “warms up” the joint, but the symptoms may worsen following exercise. In individuals with more advanced osteoarthritis, the affected joints can become more painful and therefore are difficult to use effectively, resulting in increased immobility. There is no cure for osteoarthritis, but several treatments can help alleviate the pain. Treatments may include lifestyle changes, such as weight loss and low-impact exercise, and over-the-counter or prescription medications that help to alleviate the pain and inflammation. For severe cases, joint replacement surgery (arthroplasty) may be required.

Joint replacement is a very invasive procedure, so other treatments are always tried before surgery. However arthroplasty can provide relief from chronic pain and can enhance mobility within a few months following the surgery. This type of surgery involves replacing the articular surfaces of the bones with prosthesis (artificial components). For example, in hip arthroplasty, the worn or damaged parts of the hip joint, including the head and neck of the femur and the acetabulum of the pelvis, are removed and replaced with artificial joint components. The replacement head for the femur consists of a rounded ball attached to the end of a shaft that is inserted inside the diaphysis of the femur. The acetabulum of the pelvis is reshaped and a replacement socket is fitted into its place. The parts, which are always built in advance of the surgery, are sometimes custom made to produce the best possible fit for a patient.

Gout is a form of arthritis that results from the deposition of uric acid crystals within a body joint. Usually only one or a few joints are affected, such as the big toe, knee, or ankle. The attack may only last a few days, but may return to the same or another joint. Gout occurs when the body makes too much uric acid or the kidneys do not properly excrete it. A diet with excessive fructose has been implicated in raising the chances of a susceptible individual developing gout.

Other forms of arthritis are associated with various autoimmune diseases, bacterial infections of the joint, or unknown genetic causes. Autoimmune diseases, including rheumatoid arthritis, scleroderma, or systemic lupus erythematosus, produce arthritis because the immune system of the body attacks the body joints. In rheumatoid arthritis, the joint capsule and synovial membrane become inflamed. As the disease progresses, the articular cartilage is severely damaged or destroyed, resulting in joint deformation, loss of movement, and severe disability. The most commonly involved joints are the hands, feet, and cervical spine, with corresponding joints on both sides of the body usually affected, though not always to the same extent. Rheumatoid arthritis is also associated with lung fibrosis, vasculitis (inflammation of blood vessels), coronary heart disease, and premature mortality. With no known cure, treatments are aimed at alleviating symptoms. Exercise, anti-inflammatory and pain medications, various specific disease-modifying anti-rheumatic drugs, or surgery are used to treat rheumatoid arthritis.

[caption id="" align="aligncenter" width="420"]The top panel in this figure shows a normal hip joint, and the bottom panel shows a hip joint with osteoarthritis. Figure 4. Osteoarthritis. Osteoarthritis of a synovial joint results from aging or prolonged joint wear and tear. These cause erosion and loss of the articular cartilage covering the surfaces of the bones, resulting in inflammation that causes joint stiffness and pain.[/caption]
[caption id="" align="alignleft" width="120"]QR Code representing a URL Visit this website to learn about a patient who arrives at the hospital with joint pain and weakness in his legs. What caused this patient’s weakness?[/caption]

]]> 2126 2108 5 0 9.5 Types of Body Movements https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2130 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2130

Learning Objectives

By the end of this section, you will be able to:
  • Define the different types of body movements
  • Identify the joints that allow for these motions
[caption id="" align="alignright" width="120"]QR Code representing a URL Watch this video to learn about anatomical motions. What motions involve increasing or decreasing the angle of the foot at the ankle?[/caption]

Synovial joints allow the body a tremendous range of movements. Each movement at a synovial joint results from the contraction or relaxation of the muscles that are attached to the bones on either side of the articulation. The type of movement that can be produced at a synovial joint is determined by its structural type. While the ball-and-socket joint gives the greatest range of movement at an individual joint, in other regions of the body, several joints may work together to produce a particular movement. Overall, each type of synovial joint is necessary to provide the body with its great flexibility and mobility. There are many types of movement that can occur at synovial joints (Table 1). Movement types are generally paired, with one being the opposite of the other. Body movements are always described in relation to the anatomical position of the body: upright stance, with upper limbs to the side of body and palms facing forward.

[caption id="attachment_1604" align="aligncenter" width="600"]This multi-part image shows different types of movements that are possible by different joints in the body. Figure 1. Movements of the Body, Part 1. Synovial joints give the body many ways in which to move. (a)–(b) Flexion and extension motions are in the sagittal (anterior–posterior) plane of motion. These movements take place at the shoulder, hip, elbow, knee, wrist, metacarpophalangeal, metatarsophalangeal, and interphalangeal joints. (c)–(d) Anterior bending of the head or vertebral column is flexion, while any posterior-going movement is extension. (e) Abduction and adduction are motions of the limbs, hand, fingers, or toes in the coronal (medial–lateral) plane of movement. Moving the limb or hand laterally away from the body, or spreading the fingers or toes, is abduction. Adduction brings the limb or hand toward or across the midline of the body, or brings the fingers or toes together. Circumduction is the movement of the limb, hand, or fingers in a circular pattern, using the sequential combination of flexion, adduction, extension, and abduction motions. Adduction/abduction and circumduction take place at the shoulder, hip, wrist, metacarpophalangeal, and metatarsophalangeal joints. (f) Turning of the head side to side or twisting of the body is rotation. Medial and lateral rotation of the upper limb at the shoulder or lower limb at the hip involves turning the anterior surface of the limb toward the midline of the body (medial or internal rotation) or away from the midline (lateral or external rotation).[/caption]
[caption id="attachment_1605" align="aligncenter" width="600"]This multi-part image shows different types of movements that are possible by different joints in the body. Figure 2. Movements of the Body, Part 2. (g) Supination of the forearm turns the hand to the palm forward position in which the radius and ulna are parallel, while forearm pronation turns the hand to the palm backward position in which the radius crosses over the ulna to form an "X." (h) Dorsiflexion of the foot at the ankle joint moves the top of the foot toward the leg, while plantar flexion lifts the heel and points the toes. (i) Eversion of the foot moves the bottom (sole) of the foot away from the midline of the body, while foot inversion faces the sole toward the midline. (j) Protraction of the mandible pushes the chin forward, and retraction pulls the chin back. (k) Depression of the mandible opens the mouth, while elevation closes it. (l) Opposition of the thumb brings the tip of the thumb into contact with the tip of the fingers of the same hand and reposition brings the thumb back next to the index finger.[/caption]

Flexion and Extension

Flexion and extension are movements that take place within the sagittal plane and involve anterior or posterior movements of the body or limbs. For the vertebral column, flexion (anterior flexion) is an anterior (forward) bending of the neck or body, while extension involves a posterior-directed motion, such as straightening from a flexed position or bending backward. Lateral flexion is the bending of the neck or body toward the right or left side. These movements of the vertebral column involve both the symphysis joint formed by each intervertebral disc, as well as the plane type of synovial joint formed between the inferior articular processes of one vertebra and the superior articular processes of the next lower vertebra. In the limbs, flexion decreases the angle between the bones (bending of the joint), while extension increases the angle and straightens the joint. For the upper limb, all anterior-going motions are flexion and all posterior-going motions are extension. These include anterior-posterior movements of the arm at the shoulder, the forearm at the elbow, the hand at the wrist, and the fingers at the metacarpophalangeal and interphalangeal joints. For the thumb, extension moves the thumb away from the palm of the hand, within the same plane as the palm, while flexion brings the thumb back against the index finger or into the palm. These motions take place at the first carpometacarpal joint. In the lower limb, bringing the thigh forward and upward is flexion at the hip joint, while any posterior-going motion of the thigh is extension. Note that extension of the thigh beyond the anatomical (standing) position is greatly limited by the ligaments that support the hip joint. Knee flexion is the bending of the knee to bring the foot toward the posterior thigh, and extension is the straightening of the knee. Flexion and extension movements are seen at the hinge, condyloid, saddle, and ball-and-socket joints of the limbs (see Figure 1a-d). Hyperextension is the abnormal or excessive extension of a joint beyond its normal range of motion, thus resulting in injury. Similarly, hyperflexion is excessive flexion at a joint. Hyperextension injuries are common at hinge joints such as the knee or elbow. In cases of “whiplash” in which the head is suddenly moved backward and then forward, a patient may experience both hyperextension and hyperflexion of the cervical region.

Abduction and Adduction

Abduction and adduction motions occur within the coronal plane and involve medial-lateral motions of the limbs, fingers, toes, or thumb. Abduction moves the limb laterally away from the midline of the body, while adduction is the opposing movement that brings the limb toward the body or across the midline. For example, abduction is raising the arm at the shoulder joint, moving it laterally away from the body, while adduction brings the arm down to the side of the body. Similarly, abduction and adduction at the wrist moves the hand away from or toward the midline of the body. Spreading the fingers or toes apart is also abduction, while bringing the fingers or toes together is adduction. For the thumb, abduction is the anterior movement that brings the thumb to a 90° perpendicular position, pointing straight out from the palm. Adduction moves the thumb back to the anatomical position, next to the index finger. Abduction and adduction movements are seen at condyloid, saddle, and ball-and-socket joints (see Figure 1e).

Circumduction

Circumduction is the movement of a body region in a circular manner, in which one end of the body region being moved stays relatively stationary while the other end describes a circle. It involves the sequential combination of flexion, adduction, extension, and abduction at a joint. This type of motion is found at biaxial condyloid and saddle joints, and at multiaxial ball-and-sockets joints (see Figure 1e).

Rotation

Rotation can occur within the vertebral column, at a pivot joint, or at a ball-and-socket joint. Rotation of the neck or body is the twisting movement produced by the summation of the small rotational movements available between adjacent vertebrae. At a pivot joint, one bone rotates in relation to another bone. This is a uniaxial joint, and thus rotation is the only motion allowed at a pivot joint. For example, at the atlantoaxial joint, the first cervical (C1) vertebra (atlas) rotates around the dens, the upward projection from the second cervical (C2) vertebra (axis). This allows the head to rotate from side to side as when shaking the head “no.” The proximal radioulnar joint is a pivot joint formed by the head of the radius and its articulation with the ulna. This joint allows for the radius to rotate along its length during pronation and supination movements of the forearm.

Rotation can also occur at the ball-and-socket joints of the shoulder and hip. Here, the humerus and femur rotate around their long axis, which moves the anterior surface of the arm or thigh either toward or away from the midline of the body. Movement that brings the anterior surface of the limb toward the midline of the body is called medial (internal) rotation. Conversely, rotation of the limb so that the anterior surface moves away from the midline is lateral (external) rotation (see Figure 1f). Be sure to distinguish medial and lateral rotation, which can only occur at the multiaxial shoulder and hip joints, from circumduction, which can occur at either biaxial or multiaxial joints.

Supination and Pronation

Supination and pronation are movements of the forearm. In the anatomical position, the upper limb is held next to the body with the palm facing forward. This is the supinated position of the forearm. In this position, the radius and ulna are parallel to each other. When the palm of the hand faces backward, the forearm is in the pronated position, and the radius and ulna form an X-shape.

Supination and pronation are the movements of the forearm that go between these two positions. Pronation is the motion that moves the forearm from the supinated (anatomical) position to the pronated (palm backward) position. This motion is produced by rotation of the radius at the proximal radioulnar joint, accompanied by movement of the radius at the distal radioulnar joint. The proximal radioulnar joint is a pivot joint that allows for rotation of the head of the radius. Because of the slight curvature of the shaft of the radius, this rotation causes the distal end of the radius to cross over the distal ulna at the distal radioulnar joint. This crossing over brings the radius and ulna into an X-shape position. Supination is the opposite motion, in which rotation of the radius returns the bones to their parallel positions and moves the palm to the anterior facing (supinated) position. It helps to remember that supination is the motion you use when scooping up soup with a spoon (see Figure 2g).

Dorsiflexion and Plantar Flexion

Dorsiflexion and plantar flexion are movements at the ankle joint, which is a hinge joint. Lifting the front of the foot, so that the top of the foot moves toward the anterior leg is dorsiflexion, while lifting the heel of the foot from the ground or pointing the toes downward is plantar flexion. These are the only movements available at the ankle joint (see Figure 2h).

Inversion and Eversion

Inversion and eversion are complex movements that involve the multiple plane joints among the tarsal bones of the posterior foot (intertarsal joints) and thus are not motions that take place at the ankle joint. Inversion is the turning of the foot to angle the bottom of the foot toward the midline, while eversion turns the bottom of the foot away from the midline. The foot has a greater range of inversion than eversion motion. These are important motions that help to stabilize the foot when walking or running on an uneven surface and aid in the quick side-to-side changes in direction used during active sports such as basketball, racquetball, or soccer (see Figure 2i).

Protraction and Retraction

Protraction and retraction are anterior-posterior movements of the scapula or mandible. Protraction of the scapula occurs when the shoulder is moved forward, as when pushing against something or throwing a ball. Retraction is the opposite motion, with the scapula being pulled posteriorly and medially, toward the vertebral column. For the mandible, protraction occurs when the lower jaw is pushed forward, to stick out the chin, while retraction pulls the lower jaw backward. (See Figure 2j.)

Depression and Elevation

Depression and elevation are downward and upward movements of the scapula or mandible. The upward movement of the scapula and shoulder is elevation, while a downward movement is depression. These movements are used to shrug your shoulders. Similarly, elevation of the mandible is the upward movement of the lower jaw used to close the mouth or bite on something, and depression is the downward movement that produces opening of the mouth (see Figure 2k).

Excursion

Excursion is the side to side movement of the mandible. Lateral excursion moves the mandible away from the midline, toward either the right or left side. Medial excursion returns the mandible to its resting position at the midline.

Superior Rotation and Inferior Rotation

Superior and inferior rotation are movements of the scapula and are defined by the direction of movement of the glenoid cavity. These motions involve rotation of the scapula around a point inferior to the scapular spine and are produced by combinations of muscles acting on the scapula. During superior rotation, the glenoid cavity moves upward as the medial end of the scapular spine moves downward. This is a very important motion that contributes to upper limb abduction. Without superior rotation of the scapula, the greater tubercle of the humerus would hit the acromion of the scapula, thus preventing any abduction of the arm above shoulder height. Superior rotation of the scapula is thus required for full abduction of the upper limb. Superior rotation is also used without arm abduction when carrying a heavy load with your hand or on your shoulder. You can feel this rotation when you pick up a load, such as a heavy book bag and carry it on only one shoulder. To increase its weight-bearing support for the bag, the shoulder lifts as the scapula superiorly rotates. Inferior rotation occurs during limb adduction and involves the downward motion of the glenoid cavity with upward movement of the medial end of the scapular spine.

Opposition and Reposition

Opposition is the thumb movement that brings the tip of the thumb in contact with the tip of a finger. This movement is produced at the first carpometacarpal joint, which is a saddle joint formed between the trapezium carpal bone and the first metacarpal bone. Thumb opposition is produced by a combination of flexion and abduction of the thumb at this joint. Returning the thumb to its anatomical position next to the index finger is called reposition (see Figure 2l).

Movements of the Joints (Table 1)
Type of Joint Movement Example
Pivot Uniaxial joint; allows rotational movement Atlantoaxial joint (C1–C2 vertebrae articulation); proximal radioulnar joint
Hinge Uniaxial joint; allows flexion/extension movements Knee; elbow; ankle; interphalangeal joints of fingers and toes
Condyloid Biaxial joint; allows flexion/extension, abduction/adduction, and circumduction movements Metacarpophalangeal (knuckle) joints of fingers; radiocarpal joint of wrist; metatarsophalangeal joints for toes
Saddle Biaxial joint; allows flexion/extension, abduction/adduction, and circumduction movements First carpometacarpal joint of the thumb; sternoclavicular joint
Plane Multiaxial joint; allows inversion and eversion of foot, or flexion, extension, and lateral flexion of the vertebral column Intertarsal joints of foot; superior-inferior articular process articulations between vertebrae
Ball-and-socket Multiaxial joint; allows flexion/extension, abduction/adduction, circumduction, and medial/lateral rotation movements Shoulder and hip joints

]]> 2130 2108 6 0 9.6 Anatomy of Selected Synovial Joints https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2134 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2134

Learning Objectives

By the end of this section, you will be able to
  • Describe the bones that articulate together to form selected synovial joints
  • Discuss the movements available at each joint
  • Describe the structures that support and prevent excess movements at each joint

Each synovial joint of the body is specialized to perform certain movements. The movements that are allowed are determined by the structural classification for each joint. For example, a multiaxial ball-and-socket joint has much more mobility than a uniaxial hinge joint. However, the ligaments and muscles that support a joint may place restrictions on the total range of motion available. Thus, the ball-and-socket joint of the shoulder has little in the way of ligament support, which gives the shoulder a very large range of motion. In contrast, movements at the hip joint are restricted by strong ligaments, which reduce its range of motion but confer stability during standing and weight bearing.

This section will examine the anatomy of selected synovial joints of the body. Anatomical names for most joints are derived from the names of the bones that articulate at that joint, although some joints, such as the elbow, hip, and knee joints are exceptions to this general naming scheme.

Knee Joint

The knee joint is the largest joint of the body (Figure 6). It actually consists of three articulations. The femoropatellar joint is found between the patella and the distal femur. The medial tibiofemoral joint and lateral tibiofemoral joint are located between the medial and lateral condyles of the femur and the medial and lateral condyles of the tibia. All of these articulations are enclosed within a single articular capsule. The knee functions as a hinge joint, allowing flexion and extension of the leg. This action is generated by both rolling and gliding motions of the femur on the tibia. In addition, some rotation of the leg is available when the knee is flexed, but not when extended. The knee is well constructed for weight bearing in its extended position, but is vulnerable to injuries associated with hyperextension, twisting, or blows to the medial or lateral side of the joint, particularly while weight bearing.

At the femoropatellar joint, the patella slides vertically within a groove on the distal femur. The patella is a sesamoid bone incorporated into the tendon of the quadriceps femoris muscle, the large muscle of the anterior thigh. The patella serves to protect the quadriceps tendon from friction against the distal femur. Continuing from the patella to the anterior tibia just below the knee is the patellar ligament. Acting via the patella and patellar ligament, the quadriceps femoris is a powerful muscle that acts to extend the leg at the knee. It also serves as a “dynamic ligament” to provide very important support and stabilization for the knee joint.

The medial and lateral tibiofemoral joints are the articulations between the rounded condyles of the femur and the relatively flat condyles of the tibia. During flexion and extension motions, the condyles of the femur both roll and glide over the surfaces of the tibia. The rolling action produces flexion or extension, while the gliding action serves to maintain the femoral condyles centered over the tibial condyles, thus ensuring maximal bony, weight-bearing support for the femur in all knee positions. As the knee comes into full extension, the femur undergoes a slight medial rotation in relation to tibia. The rotation results because the lateral condyle of the femur is slightly smaller than the medial condyle. Thus, the lateral condyle finishes its rolling motion first, followed by the medial condyle. The resulting small medial rotation of the femur serves to “lock” the knee into its fully extended and most stable position. Flexion of the knee is initiated by a slight lateral rotation of the femur on the tibia, which “unlocks” the knee. This lateral rotation motion is produced by the popliteus muscle of the posterior leg.

Located between the articulating surfaces of the femur and tibia are two articular discs, the medial meniscus and lateral meniscus (see Figure 6b). Each is a C-shaped fibrocartilage structure that is thin along its inside margin and thick along the outer margin. They are attached to their tibial condyles, but do not attach to the femur. While both menisci are free to move during knee motions, the medial meniscus shows less movement because it is anchored at its outer margin to the articular capsule and tibial collateral ligament. The menisci provide padding between the bones and help to fill the gap between the round femoral condyles and flattened tibial condyles. Some areas of each meniscus lack an arterial blood supply and thus these areas heal poorly if damaged.

The knee joint has multiple ligaments that provide support, particularly in the extended position (see Figure 6c). Outside of the articular capsule, located at the sides of the knee, are two extrinsic ligaments. The fibular collateral ligament (lateral collateral ligament) is on the lateral side and spans from the lateral epicondyle of the femur to the head of the fibula. The tibial collateral ligament (medial collateral ligament) of the medial knee runs from the medial epicondyle of the femur to the medial tibia. As it crosses the knee, the tibial collateral ligament is firmly attached on its deep side to the articular capsule and to the medial meniscus, an important factor when considering knee injuries. In the fully extended knee position, both collateral ligaments are taut (tight), thus serving to stabilize and support the extended knee and preventing side-to-side or rotational motions between the femur and tibia.

The articular capsule of the posterior knee is thickened by intrinsic ligaments that help to resist knee hyperextension. Inside the knee are two intracapsular ligaments, the anterior cruciate ligament and posterior cruciate ligament. These ligaments are anchored inferiorly to the tibia at the intercondylar eminence, the roughened area between the tibial condyles. The cruciate ligaments are named for whether they are attached anteriorly or posteriorly to this tibial region. Each ligament runs diagonally upward to attach to the inner aspect of a femoral condyle. The cruciate ligaments are named for the X-shape formed as they pass each other (cruciate means “cross”). The posterior cruciate ligament is the stronger ligament. It serves to support the knee when it is flexed and weight bearing, as when walking downhill. In this position, the posterior cruciate ligament prevents the femur from sliding anteriorly off the top of the tibia. The anterior cruciate ligament becomes tight when the knee is extended, and thus resists hyperextension.

[caption id="" align="aligncenter" width="620"]This image shows the different views of the knee joint. The top, left panel shows the sagittal view of the right knee joint. The top, left panel shows the superior view of the right tibia, identifying the ligaments. The bottom, right panel shows the anterior view of the right knee. Figure 6. Knee Joint. (a) The knee joint is the largest joint of the body. (b)–(c) It is supported by the tibial and fibular collateral ligaments located on the sides of the knee outside of the articular capsule, and the anterior and posterior cruciate ligaments found inside the capsule. The medial and lateral menisci provide padding and support between the femoral condyles and tibial condyles.[/caption]
[caption id="" align="alignleft" width="120"]QR Code representing a URL Watch this video to learn more about the flexion and extension of the knee, as the femur both rolls and glides on the tibia to maintain stable contact between the bones in all knee positions.[/caption]

[caption id="" align="alignright" width="120"]QR Code representing a URL Watch this video to learn more about the anatomy of the knee joint, including bones, joints, muscles, nerves, and blood vessels.[/caption]
]]> 2134 2108 7 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2137 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2137 [caption id="" align="aligncenter" width="600"]This photograph shows a man playing tennis. Figure 1. Tennis Player. Athletes rely on toned skeletal muscles to supply the force required for movement. (credit: Emmanuel Huybrechts/flickr)[/caption]

Learning Objectives

After studying this chapter, you will be able to:
  • Explain the organization of muscle tissue
  • Describe the function and structure of skeletal, cardiac muscle, and smooth muscle
  • Explain how muscles work with tendons to move the body
  • Describe how muscles contract and relax
  • Define the process of muscle metabolism
  • Explain how the nervous system controls muscle tension
  • Relate the connections between exercise and muscle performance
  • Explain the development and regeneration of muscle tissue
When most people think of muscles, they think of the muscles that are visible just under the skin, particularly of the limbs. These are skeletal muscles, so-named because most of them move the skeleton. But there are two other types of muscle in the body, with distinctly different jobs. Cardiac muscle, found in the heart, is concerned with pumping blood through the circulatory system. Smooth muscle is concerned with various involuntary movements, such as having one’s hair stand on end when cold or frightened, or moving food through the digestive system. This chapter will examine the structure and function of these three types of muscles.]]> 2137 2135 1 0 10.1 Overview of Muscle Tissues https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2139 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2139

Learning Objectives

By the end of this section, you will be able to:
  • Describe the different types of muscle
  • Explain contractibility and extensibility

Muscle is one of the four primary tissue types of the body, and the body contains three types of muscle tissue: skeletal muscle, cardiac muscle, and smooth muscle (Figure 1). All three muscle tissues have some properties in common; they all exhibit a quality called excitability as their plasma membranes can change their electrical states (from polarized to depolarized) and send an electrical wave called an action potential along the entire length of the membrane. While the nervous system can influence the excitability of cardiac and smooth muscle to some degree, skeletal muscle completely depends on signaling from the nervous system to work properly. On the other hand, both cardiac muscle and smooth muscle can respond to other stimuli, such as hormones and local stimuli.

[caption id="" align="aligncenter" width="320"]This figure show the micrographs of skeletal muscle, smooth muscle, and cardiac muscle cells. Figure 1. The Three Types of Muscle Tissue. The body contains three types of muscle tissue: (a) skeletal muscle, (b) smooth muscle, and (c) cardiac muscle. From top, LM × 1600, LM × 1600, LM × 1600. (Micrographs provided by the Regents of University of Michigan Medical School © 2012)[/caption]

The muscles all begin the actual process of contracting (shortening) when a protein called actin is pulled by a protein called myosin. This occurs in striated muscle (skeletal and cardiac) after specific binding sites on the actin have been exposed in response to the interaction between calcium ions (Ca++) and proteins (troponin and tropomyosin) that “shield” the actin-binding sites. Ca++ also is required for the contraction of smooth muscle, although its role is different: here Ca++ activates enzymes, which in turn activate myosin heads. All muscles require adenosine triphosphate (ATP) to continue the process of contracting, and they all relax when the Ca++ is removed and the actin-binding sites are re-shielded.

A muscle can return to its original length when relaxed due to a quality of muscle tissue called elasticity. It can recoil back to its original length due to elastic fibers. Muscle tissue also has the quality of extensibility; it can stretch or extend. Contractility allows muscle tissue to pull on its attachment points and shorten with force.

Differences among the three muscle types include the microscopic organization of their contractile proteins—actin and myosin. The actin and myosin proteins are arranged very regularly in the cytoplasm of individual muscle cells (referred to as fibers) in both skeletal muscle and cardiac muscle, which creates a pattern, or stripes, called striations. The striations are visible with a light microscope under high magnification (see Figure 1). Skeletal muscle fibers are multinucleated structures that compose the skeletal muscle. Cardiac muscle fibers each have one to two nuclei and are physically and electrically connected to each other so that the entire heart contracts as one unit (called a syncytium).

Because the actin and myosin are not arranged in such regular fashion in smooth muscle, the cytoplasm of a smooth muscle fiber (which has only a single nucleus) has a uniform, nonstriated appearance (resulting in the name smooth muscle). However, the less organized appearance of smooth muscle should not be interpreted as less efficient. Smooth muscle in the walls of arteries is a critical component that regulates blood pressure necessary to push blood through the circulatory system; and smooth muscle in the skin, visceral organs, and internal passageways is essential for moving all materials through the body.

]]> 2139 2135 2 0 10.3 Muscle Fiber Contraction and Relaxation https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2154 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2154

Learning Objectives

By the end of this section, you will be able to:
  • Describe the components involved in a muscle contraction
  • Explain how muscles contract and relax
  • Describe the sliding filament model of muscle contraction

The sequence of events that result in the contraction of an individual muscle fiber begins with a signal—the neurotransmitter, ACh—from the motor neuron innervating that fiber. The local membrane of the fiber will depolarize as positively charged sodium ions (Na+) enter, triggering an action potential that spreads to the rest of the membrane will depolarize, including the T-tubules. This triggers the release of calcium ions (Ca++) from storage in the sarcoplasmic reticulum (SR). The Ca++ then initiates contraction, which is sustained by ATP (Figure 1). As long as Ca++ ions remain in the sarcoplasm to bind to troponin, which keeps the actin-binding sites “unshielded,” and as long as ATP is available to drive the cross-bridge cycling and the pulling of actin strands by myosin, the muscle fiber will continue to shorten to an anatomical limit.

[caption id="" align="aligncenter" width="400"]The top panel in this figure shows the interaction of a motor neuron with a muscle fiber and how the release of acetylcholine into the muscle cells leads to the release of calcium. The middle panel shows how calcium release activates troponin and leads to muscle contraction. The bottom panel shows an image of a muscle fiber being shortened and producing tension. Figure 1. Contraction of a Muscle Fiber. A cross-bridge forms between actin and the myosin heads triggering contraction. As long as Ca++ ions remain in the sarcoplasm to bind to troponin, and as long as ATP is available, the muscle fiber will continue to shorten.[/caption]

Muscle contraction usually stops when signaling from the motor neuron ends, which repolarizes the sarcolemma and T-tubules, and closes the voltage-gated calcium channels in the SR. Ca++ ions are then pumped back into the SR, which causes the tropomyosin to reshield (or re-cover) the binding sites on the actin strands. A muscle also can stop contracting when it runs out of ATP and becomes fatigued (Figure 2).

[caption id="" align="aligncenter" width="380"]The top panel in this figure shows the interaction of a motor neuron with a muscle fiber and how calcium is being absorbed into the muscle fiber. This results in the relaxation of the thin and thick filaments as shown in the bottom panel. Figure 2. Relaxation of a Muscle Fiber. Ca++ ions are pumped back into the SR, which causes the tropomyosin to reshield the binding sites on the actin strands. A muscle may also stop contracting when it runs out of ATP and becomes fatigued.[/caption]
[caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this video to learn more about the role of calcium.[/caption]
The molecular events of muscle fiber shortening occur within the fiber’s sarcomeres (see Figure 3). The contraction of a striated muscle fiber occurs as the sarcomeres, linearly arranged within myofibrils, shorten as myosin heads pull on the actin filaments.

The region where thick and thin filaments overlap has a dense appearance, as there is little space between the filaments. This zone where thin and thick filaments overlap is very important to muscle contraction, as it is the site where filament movement starts. Thin filaments, anchored at their ends by the Z-discs, do not extend completely into the central region that only contains thick filaments, anchored at their bases at a spot called the M-line. A myofibril is composed of many sarcomeres running along its length; thus, myofibrils and muscle cells contract as the sarcomeres contract.

The Sliding Filament Model of Contraction

When signaled by a motor neuron, a skeletal muscle fiber contracts as the thin filaments are pulled and then slide past the thick filaments within the fiber’s sarcomeres. This process is known as the sliding filament model of muscle contraction (Figure 3). The sliding can only occur when myosin-binding sites on the actin filaments are exposed by a series of steps that begins with Ca++ entry into the sarcoplasm.

[caption id="" align="aligncenter" width="450"]This diagram shows how muscle contracts. The top panel shows the stretched filaments and the bottom panel shows the compressed filaments. Figure 3. The Sliding Filament Model of Muscle Contraction. When a sarcomere contracts, the Z lines move closer together, and the I band becomes smaller. The A band stays the same width. At full contraction, the thin and thick filaments overlap.[/caption]
Tropomyosin is a protein that winds around the chains of the actin filament and covers the myosin-binding sites to prevent actin from binding to myosin. Tropomyosin binds to troponin to form a troponin-tropomyosin complex. The troponin-tropomyosin complex prevents the myosin “heads” from binding to the active sites on the actin microfilaments. Troponin also has a binding site for Ca++ ions.

To initiate muscle contraction, tropomyosin has to expose the myosin-binding site on an actin filament to allow cross-bridge formation between the actin and myosin microfilaments. The first step in the process of contraction is for Ca++ to bind to troponin so that tropomyosin can slide away from the binding sites on the actin strands. This allows the myosin heads to bind to these exposed binding sites and form cross-bridges. The thin filaments are then pulled by the myosin heads to slide past the thick filaments toward the center of the sarcomere. But each head can only pull a very short distance before it has reached its limit and must be “re-cocked” before it can pull again, a step that requires ATP.

ATP and Muscle Contraction

For thin filaments to continue to slide past thick filaments during muscle contraction, myosin heads must pull the actin at the binding sites, detach, re-cock, attach to more binding sites, pull, detach, re-cock, etc. This repeated movement is known as the cross-bridge cycle. This motion of the myosin heads is similar to the oars when an individual rows a boat: The paddle of the oars (the myosin heads) pull, are lifted from the water (detach), repositioned (re-cocked) and then immersed again to pull (Figure 4). Each cycle requires energy, and the action of the myosin heads in the sarcomeres repetitively pulling on the thin filaments also requires energy, which is provided by ATP.

[caption id="" align="aligncenter" width="450"]This multipart figure shows the mechanism of skeletal muscle contraction. In the top panel, the ADP and inorganic phosphate molecules are bound to the myosin motor head. In the middle panel, the ADP and phosphate come off the myosin motor and the direction of the power stroke is shown. In the bottom panel, a molecule of ATP is shown to bind the myosin motor head and the motor is reset. Figure 4. Skeletal Muscle Contraction. (a) The active site on actin is exposed as calcium binds to troponin. (b) The myosin head is attracted to actin, and myosin binds actin at its actin-binding site, forming the cross-bridge. (c) During the power stroke, the phosphate generated in the previous contraction cycle is released. This results in the myosin head pivoting toward the center of the sarcomere, after which the attached ADP and phosphate group are released. (d) A new molecule of ATP attaches to the myosin head, causing the cross-bridge to detach. (e) The myosin head hydrolyzes ATP to ADP and phosphate, which returns the myosin to the cocked position.[/caption]

Cross-bridge formation occurs when the myosin head attaches to the actin while adenosine diphosphate (ADP) and inorganic phosphate (Pi) are still bound to myosin (Figure 4a,b). Pi is then released, causing myosin to form a stronger attachment to the actin, after which the myosin head moves toward the M-line, pulling the actin along with it. As actin is pulled, the filaments move approximately 10 nm toward the M-line. This movement is called the power stroke, as movement of the thin filament occurs at this step (Figure 4c). In the absence of ATP, the myosin head will not detach from actin.

One part of the myosin head attaches to the binding site on the actin, but the head has another binding site for ATP. ATP binding causes the myosin head to detach from the actin (Figure 4d). After this occurs, ATP is converted to ADP and Pi by the intrinsic ATPase activity of myosin. The energy released during ATP hydrolysis changes the angle of the myosin head into a cocked position (Figure 4e). The myosin head is now in position for further movement.

When the myosin head is cocked, myosin is in a high-energy configuration. This energy is expended as the myosin head moves through the power stroke, and at the end of the power stroke, the myosin head is in a low-energy position. After the power stroke, ADP is released; however, the formed cross-bridge is still in place, and actin and myosin are bound together. As long as ATP is available, it readily attaches to myosin, the cross-bridge cycle can recur, and muscle contraction can continue.

Note that each thick filament of roughly 300 myosin molecules has multiple myosin heads, and many cross-bridges form and break continuously during muscle contraction. Multiply this by all of the sarcomeres in one myofibril, all the myofibrils in one muscle fiber, and all of the muscle fibers in one skeletal muscle, and you can understand why so much energy (ATP) is needed to keep skeletal muscles working. In fact, it is the loss of ATP that results in the rigor mortis observed soon after someone dies. With no further ATP production possible, there is no ATP available for myosin heads to detach from the actin-binding sites, so the cross-bridges stay in place, causing the rigidity in the skeletal muscles.

Sources of ATP

ATP supplies the energy for muscle contraction to take place. In addition to its direct role in the cross-bridge cycle, ATP also provides the energy for the active-transport Ca++ pumps in the SR. Muscle contraction does not occur without sufficient amounts of ATP. The amount of ATP stored in muscle is very low, only sufficient to power a few seconds worth of contractions. As it is broken down, ATP must therefore be regenerated and replaced quickly to allow for sustained contraction. There are three mechanisms by which ATP can be regenerated: creatine phosphate metabolism, anaerobic glycolysis, fermentation and aerobic respiration.

Creatine phosphate is a molecule that can store energy in its phosphate bonds. In a resting muscle, excess ATP transfers its energy to creatine, producing ADP and creatine phosphate. This acts as an energy reserve that can be used to quickly create more ATP. When the muscle starts to contract and needs energy, creatine phosphate transfers its phosphate back to ADP to form ATP and creatine. This reaction is catalyzed by the enzyme creatine kinase and occurs very quickly; thus, creatine phosphate-derived ATP powers the first few seconds of muscle contraction. However, creatine phosphate can only provide approximately 15 seconds worth of energy, at which point another energy source has to be used (Figure 5).

[caption id="" align="aligncenter" width="430"]This figure shows the metabolic processes in muscle. The top panel shows the reactions in resting muscle. The middle panel shows glycolysis and aerobic respiration and the bottom panel shows cellular respiration in mitochondria. Figure 5. Muscle Metabolism. (a) Some ATP is stored in a resting muscle. As contraction starts, it is used up in seconds. More ATP is generated from creatine phosphate for about 15 seconds. (b) Each glucose molecule produces two ATP and two molecules of pyruvic acid, which can be used in aerobic respiration or converted to lactic acid. If oxygen is not available, pyruvic acid is converted to lactic acid, which may contribute to muscle fatigue. This occurs during strenuous exercise when high amounts of energy are needed but oxygen cannot be sufficiently delivered to muscle. (c) Aerobic respiration is the breakdown of glucose in the presence of oxygen (O2) to produce carbon dioxide, water, and ATP. Approximately 95 percent of the ATP required for resting or moderately active muscles is provided by aerobic respiration, which takes place in mitochondria.[/caption]

As the ATP produced by creatine phosphate is depleted, muscles turn to glycolysis as an ATP source. Glycolysis is an anaerobic (non-oxygen-dependent) process that breaks down glucose (sugar) to produce ATP; however, glycolysis cannot generate ATP as quickly as creatine phosphate. Thus, the switch to glycolysis results in a slower rate of ATP availability to the muscle. The sugar used in glycolysis can be provided by blood glucose or by metabolizing glycogen that is stored in the muscle. The breakdown of one glucose molecule produces two ATP and two molecules of pyruvic acid, which can be used in aerobic respiration or when oxygen levels are low, converted to lactic acid (Figure 5b).

If oxygen is available, pyruvic acid is used in aerobic respiration. However, if oxygen is not available, pyruvic acid is converted to lactic acid, which may contribute to muscle fatigue. This conversion allows the recycling of the enzyme NAD+ from NADH, which is needed for glycolysis to continue. This occurs during strenuous exercise when high amounts of energy are needed but oxygen cannot be sufficiently delivered to muscle. Glycolysis itself cannot be sustained for very long (approximately 1 minute of muscle activity), but it is useful in facilitating short bursts of high-intensity output. This is because glycolysis does not utilize glucose very efficiently, producing a net gain of two ATPs per molecule of glucose, and the end product of lactic acid, which may contribute to muscle fatigue as it accumulates.

Aerobic respiration is the breakdown of glucose or other nutrients in the presence of oxygen (O2) to produce carbon dioxide, water, and ATP. Approximately 95 percent of the ATP required for resting or moderately active muscles is provided by aerobic respiration, which takes place in mitochondria. The inputs for aerobic respiration include glucose circulating in the bloodstream, pyruvic acid, and fatty acids. Aerobic respiration is much more efficient than anaerobic glycolysis, producing approximately 36 ATPs per molecule of glucose versus four from glycolysis. However, aerobic respiration cannot be sustained without a steady supply of O2 to the skeletal muscle and is much slower (Figure 5c). To compensate, muscles store small amount of excess oxygen in proteins call myoglobin, allowing for more efficient muscle contractions and less fatigue. Aerobic training also increases the efficiency of the circulatory system so that O2 can be supplied to the muscles for longer periods of time.

Muscle fatigue occurs when a muscle can no longer contract in response to signals from the nervous system. The exact causes of muscle fatigue are not fully known, although certain factors have been correlated with the decreased muscle contraction that occurs during fatigue. ATP is needed for normal muscle contraction, and as ATP reserves are reduced, muscle function may decline. This may be more of a factor in brief, intense muscle output rather than sustained, lower intensity efforts. Lactic acid buildup may lower intracellular pH, affecting enzyme and protein activity. Imbalances in Na+ and K+ levels as a result of membrane depolarization may disrupt Ca++ flow out of the SR. Long periods of sustained exercise may damage the SR and the sarcolemma, resulting in impaired Ca++ regulation.

Intense muscle activity results in an oxygen debt, which is the amount of oxygen needed to compensate for ATP produced without oxygen during muscle contraction. Oxygen is required to restore ATP and creatine phosphate levels, convert lactic acid to pyruvic acid, and, in the liver, to convert lactic acid into glucose or glycogen. Other systems used during exercise also require oxygen, and all of these combined processes result in the increased breathing rate that occurs after exercise. Until the oxygen debt has been met, oxygen intake is elevated, even after exercise has stopped.

Relaxation of a Skeletal Muscle

Relaxing skeletal muscle fibers, and ultimately, the skeletal muscle, begins with the motor neuron, which stops releasing its chemical signal, ACh, into the synapse at the NMJ. The muscle fiber will repolarize, which closes the gates in the SR where Ca++ was being released. ATP-driven pumps will move Ca++ out of the sarcoplasm back into the SR. This results in the “reshielding” of the actin-binding sites on the thin filaments. Without the ability to form cross-bridges between the thin and thick filaments, the muscle fiber loses its tension and relaxes.

Muscle Strength

The number of skeletal muscle fibers in a given muscle is genetically determined and does not change. Muscle strength is directly related to the amount of myofibrils and sarcomeres within each fiber. Factors, such as hormones and stress (and artificial anabolic steroids), acting on the muscle can increase the production of sarcomeres and myofibrils within the muscle fibers, a change called hypertrophy, which results in the increased mass and bulk in a skeletal muscle. Likewise, decreased use of a skeletal muscle results in atrophy, where the number of sarcomeres and myofibrils disappear (but not the number of muscle fibers). It is common for a limb in a cast to show atrophied muscles when the cast is removed, and certain diseases, such as polio, show atrophied muscles.

Disorders of the Muscular System

Duchenne muscular dystrophy (DMD) is a progressive weakening of the skeletal muscles. It is one of several diseases collectively referred to as “muscular dystrophy.” DMD is caused by a lack of the protein dystrophin, which helps the thin filaments of myofibrils bind to the sarcolemma. Without sufficient dystrophin, muscle contractions cause the sarcolemma to tear, causing an influx of Ca++, leading to cellular damage and muscle fiber degradation. Over time, as muscle damage accumulates, muscle mass is lost, and greater functional impairments develop.

DMD is an inherited disorder caused by an abnormal X chromosome. It primarily affects males, and it is usually diagnosed in early childhood. DMD usually first appears as difficulty with balance and motion, and then progresses to an inability to walk. It continues progressing upward in the body from the lower extremities to the upper body, where it affects the muscles responsible for breathing and circulation. It ultimately causes death due to respiratory failure, and those afflicted do not usually live past their 20s.

Because DMD is caused by a mutation in the gene that codes for dystrophin, it was thought that introducing healthy myoblasts into patients might be an effective treatment. Myoblasts are the embryonic cells responsible for muscle development, and ideally, they would carry healthy genes that could produce the dystrophin needed for normal muscle contraction. This approach has been largely unsuccessful in humans. A recent approach has involved attempting to boost the muscle’s production of utrophin, a protein similar to dystrophin that may be able to assume the role of dystrophin and prevent cellular damage from occurring.

]]> 2154 2135 4 0 10.4 Nervous System Control of Muscle Tension https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2157 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2157

Learning Objectives

By the end of this section, you will be able to:
  • Explain concentric, isotonic, and eccentric contractions
  • Describe the length-tension relationship
  • Describe the three phases of a muscle twitch
  • Define wave summation, tetanus, and treppe

To move an object, referred to as load, the sarcomeres in the muscle fibers of the skeletal muscle must shorten. The force generated by the contraction of the muscle (or shortening of the sarcomeres) is called muscle tension. However, muscle tension also is generated when the muscle is contracting against a load that does not move, resulting in two main types of skeletal muscle contractions: isotonic contractions and isometric contractions.

In isotonic contractions, where the tension in the muscle stays constant, a load is moved as the length of the muscle changes (shortens). There are two types of isotonic contractions: concentric and eccentric. A concentric contraction involves the muscle shortening to move a load. An example of this is the biceps brachii muscle contracting when a hand weight is brought upward with increasing muscle tension. As the biceps brachii contract, the angle of the elbow joint decreases as the forearm is brought toward the body. Here, the biceps brachii contracts as sarcomeres in its muscle fibers are shortening and cross-bridges form; the myosin heads pull the actin. An eccentric contraction occurs as the muscle tension diminishes and the muscle lengthens. In this case, the hand weight is lowered in a slow and controlled manner as the amount of cross-bridges being activated by nervous system stimulation decreases. In this case, as tension is released from the biceps brachii, the angle of the elbow joint increases. Eccentric contractions are also used for movement and balance of the body.

An isometric contraction occurs as the muscle produces tension without changing the angle of a skeletal joint. Isometric contractions involve sarcomere shortening and increasing muscle tension, but do not move a load, as the force produced cannot overcome the resistance provided by the load. For example, if one attempts to lift a hand weight that is too heavy, there will be sarcomere activation and shortening to a point, and ever-increasing muscle tension, but no change in the angle of the elbow joint. In everyday living, isometric contractions are active in maintaining posture and maintaining bone and joint stability. However, holding your head in an upright position occurs not because the muscles cannot move the head, but because the goal is to remain stationary and not produce movement. Most actions of the body are the result of a combination of isotonic and isometric contractions working together to produce a wide range of outcomes (Figure 1).

[caption id="" align="aligncenter" width="390"]This figure shows the different types of muscle contraction and the associated body movements. The top panel shows concentric contraction, the middle panel shows eccentric contraction, and the bottom panel shows isometric contraction. Figure 1. Types of Muscle Contractions. During isotonic contractions, muscle length changes to move a load. During isometric contractions, muscle length does not change because the load exceeds the tension the muscle can generate.[/caption]

All of these muscle activities are under the exquisite control of the nervous system. Neural control regulates concentric, eccentric and isometric contractions, muscle fiber recruitment, and muscle tone. A crucial aspect of nervous system control of skeletal muscles is the role of motor units.

Motor Units

As you have learned, every skeletal muscle fiber must be innervated by the axon terminal of a motor neuron in order to contract. Each muscle fiber is innervated by only one motor neuron. The actual group of muscle fibers in a muscle innervated by a single motor neuron is called a motor unit. The size of a motor unit is variable depending on the nature of the muscle.

A small motor unit is an arrangement where a single motor neuron supplies a small number of muscle fibers in a muscle. Small motor units permit very fine motor control of the muscle. The best example in humans is the small motor units of the extraocular eye muscles that move the eyeballs. There are thousands of muscle fibers in each muscle, but every six or so fibers are supplied by a single motor neuron, as the axons branch to form synaptic connections at their individual NMJs. This allows for exquisite control of eye movements so that both eyes can quickly focus on the same object. Small motor units are also involved in the many fine movements of the fingers and thumb of the hand for grasping, texting, etc.

A large motor unit is an arrangement where a single motor neuron supplies a large number of muscle fibers in a muscle. Large motor units are concerned with simple, or “gross,” movements, such as powerfully extending the knee joint. The best example is the large motor units of the thigh muscles or back muscles, where a single motor neuron will supply thousands of muscle fibers in a muscle, as its axon splits into thousands of branches.

There is a wide range of motor units within many skeletal muscles, which gives the nervous system a wide range of control over the muscle. The small motor units in the muscle will have smaller, lower-threshold motor neurons that are more excitable, firing first to their skeletal muscle fibers, which also tend to be the smallest. Activation of these smaller motor units, results in a relatively small degree of contractile strength (tension) generated in the muscle. As more strength is needed, larger motor units, with bigger, higher-threshold motor neurons are enlisted to activate larger muscle fibers. This increasing activation of motor units produces an increase in muscle contraction known as recruitment. As more motor units are recruited, the muscle contraction grows progressively stronger. In some muscles, the largest motor units may generate a contractile force of 50 times more than the smallest motor units in the muscle. This allows a feather to be picked up using the biceps brachii arm muscle with minimal force, and a heavy weight to be lifted by the same muscle by recruiting the largest motor units.

When necessary, the maximal number of motor units in a muscle can be recruited simultaneously, producing the maximum force of contraction for that muscle, but this cannot last for very long because of the energy requirements to sustain the contraction. To prevent complete muscle fatigue, motor units are generally not all simultaneously active, but instead some motor units rest while others are active, which allows for longer muscle contractions. The nervous system uses recruitment as a mechanism to efficiently utilize a skeletal muscle.

The Length-Tension Range of a Sarcomere

When a skeletal muscle fiber contracts, myosin heads attach to actin to form cross-bridges followed by the thin filaments sliding over the thick filaments as the heads pull the actin, and this results in sarcomere shortening, creating the tension of the muscle contraction. The cross-bridges can only form where thin and thick filaments already overlap, so that the length of the sarcomere has a direct influence on the force generated when the sarcomere shortens. This is called the length-tension relationship.

The ideal length of a sarcomere to produce maximal tension occurs at 80 percent to 120 percent of its resting length, with 100 percent being the state where the medial edges of the thin filaments are just at the most-medial myosin heads of the thick filaments (Figure 2). This length maximizes the overlap of actin-binding sites and myosin heads. If a sarcomere is stretched past this ideal length (beyond 120 percent), thick and thin filaments do not overlap sufficiently, which results in less tension produced. If a sarcomere is shortened beyond 80 percent, the zone of overlap is reduced with the thin filaments jutting beyond the last of the myosin heads and shrinks the H zone, which is normally composed of myosin tails. Eventually, there is nowhere else for the thin filaments to go and the amount of tension is diminished. If the muscle is stretched to the point where thick and thin filaments do not overlap at all, no cross-bridges can be formed, and no tension is produced in that sarcomere. This amount of stretching does not usually occur, as accessory proteins and connective tissue oppose extreme stretching.

[caption id="" align="aligncenter" width="380"]A graph shows the percent sarcomere length on the x-axis and tension on the y-axis. As the length of the sarcomere increases, the tension first increases, and then decreases. Figure 2. The Ideal Length of a Sarcomere. Sarcomeres produce maximal tension when thick and thin filaments overlap between about 80 percent to 120 percent.[/caption]

Muscle Tone

Skeletal muscles are rarely completely relaxed, or flaccid. Even if a muscle is not producing movement, it is contracted a small amount to maintain its contractile proteins and produce muscle tone. The tension produced by muscle tone allows muscles to continually stabilize joints and maintain posture.

Muscle tone is accomplished by a complex interaction between the nervous system and skeletal muscles that results in the activation of a few motor units at a time, most likely in a cyclical manner. In this manner, muscles never fatigue completely, as some motor units can recover while others are active.

The absence of the low-level contractions that lead to muscle tone is referred to as hypotonia or atrophy, and can result from damage to parts of the central nervous system (CNS), such as the cerebellum, or from loss of innervations to a skeletal muscle, as in poliomyelitis. Hypotonic muscles have a flaccid appearance and display functional impairments, such as weak reflexes. Conversely, excessive muscle tone is referred to as hypertonia, accompanied by hyperreflexia (excessive reflex responses), often the result of damage to upper motor neurons in the CNS. Hypertonia can present with muscle rigidity (as seen in Parkinson’s disease) or spasticity, a phasic change in muscle tone, where a limb will “snap” back from passive stretching (as seen in some strokes).

]]> 2157 2135 5 0 10.6 Exercise and Muscle Performance https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2161 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2161

Learning Objectives

By the end of this section, you will be able to:
  • Describe hypertrophy and atrophy
  • Explain how resistance exercise builds muscle
  • Explain how performance-enhancing substances affect muscle

Physical training alters the appearance of skeletal muscles and can produce changes in muscle performance. Conversely, a lack of use can result in decreased performance and muscle appearance. Although muscle cells can change in size, new cells are not formed when muscles grow. Instead, structural proteins are added to muscle fibers in a process called hypertrophy, so cell diameter increases. The reverse, when structural proteins are lost and muscle mass decreases, is called atrophy. Age-related muscle atrophy is called sarcopenia. Cellular components of muscles can also undergo changes in response to changes in muscle use.

Endurance Exercise

Slow fibers are predominantly used in endurance exercises that require little force but involve numerous repetitions. The aerobic metabolism used by slow-twitch fibers allows them to maintain contractions over long periods. Endurance training modifies these slow fibers to make them even more efficient by producing more mitochondria to enable more aerobic metabolism and more ATP production. Endurance exercise can also increase the amount of myoglobin in a cell, as increased aerobic respiration increases the need for oxygen. Myoglobin is found in the sarcoplasm and acts as an oxygen storage supply for the mitochondria.

The training can trigger the formation of more extensive capillary networks around the fiber, a process called angiogenesis, to supply oxygen and remove metabolic waste. To allow these capillary networks to supply the deep portions of the muscle, muscle mass does not greatly increase in order to maintain a smaller area for the diffusion of nutrients and gases. All of these cellular changes result in the ability to sustain low levels of muscle contractions for greater periods without fatiguing.

The proportion of SO muscle fibers in muscle determines the suitability of that muscle for endurance, and may benefit those participating in endurance activities. Postural muscles have a large number of SO fibers and relatively few FO and FG fibers, to keep the back straight (Figure 1). Endurance athletes, like marathon-runners also would benefit from a larger proportion of SO fibers, but it is unclear if the most-successful marathoners are those with naturally high numbers of SO fibers, or whether the most successful marathon runners develop high numbers of SO fibers with repetitive training. Endurance training can result in overuse injuries such as stress fractures and joint and tendon inflammation.

[caption id="" align="aligncenter" width="380"]This photograph shows some runners in a race. Figure 1. Marathoners. Long-distance runners have a large number of SO fibers and relatively few FO and FG fibers. (credit: “Tseo2”/Wikimedia Commons)[/caption]

Resistance Exercise

Resistance exercises, as opposed to endurance exercise, require large amounts of FG fibers to produce short, powerful movements that are not repeated over long periods. The high rates of ATP hydrolysis and cross-bridge formation in FG fibers result in powerful muscle contractions. Muscles used for power have a higher ratio of FG to SO/FO fibers, and trained athletes possess even higher levels of FG fibers in their muscles. Resistance exercise affects muscles by increasing the formation of myofibrils, thereby increasing the thickness of muscle fibers. This added structure causes hypertrophy, or the enlargement of muscles, exemplified by the large skeletal muscles seen in body builders and other athletes (Figure 2). Because this muscular enlargement is achieved by the addition of structural proteins, athletes trying to build muscle mass often ingest large amounts of protein.

[caption id="" align="aligncenter" width="350"]This photograph shows a man flexing his muscles. Figure 2. Hypertrophy. Body builders have a large number of FG fibers and relatively few FO and SO fibers. (credit: Lin Mei/flickr)[/caption]

Except for the hypertrophy that follows an increase in the number of sarcomeres and myofibrils in a skeletal muscle, the cellular changes observed during endurance training do not usually occur with resistance training. There is usually no significant increase in mitochondria or capillary density. However, resistance training does increase the development of connective tissue, which adds to the overall mass of the muscle and helps to contain muscles as they produce increasingly powerful contractions. Tendons also become stronger to prevent tendon damage, as the force produced by muscles is transferred to tendons that attach the muscle to bone.

For effective strength training, the intensity of the exercise must continually be increased. For instance, continued weight lifting without increasing the weight of the load does not increase muscle size. To produce ever-greater results, the weights lifted must become increasingly heavier, making it more difficult for muscles to move the load. The muscle then adapts to this heavier load, and an even heavier load must be used if even greater muscle mass is desired.

If done improperly, resistance training can lead to overuse injuries of the muscle, tendon, or bone. These injuries can occur if the load is too heavy or if the muscles are not given sufficient time between workouts to recover or if joints are not aligned properly during the exercises. Cellular damage to muscle fibers that occurs after intense exercise includes damage to the sarcolemma and myofibrils. This muscle damage contributes to the feeling of soreness after strenuous exercise, but muscles gain mass as this damage is repaired, and additional structural proteins are added to replace the damaged ones. Overworking skeletal muscles can also lead to tendon damage and even skeletal damage if the load is too great for the muscles to bear.

Performance-Enhancing Substances

Some athletes attempt to boost their performance by using various agents that may enhance muscle performance. Anabolic steroids are one of the more widely known agents used to boost muscle mass and increase power output. Anabolic steroids are a form of testosterone, a male sex hormone that stimulates muscle formation, leading to increased muscle mass.

Endurance athletes may also try to boost the availability of oxygen to muscles to increase aerobic respiration by using substances such as erythropoietin (EPO), a hormone normally produced in the kidneys, which triggers the production of red blood cells. The extra oxygen carried by these blood cells can then be used by muscles for aerobic respiration. Human growth hormone (hGH) is another supplement, and although it can facilitate building muscle mass, its main role is to promote the healing of muscle and other tissues after strenuous exercise. Increased hGH may allow for faster recovery after muscle damage, reducing the rest required after exercise, and allowing for more sustained high-level performance.

Although performance-enhancing substances often do improve performance, most are banned by governing bodies in sports and are illegal for nonmedical purposes. Their use to enhance performance raises ethical issues of cheating because they give users an unfair advantage over nonusers. A greater concern, however, is that their use carries serious health risks. The side effects of these substances are often significant, nonreversible, and in some cases fatal. The physiological strain caused by these substances is often greater than what the body can handle, leading to effects that are unpredictable and dangerous. Anabolic steroid use has been linked to infertility, aggressive behavior, cardiovascular disease, and brain cancer.

Similarly, some athletes have used creatine to increase power output. Creatine phosphate provides quick bursts of ATP to muscles in the initial stages of contraction. Increasing the amount of creatine available to cells is thought to produce more ATP and therefore increase explosive power output, although its effectiveness as a supplement has been questioned.

Everyday Connection

Aging and Muscle Tissue Although atrophy due to disuse can often be reversed with exercise, muscle atrophy with age, referred to as sarcopenia, is irreversible. This is a primary reason why even highly trained athletes succumb to declining performance with age. This decline is noticeable in athletes whose sports require strength and powerful movements, such as sprinting, whereas the effects of age are less noticeable in endurance athletes such as marathon runners or long-distance cyclists. As muscles age, muscle fibers die, and they are replaced by connective tissue and adipose tissue (Figure 3). Because those tissues cannot contract and generate force as muscle can, muscles lose the ability to produce powerful contractions. The decline in muscle mass causes a loss of strength, including the strength required for posture and mobility. This may be caused by a reduction in FG fibers that hydrolyze ATP quickly to produce short, powerful contractions. Muscles in older people sometimes possess greater numbers of SO fibers, which are responsible for longer contractions and do not produce powerful movements. There may also be a reduction in the size of motor units, resulting in fewer fibers being stimulated and less muscle tension being produced.

[caption id="" align="aligncenter" width="325"]This image shows muscle atrophy. The left panel shows normal muscle and the right panel shows atrophied muscle. Figure 3. Atrophy. Muscle mass is reduced as muscles atrophy with disuse.[/caption]

Sarcopenia can be delayed to some extent by exercise, as training adds structural proteins and causes cellular changes that can offset the effects of atrophy. Increased exercise can produce greater numbers of cellular mitochondria, increase capillary density, and increase the mass and strength of connective tissue. The effects of age-related atrophy are especially pronounced in people who are sedentary, as the loss of muscle cells is displayed as functional impairments such as trouble with locomotion, balance, and posture. This can lead to a decrease in quality of life and medical problems, such as joint problems because the muscles that stabilize bones and joints are weakened. Problems with locomotion and balance can also cause various injuries due to falls.

]]> 2161 2135 6 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2164 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2164 [caption id="" align="aligncenter" width="500"]This photo shows a man executing a complicated yoga pose. Figure 1. A Body in Motion. The muscular system allows us to move, flex and contort our bodies. Practicing yoga, as pictured here, is a good example of the voluntary use of the muscular system. (credit: Dmitry Yanchylenko)[/caption]

Chapter Objectives

After studying this chapter, you will be able to:
  • Describe the actions and roles of agonists and antagonists
  • Explain the structure and organization of muscle fascicles and their role in generating force
  • Explain the criteria used to name skeletal muscles
  • Identify the skeletal muscles and their actions on the skeleton and soft tissues of the body
  • Identify the origins and insertions of skeletal muscles and the prime movements
Think about the things that you do each day—talking, walking, sitting, standing, and running—all of these activities require movement of particular skeletal muscles. Skeletal muscles are even used during sleep. The diaphragm is a sheet of skeletal muscle that has to contract and relax for you to breathe day and night. If you recall from your study of the skeletal system and joints, body movement occurs around the joints in the body. The focus of this chapter is on skeletal muscle organization. The system to name skeletal muscles will be explained; in some cases, the muscle is named by its shape, and in other cases it is named by its location or attachments to the skeleton. If you understand the meaning of the name of the muscle, often it will help you remember its location and/or what it does. This chapter also will describe how skeletal muscles are arranged to accomplish movement, and how other muscles may assist, or be arranged on the skeleton to resist or carry out the opposite movement. The actions of the skeletal muscles will be covered in a regional manner, working from the head down to the toes.
]]> 2164 2162 1 0 11.1 Interactions of Skeletal Muscles, Their Fascicle Arrangement, and Their Lever Systems https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2171 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2171

Learning Objectives

By the end of this section, you will be able to:
  • Compare and contrast agonist and antagonist muscles
  • Describe how fascicles are arranged within a skeletal muscle
  • Explain the major events of a skeletal muscle contraction within a muscle in generating force

To move the skeleton, the tension created by the contraction of the fibers in most skeletal muscles is transferred to the tendons. The tendons are strong bands of dense, regular connective tissue that connect muscles to bones. The bone connection is why this muscle tissue is called skeletal muscle.

Interactions of Skeletal Muscles in the Body

To pull on a bone, that is, to change the angle at its synovial joint, which essentially moves the skeleton, a skeletal muscle must also be attached to a fixed part of the skeleton. The moveable end of the muscle that attaches to the bone being pulled is called the muscle’s insertion, and the end of the muscle attached to a fixed (stabilized) bone is called the origin. During forearm flexion—bending the elbow—the brachioradialis assists the brachialis.

Although a number of muscles may be involved in an action, the principal muscle involved is called the prime mover, or agonist. To lift a cup, a muscle called the biceps brachii is actually the prime mover; however, because it can be assisted by the brachialis, the brachialis is called a synergist in this action (Figure 1). A synergist can also be a fixator that stabilizes the bone that is the attachment for the prime mover’s origin.

[caption id="" align="aligncenter" width="420"]This diagram shows two separate hands holding a glass of liquid. The biceps muscles are highlighted in pink. Figure 1. Prime Movers and Synergists. The biceps brachii flex the lower arm. The brachoradialis, in the forearm, and brachialis, located deep to the biceps in the upper arm, are both synergists that aid in this motion.[/caption]

A muscle with the opposite action of the prime mover is called an antagonist. Antagonists play two important roles in muscle function: (1) they maintain body or limb position, such as holding the arm out or standing erect; and (2) they control rapid movement, as in shadow boxing without landing a punch or the ability to check the motion of a limb.

For example, to extend the knee, a group of four muscles called the quadriceps femoris in the anterior compartment of the thigh are activated (and would be called the agonists of knee extension). However, to flex the knee joint, an opposite or antagonistic set of muscles called the hamstrings is activated.

As you can see, these terms would also be reversed for the opposing action. If you consider the first action as the knee bending, the hamstrings would be called the agonists and the quadriceps femoris would then be called the antagonists. See Table 1 for a list of some agonists and antagonists.

Agonist and Antagonist Skeletal Muscle Pairs (Table 1)
Agonist Antagonist Movement
Biceps brachii: in the anterior compartment of the arm Triceps brachii: in the posterior compartment of the arm The biceps brachii flexes the forearm, whereas the triceps brachii extends it.
Hamstrings: group of three muscles in the posterior compartment of the thigh Quadriceps femoris: group of four muscles in the anterior compartment of the thigh The hamstrings flex the leg, whereas the quadriceps femoris extend it.
Flexor digitorum superficialis and flexor digitorum profundus: in the anterior compartment of the forearm Extensor digitorum: in the posterior compartment of the forearm The flexor digitorum superficialis and flexor digitorum profundus flex the fingers and the hand at the wrist, whereas the extensor digitorum extends the fingers and the hand at the wrist.

There are also skeletal muscles that do not pull against the skeleton for movements. For example, there are the muscles that produce facial expressions. The insertions and origins of facial muscles are in the skin, so that certain individual muscles contract to form a smile or frown, form sounds or words, and raise the eyebrows. There also are skeletal muscles in the tongue, and the external urinary and anal sphincters that allow for voluntary regulation of urination and defecation, respectively. In addition, the diaphragm contracts and relaxes to change the volume of the pleural cavities but it does not move the skeleton to do this.

The Lever System of Muscle and Bone Interactions

Skeletal muscles do not work by themselves. Muscles are arranged in pairs based on their functions. For muscles attached to the bones of the skeleton, the connection determines the force, speed, and range of movement. These characteristics depend on each other and can explain the general organization of the muscular and skeletal systems.

The skeleton and muscles act together to move the body. Have you ever used the back of a hammer to remove a nail from wood? The handle acts as a lever and the head of the hammer acts as a fulcrum, the fixed point that the force is applied to when you pull back or push down on the handle. The effort applied to this system is the pulling or pushing on the handle to remove the nail, which is the load, or “resistance” to the movement of the handle in the system. Our musculoskeletal system works in a similar manner, with bones being stiff levers and the articular endings of the bones—encased in synovial joints—acting as fulcrums. The load would be an object being lifted or any resistance to a movement (your head is a load when you are lifting it), and the effort, or applied force, comes from contracting skeletal muscle.

There are several types of lever systems in the body that are identified as either first-class, second-class, or third-class levers. First-class levers are the most simple types of lever, where the balance depends on the distance between the effort and the load from the fulcrum, and the size of the load. In the body the best example of this is the way your head is raised off your chest. As shown in Figure 3 the posterior neck muscles act as the effort, the facial skeleton is the load, and the atlanto-occipital joint behaves as the fulcrum. [caption id="" align="alignnone" width="648"]image Figure 2. First class lever.[/caption]   [caption id="" align="alignnone" width="615"]image Figure 3. First class lever as seen in the body. (credit: sarahmckinnon/flickr.com, original image: celtibere/pixabay.com)[/caption] Second-class levers are levers where the load is applied between the effort and the fulcrum. The effort is closer to the load than the fulcrum, which allows a large load to be moved by a small amount of effort. However this means that the load will be moved at a slower pace, and can only be moved a short distance. Any time you stand up on your toes as shown in Figure 5, you are using a second class lever. The weight of your body acts as the load, your calf muscles are the effort, and the joints in the balls of your feet act as fulcrums. [embed]https://www.flickr.com/photos/144136128@N02/29327887125/[/embed]   [caption id="" align="alignnone" width="682"]image Figure 5. Second class lever. (credit: sarahmckinnon/flickr.com, original image: thenarratographer/flickr.com)[/caption] Third Class levers are the most common type of levers in your body. The effort is applied between the fulcrum and the load, which allows the load to be moved quickly over large distances. When you lift your hand by flexing your bicep you are using a third class lever. The Elbow joint acts as the fulcrum, the insertion of the biceps brachii becomes the effort, and the weight of your hand is the load being lifted (Figure 3). [caption id="" align="alignnone" width="647"]image Figure 6. Third class Lever.[/caption] [caption id="" align="alignnone" width="621"]image Figure 7. Third class lever. (credit: sarahmckinnon/flickr.com)[/caption]
]]> 2171 2162 2 0 Simple Second Class Lever]]> 11.3 Axial Muscles of the Head, Neck, and Back https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2177 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2177

Learning Objectives

By the end of this section, you will be able to:
  • Identify the axial muscles of the face, head, and neck
  • Identify the movement and function of the face, head, and neck muscles

The skeletal muscles are divided into axial (muscles of the trunk and head) and appendicular (muscles of the arms and legs) categories. This system reflects the bones of the skeleton system, which are also arranged in this manner. The axial muscles are grouped based on location, function, or both. Some of the axial muscles may seem to blur the boundaries because they cross over to the appendicular skeleton. The first grouping of the axial muscles you will review includes the muscles of the head and neck, then you will review the muscles of the vertebral column, and finally you will review the oblique and rectus muscles.

Muscles That Move the Head

The head, attached to the top of the vertebral column, is balanced, moved, and rotated by the neck muscles (Table 5). When these muscles act unilaterally, the head rotates. When they contract bilaterally, the head flexes or extends. The major muscle that laterally flexes and rotates the head is the sternocleidomastoid. In addition, both muscles working together are the flexors of the head. Place your fingers on both sides of the neck and turn your head to the left and to the right. You will feel the movement originate there. This muscle divides the neck into anterior and posterior triangles when viewed from the side (Figure 8).

[caption id="" align="aligncenter" width="655"]The left panel shows the lateral view of the neck. The middle panel shows the superficial neck muscles, and the right panel shows the deep neck muscles Figure 8. Posterior and Lateral Views of the Neck. The superficial and deep muscles of the neck are responsible for moving the head, cervical vertebrae, and scapulas.[/caption]
Muscles That Move the Head (Table 5)
Movement Target Target motion direction Prime mover Origin Insertion
Rotates and tilts head to the side; tilts head forward Skull; vertebrae Individually: rotates head to opposite side; bilaterally: flexion Sternocleidomastoid Sternum; clavicle Temporal bone (mastoid process); occipital bone
Rotates and tilts head backward Skull; vertebrae Individually: laterally flexes and rotates head to same side; bilaterally: extension Semispinalis capitis Transverse and articular processes of cervical and thoracic vertebra Occipital bone
Rotates and tilts head to the side; tilts head backward Skull; vertebrae Individually: laterally flexes and rotates head to same side; bilaterally: extension Splenius capitis Spinous processes of cervical and thoracic vertebra Temporal bone (mastoid process); occipital bone
Rotates and tilts head to the side; tilts head backward Skull; vertebrae Individually: laterally flexes and rotates head to same side; bilaterally: extension Longissimus capitis Transverse and articular processes of cervical and thoracic vertebra Temporal bone (mastoid process)

Muscles of the Posterior Neck and the Back

The posterior muscles of the neck are primarily concerned with head movements, like extension. The back muscles stabilize and move the vertebral column, and are grouped according to the lengths and direction of the fascicles.

The splenius muscles originate at the midline and run laterally and superiorly to their insertions. From the sides and the back of the neck, the splenius capitis inserts onto the head region, and the splenius cervicis extends onto the cervical region. These muscles can extend the head, laterally flex it, and rotate it (Figure 9).

[caption id="" align="aligncenter" width="520"]The top left panel shows a lateral view of the muscles of the neck, and the bottom left panel shows the posterior view of the superficial and deep muscles of the neck. The center panel shows the deep muscles of the back, and the right panel shows the deep spinal muscles. Figure 9. Muscles of the Neck and Back. The large, complex muscles of the neck and back move the head, shoulders, and vertebral column.[/caption]

The erector spinae group forms the majority of the muscle mass of the back and it is the primary extensor of the vertebral column. It controls flexion, lateral flexion, and rotation of the vertebral column, and maintains the lumbar curve. The erector spinae comprises the iliocostalis (laterally placed) group, the longissimus (intermediately placed) group, and the spinalis (medially placed) group.

The iliocostalis group includes the iliocostalis cervicis, associated with the cervical region; the iliocostalis thoracis, associated with the thoracic region; and the iliocostalis lumborum, associated with the lumbar region. The three muscles of the longissimus group are the longissimus capitis, associated with the head region; the longissimus cervicis, associated with the cervical region; and the longissimus thoracis, associated with the thoracic region. The third group, the spinalis group, comprises the spinalis capitis (head region), the spinalis cervicis (cervical region), and the spinalis thoracis (thoracic region).

The transversospinales muscles run from the transverse processes to the spinous processes of the vertebrae. Similar to the erector spinae muscles, the semispinalis muscles in this group are named for the areas of the body with which they are associated. The semispinalis muscles include the semispinalis capitis, the semispinalis cervicis, and the semispinalis thoracis. The multifidus muscle of the lumbar region helps extend and laterally flex the vertebral column.

Important in the stabilization of the vertebral column is the segmental muscle group, which includes the interspinales and intertransversarii muscles. These muscles bring together the spinous and transverse processes of each consecutive vertebra. Finally, the scalene muscles work together to flex, laterally flex, and rotate the head. They also contribute to deep inhalation. The scalene muscles include the anterior scalene muscle (anterior to the middle scalene), the middle scalene muscle (the longest, intermediate between the anterior and posterior scalenes), and the posterior scalene muscle (the smallest, posterior to the middle scalene).

]]> 2177 2162 4 0 11.4 Axial Muscles of the Abdominal Wall, and Thorax https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2181 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2181

Learning Objectives

By the end of this section, you will be able to:
  • Identify the intrinsic skeletal muscles of the back and neck, and the skeletal muscles of the abdominal wall and thorax
  • Identify the movement and function of the intrinsic skeletal muscles of the back and neck, and the skeletal muscles of the abdominal wall and thorax

It is a complex job to balance the body on two feet and walk upright. The muscles of the vertebral column, thorax, and abdominal wall extend, flex, and stabilize different parts of the body’s trunk. The deep muscles of the core of the body help maintain posture as well as carry out other functions. The brain sends out electrical impulses to these various muscle groups to control posture by alternate contraction and relaxation. This is necessary so that no single muscle group becomes fatigued too quickly. If any one group fails to function, body posture will be compromised.

Muscles of the Abdomen

There are four pairs of abdominal muscles that cover the anterior and lateral abdominal region and meet at the anterior midline. These muscles of the anterolateral abdominal wall can be divided into four groups: the external obliques, the internal obliques, the transversus abdominis, and the rectus abdominis (Figure 1 and Table 6).

[caption id="" align="aligncenter" width="520"]The top panel shows the lateral view of the superficial and deep abdominal muscles. The bottom panel shows the anterior view of the posterior abdominal muscles. Figure 1. Muscles of the Abdomen. (a) The anterior abdominal muscles include the medially located rectus abdominis, which is covered by a sheet of connective tissue called the rectus sheath. On the flanks of the body, medial to the rectus abdominis, the abdominal wall is composed of three layers. The external oblique muscles form the superficial layer, while the internal oblique muscles form the middle layer, and the transverses abdominus forms the deepest layer. (b) The muscles of the lower back move the lumbar spine but also assist in femur movements.[/caption]
Muscles of the Abdomen (Table 6)
Movement Target Target motion direction Prime mover Origin Insertion
Twisting at waist; also bending to the side Vertebral column Supination; lateral flexion External obliques; internal obliques Ribs 5–12; ilium Ribs 7–10; linea alba; ilium
Squeezing abdomen during forceful exhalations, defecation, urination, and childbirth Abdominal cavity Compression Transversus abdominus Ilium; ribs 5–10 Sternum; linea alba; pubis
Sitting up Vertebral column Flexion Rectus abdominis Pubis Sternum; ribs 5 and 7
Bending to the side Vertebral column Lateral flexion Quadratus lumborum Ilium; ribs 5–10 Rib 12; vertebrae L1–L4

There are three flat skeletal muscles in the antero-lateral wall of the abdomen. The external oblique, closest to the surface, extend inferiorly and medially, in the direction of sliding one’s four fingers into pants pockets. Perpendicular to it is the intermediate internal oblique, extending superiorly and medially, the direction the thumbs usually go when the other fingers are in the pants pocket. The deep muscle, the transversus abdominis, is arranged transversely around the abdomen, similar to the front of a belt on a pair of pants. This arrangement of three bands of muscles in different orientations allows various movements and rotations of the trunk. The three layers of muscle also help to protect the internal abdominal organs in an area where there is no bone.

The linea alba is a white, fibrous band that is made of the bilateral rectus sheaths that join at the anterior midline of the body. These enclose the rectus abdominis muscles (a pair of long, linear muscles, commonly called the “sit-up” muscles) that originate at the pubic crest and symphysis, and extend the length of the body’s trunk. Each muscle is segmented by three transverse bands of collagen fibers called the tendinous intersections. This results in the look of “six-pack abs,” as each segment hypertrophies on individuals at the gym who do many sit-ups.

The posterior abdominal wall is formed by the lumbar vertebrae, parts of the ilia of the hip bones, psoas major and iliacus muscles, and quadratus lumborum muscle. This part of the core plays a key role in stabilizing the rest of the body and maintaining posture.

Muscles of the Thorax

The muscles of the chest serve to facilitate breathing by changing the size of the thoracic cavity (Table 7). When you inhale, your chest rises because the cavity expands. Alternately, when you exhale, your chest falls because the thoracic cavity decreases in size.

Muscles of the Thorax (Table 7)
Movement Target Target motion direction Prime mover Origin Insertion
Inhalation; exhalation Thoracic cavity Compression; expansion Diaphragm Sternum; ribs 6–12; lumbar vertebrae Central tendon
Inhalation;exhalation Ribs Elevation (expands thoracic cavity) External intercostals Rib superior to each intercostal muscle Rib inferior to each intercostal muscle
Forced exhalation Ribs Movement along superior/inferior axis to bring ribs closer together Internal intercostals Rib inferior to each intercostal muscle Rib superior to each intercostal muscle

The Diaphragm

The change in volume of the thoracic cavity during breathing is due to the alternate contraction and relaxation of the diaphragm (Figure 2). It separates the thoracic and abdominal cavities, and is dome-shaped at rest. The superior surface of the diaphragm is convex, creating the elevated floor of the thoracic cavity. The inferior surface is concave, creating the curved roof of the abdominal cavity.

[caption id="" align="aligncenter" width="450"]This figure shows the inferior view of the diaphragm with the major parts labeled. Figure 2. Muscles of the Diaphragm. The diaphragm separates the thoracic and abdominal cavities.[/caption]

Defecating, urination, and even childbirth involve cooperation between the diaphragm and abdominal muscles (this cooperation is referred to as the “Valsalva maneuver”). You hold your breath by a steady contraction of the diaphragm; this stabilizes the volume and pressure of the peritoneal cavity. When the abdominal muscles contract, the pressure cannot push the diaphragm up, so it increases pressure on the intestinal tract (defecation), urinary tract (urination), or reproductive tract (childbirth).

The inferior surface of the pericardial sac and the inferior surfaces of the pleural membranes (parietal pleura) fuse onto the central tendon of the diaphragm. To the sides of the tendon are the skeletal muscle portions of the diaphragm, which insert into the tendon while having a number of origins including the xiphoid process of the sternum anteriorly, the inferior six ribs and their cartilages laterally, and the lumbar vertebrae and 12th ribs posteriorly.

The diaphragm also includes three openings for the passage of structures between the thorax and the abdomen. The inferior vena cava passes through the caval opening, and the esophagus and attached nerves pass through the esophageal hiatus. The aorta, thoracic duct, and azygous vein pass through the aortic hiatus of the posterior diaphragm.

The Intercostal Muscles

There are three sets of muscles, called intercostal muscles, which span each of the intercostal spaces. The principal role of the intercostal muscles is to assist in breathing by changing the dimensions of the rib cage (Figure 3).

[caption id="" align="aligncenter" width="620"]This figure shows the muscles in the thorax. The left panel shows the ribs, the major bones, and the muscles connecting them. The right panel shows a magnified view of the sternum and labels the muscles. Figure 3. Intercostal Muscles. The external intercostals are located laterally on the sides of the body. The internal intercostals are located medially near the sternum. The innermost intercostals are located deep to both the internal and external intercostals.[/caption]
The 11 pairs of superficial external intercostal muscles aid in inspiration of air during breathing because when they contract, they raise the rib cage, which expands it. The 11 pairs of internal intercostal muscles, just under the externals, are used for expiration because they draw the ribs together to constrict the rib cage. The innermost intercostal muscles are the deepest, and they act as synergists for the action of the internal intercostals.
]]> 2181 2162 5 0 11.5 Muscles of the Pectoral Girdle and Upper Limbs https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2187 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2187

Learning Objectives

By the end of this section, you will be able to:
  • Identify the muscles of the pectoral girdle and upper limbs
  • Identify the movement and function of the pectoral girdle and upper limbs

Muscles of the shoulder and upper limb can be divided into four groups: muscles that stabilize and position the pectoral girdle, muscles that move the arm, muscles that move the forearm, and muscles that move the wrists, hands, and fingers. The pectoral girdle, or shoulder girdle, consists of the lateral ends of the clavicle and scapula, along with the proximal end of the humerus, and the muscles covering these three bones to stabilize the shoulder joint. The girdle creates a base from which the head of the humerus, in its ball-and-socket joint with the glenoid fossa of the scapula, can move the arm in multiple directions.

Muscles That Position the Pectoral Girdle

Muscles that position the pectoral girdle are located either on the anterior thorax or on the posterior thorax (Figure 1 and Table 8). The anterior muscles include the subclavius, pectoralis minor, and serratus anterior. The posterior muscles include the trapezius, rhomboid major, and rhomboid minor. When the rhomboids are contracted, your scapula moves medially, which can pull the shoulder and upper limb posteriorly.

[caption id="" align="aligncenter" width="540"]The left panel shows the anterior lateral view of the pectoral girdle muscle, and the right panel shows the posterior view of the pectoral girdle muscle. Figure 1. Muscles That Position the Pectoral Girdle. The muscles that stabilize the pectoral girdle make it a steady base on which other muscles can move the arm. Note that the pectoralis major and deltoid, which move the humerus, are cut here to show the deeper positioning muscles.[/caption]
Muscles that Position the Pectoral Girdle (Table 8)
Position in the thorax Movement Target Target motion direction Prime mover Origin Insertion
Anterior thorax Stabilizes clavicle during movement by depressing it Clavicle Depression Subclavius First rib Inferior surface of clavicle
Anterior thorax Rotates shoulder anteriorly (throwing motion); assists with inhalation Scapula; ribs Scapula: depresses; ribs: elevates Pectoralis minor Anterior surfaces of certain ribs (2–4 or 3–5) Coracoid process of scapula
Anterior thorax Moves arm from side of body to front of body; assists with inhalation Scapula; ribs Scapula: protracts; ribs: elevates Serratus anterior Muscle slips from certain ribs (1–8 or 1–9) Anterior surface of vertebral border of scapula
Posterior thorax Elevates shoulders (shrugging); pulls shoulder blades together; tilts head backwards Scapula; cervical spine Scapula: rotests inferiorly, retracts, elevates, and depresses; spine: extends Trapezius Skull; vertebral column Acromion and spine of scapula; clavicle
Posterior thorax Stabilizes scapula during pectoral girdle movement Scapula Retracts; rotates inferiorly Rhomboid major Thoracic vertebrae (T2–T5) Medial border of scapula
Posterior thorax Stabilizes scapula during pectoral girdle movement Scapula Retracts; rotates inferiorly Rhomboid minor Cervical and thoracic vertebrae (C7 and T1) Medial border of scapula

Muscles That Move the Humerus

Similar to the muscles that position the pectoral girdle, muscles that cross the shoulder joint and move the humerus bone of the arm include both axial and scapular muscles (Figure 2 and Figure 3). The two axial muscles are the pectoralis major and the latissimus dorsi. The pectoralis major is thick and fan-shaped, covering much of the superior portion of the anterior thorax. The broad, triangular latissimus dorsi is located on the inferior part of the back, where it inserts into a thick connective tissue shealth called an aponeurosis.
[caption id="" align="aligncenter" width="520"]The top left panel shows the lateral view of the pectoral and back muscles. The top right panel shows the posterior view of the right deltoid and the left back muscle. The bottom left panel shows the anterior view of the deep muscles of the left shoulder, and the bottom right panel shows the deep muscles of the left shoulder. Figure 2. Muscles That Move the Humerus. (a, c) The muscles that move the humerus anteriorly are generally located on the anterior side of the body and originate from the sternum (e.g., pectoralis major) or the anterior side of the scapula (e.g., subscapularis). (b) The muscles that move the humerus superiorly generally originate from the superior surfaces of the scapula and/or the clavicle (e.g., deltoids). The muscles that move the humerus inferiorly generally originate from middle or lower back (e.g., latissiumus dorsi). (d) The muscles that move the humerus posteriorly are generally located on the posterior side of the body and insert into the scapula (e.g., infraspinatus).[/caption]
[caption id="" align="aligncenter" width="520"]This table describes the muscles that move the humerus. The pectoralis major is an axial muscle that brings the elbows together and moves the elbows up (as during an uppercut punch). It originates in the clavicle, sternum, cartilage of ribs 1 through 6 or 1 through 7, and the aponeurosis of the external oblique muscle. The latissimus dorsi is an axial muscle that moves the elbow back (as in elbowing someone standing behind you) or spreads the elbows apart. It originates in the thoracic vertebrae (T7 through T12), the lower vertebrae, ribs 9 through 12, and the iliac crest. The deltoid is a scapular muscle that lifts arms at the shoulder. It originates in the trapezius, clavicle, acromion, and spine of scapula. The subscapularis is a scapular muscle that assists the pectoralis major in bringing the elbows together and stabilizes the shoulder joint during movement of the pectoral girdle. It originates in the subscapular fossa of the scapula. The supraspinatus is a scapular muscle that rotates the elbow outwards, as during a tennis swing. It originates in the supraspinous fossa of the scapula. The infraspinatus is a scapular muscle that rotates the elbow outwards, as during a tennis swing. It originates in the infraspinous fossa of the scapula. The teres major is a scapular muscle that assists the infraspinatus in rotating the elbow outwards. It originates in the posterior surface of the scapula. The teres minor is a scapular muscle that assists the infraspinatus in rotating the elbow outwards. It originates in the lateral border of the dorsal scapular surface. The coracobra chialis is a scapular muscle that moves the elbow up and across the body, as when putting a hand on the chest. It originates in the coracoid process of the scapula. Figure 3. Muscles That Move the Humerus[/caption]
The rest of the shoulder muscles originate on the scapula. The anatomical and ligamental structure of the shoulder joint and the arrangements of the muscles covering it, allows the arm to carry out different types of movements. The deltoid, the thick muscle that creates the rounded lines of the shoulder is the major abductor of the arm, but it also facilitates flexing and medial rotation, as well as extension and lateral rotation. The subscapularis originates on the anterior scapula and medially rotates the arm. Named for their locations, the supraspinatus (superior to the spine of the scapula) and the infraspinatus (inferior to the spine of the scapula) abduct the arm, and laterally rotate the arm, respectively. The thick and flat teres major is inferior to the teres minor and extends the arm, and assists in adduction and medial rotation of it. The long teres minor laterally rotates and extends the arm. Finally, the coracobrachialis flexes and adducts the arm.

The tendons of the deep subscapularis, supraspinatus, infraspinatus, and teres minor connect the scapula to the humerus, forming the rotator cuff (musculotendinous cuff), the circle of tendons around the shoulder joint. When baseball pitchers undergo shoulder surgery it is usually on the rotator cuff, which becomes pinched and inflamed, and may tear away from the bone due to the repetitive motion of bring the arm overhead to throw a fast pitch.

Muscles That Move the Forearm

The forearm, made of the radius and ulna bones, has four main types of action at the hinge of the elbow joint: flexion, extension, pronation, and supination. The forearm flexors include the biceps brachii, brachialis, and brachioradialis. The extensors are the triceps brachii and anconeus. The pronators are the pronator teres and the pronator quadratus, and the supinator is the only one that turns the forearm anteriorly. When the forearm faces anteriorly, it is supinated. When the forearm faces posteriorly, it is pronated.

The biceps brachii, brachialis, and brachioradialis flex the forearm. The two-headed biceps brachii crosses the shoulder and elbow joints to flex the forearm, also taking part in supinating the forearm at the radioulnar joints and flexing the arm at the shoulder joint. Deep to the biceps brachii, the brachialis provides additional power in flexing the forearm. Finally, the brachioradialis can flex the forearm quickly or help lift a load slowly. These muscles and their associated blood vessels and nerves form the anterior compartment of the arm (anterior flexor compartment of the arm) (Figure 4 and Figure 5).

[caption id="" align="aligncenter" width="630"]This multipart figure shows the different muscles that move the forearm. The major muscle groups are labeled. Figure 4. Muscles That Move the Forearm. The muscles originating in the upper arm flex, extend, pronate, and supinate the forearm. The muscles originating in the forearm move the wrists, hands, and fingers.[/caption]
[caption id="" align="aligncenter" width="520"]This table describes the muscles that move the forearm. The biceps brachii are anterior muscles that perform a bicep curl; they also allow the palm of the hand to point toward the body while flexing. They originate in the coracoid process and the tubercle above the glenoid cavity. The brachialis originates in the front of the distal humerus. The brachioradialis is an anterior muscle that assists and stablizes the elbow during bicep-curl motion. It originates in the lateral supracondylar ridge at the distal end of the humerus. The triceps brachii are posterior muscles that extend the forearm, as during a punch. They originate in the infraglenoid tubercle of the scapula, the posterior shaft of the humerus, and the posterior humeral shaft distal to the radial groove. The aconeus is a posterior muscle that assists in extending the forearm; it also allows the forearm to extend away from the body. It originates in the lateral epicondyle of the humerus. The pronator teres is an anterior muscle that turns the hand palm-down. It originates in the medial epicondyle of the humerus and the coronoid process of the ulna. The pronator quadratus is an anterior muscle that assists in turning the hand palm-down. It originates in the distal portion of the anterior ulnar shaft. The supinator is a posterior muscle that turns the hand palm-down. It originates in the lateral epicondyle of the humerus and the proximal ulna. Figure 5. Muscles That Move the Forearm[/caption]
]]> 2187 2162 6 0 11.6 Appendicular Muscles of the Pelvic Girdle and Lower Limbs https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2193 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2193

Learning Objectives

By the end of this section, you will be able to:
  • Identify the appendicular muscles of the pelvic girdle and lower limb
  • Identify the movement and function of the pelvic girdle and lower limb

The appendicular muscles of the lower body position and stabilize the pelvic girdle, which serves as a foundation for the lower limbs. Comparatively, there is much more movement at the pectoral girdle than at the pelvic girdle. There is very little movement of the pelvic girdle because of its connection with the sacrum at the base of the axial skeleton. The pelvic girdle is less range of motion because it was designed to stabilize and support the body.

Muscles of the Thigh

What would happen if the pelvic girdle, which attaches the lower limbs to the torso, were capable of the same range of motion as the pectoral girdle? For one thing, walking would expend more energy if the heads of the femurs were not secured in the acetabula of the pelvis. The body’s center of gravity is in the area of the pelvis. If the center of gravity were not to remain fixed, standing up would be difficult as well. Therefore, what the leg muscles lack in range of motion and versatility, they make up for in size and power, facilitating the body’s stabilization, posture, and movement.

Gluteal Region Muscles That Move the Femur

Most muscles that insert on the femur (the thigh bone) and move it, originate on the pelvic girdle. The psoas major and iliacus make up the iliopsoas group. Some of the largest and most powerful muscles in the body are the gluteal muscles or gluteal group. The gluteus maximus is the largest; deep to the gluteus maximus is the gluteus medius, and deep to the gluteus medius is the gluteus minimus, the smallest of the trio (Figure 1 and Figure 2).

[caption id="" align="aligncenter" width="450"]The left panel shows the superficial pelvic and thigh muscles, the center panel shows the deep pelvic and thigh muscles. The right panel shows the posterior view of the pelvic and thigh muscles. Figure 1. Hip and Thigh Muscles. The large and powerful muscles of the hip that move the femur generally originate on the pelvic girdle and insert into the femur. The muscles that move the lower leg typically originate on the femur and insert into the bones of the knee joint. The anterior muscles of the femur extend the lower leg but also aid in flexing the thigh. The posterior muscles of the femur flex the lower leg but also aid in extending the thigh. A combination of gluteal and thigh muscles also adduct, abduct, and rotate the thigh and lower leg.[/caption]
[caption id="" align="aligncenter" width="550"]This table describes gluteal region muscles that move the femur. These muscles make up the iliopsoas group. The psoas major raises the knee at the hip, as if performing a knee attack; it also assists the lateral rotators in twisting the thigh (and lower leg) outward, and assists with bending over and maintaining posture. It originates in the lumbar vertebrae (L1 through L5) and thoracic vertebra (T12). The iliacus raises the knee at the hip, as if performing a knee attack; it also assists the lateral rotators in twisting the thigh (and lower leg) outward, and assists with bending over and maintaining posture. It originates in the iliac fossa, iliac crest, and lateral sacrum. These muscles make up the gluteal group. The gluteous maximus lowers the knee and moves the thigh back, as when getting ready to kick a ball. It originates in the dorsal ilium, sacrum, and coccyx. The gluteus medius opens the thigh, as when doing a split. It originates in the lateral surface of the ilium. The gluteus minimus brings the thighs back together. It originates in the external surface of the ilium. The tensor fascia lata assists with raising the knee at the hip and opening the thighs; it also maintains posture by stabilizing the iliotibial track, which connects to the knee. It originates in the anterior aspect of the iliac crest and the anterior superior iliac spine. These muscles make up the lateral rotators. The piriformis twists the thigh (and lower leg) outward; it also maintains posture by stabilizing the hip joint. It originates in the anterolateral surface of the sacrum. The obturator internus twists the thigh (and lower leg) outward; it also maintains posture by stabilizing the hip joint. It originates in the inner surface of the obturator membrane, the greater sciatic notch, and the margins of the obturator foramen. The superior gemellus twists the thigh (and lower leg) outward; it also maintains posture by stabilizing the hip joint. It originates in the ischial spine. The inferior gemellus twists the thigh (and lower leg) outward; it also maintains posture by stabilizing the hip joint. It originates in the ischial tuberosity. The quatratus femoris twists the thigh (and lower leg) outward; it also maints posture by stabilizing the hip joint. It originates in the ischial tuberosity. These muscles are adductors. The adductor longus brings the thighs back together; it also assists with raising the knee. It originates in the pubis near the pubic symphysis. The adductor brevis brings the thighs back together; it also assists with raising the knee. It originates in teh body of the pubis and in the inferior ramus of the pubis. The adductor magnus brings the thighs back together; it also assists with raising the knee and moving the thigh back. It originates in the ischial rami, the pubic rami, and the ischial tuberosity. The pectineus opens the thigh; it also assists with raising the knee and turning the thigh (and lower leg) inward. It originates in the pectineal line of the pubis. Figure 2. Gluteal Region Muscles That Move the Femur[/caption]

The tensor fascia latae is a thick, squarish muscle in the superior aspect of the lateral thigh. It acts as a synergist of the gluteus medius and iliopsoas in flexing and abducting the thigh. It also helps stabilize the lateral aspect of the knee by pulling on the iliotibial tract (band), making it taut. Deep to the gluteus maximus, the piriformis, obturator internus, obturator externus, superior gemellus, inferior gemellus, and quadratus femoris laterally rotate the femur at the hip.

The adductor longus, adductor brevis, and adductor magnus can both medially and laterally rotate the thigh depending on the placement of the foot. The adductor longus flexes the thigh, whereas the adductor magnus extends it. The pectineus adducts and flexes the femur at the hip as well. The pectineus is located in the femoral triangle, which is formed at the junction between the hip and the leg and also includes the femoral nerve, the femoral artery, the femoral vein, and the deep inguinal lymph nodes.

Thigh Muscles That Move the Femur, Tibia, and Fibula

Deep fascia in the thigh separates it into medial, anterior, and posterior compartments (see Figure 1 and Figure 3). The muscles in the medial compartment of the thigh are responsible for adducting the femur at the hip. Along with the adductor longus, adductor brevis, adductor magnus, and pectineus, the strap-like gracilis adducts the thigh in addition to flexing the leg at the knee.

[caption id="" align="aligncenter" width="520"]This table describes the thigh muscles that move the femur, tibia, and fibula. The medial compartment of the thigh consists of the gracilis, which moves the back of the lower legs up toward the buttocks, as when kneeling; it also assists in opening the thighs. It originates in the inferior ramus, the body of the pubis, and the ischial ramus. These muscles, the quadriceps femoris group, make up the anterior compartment of the thigh. The rectus femoris moves the lower leg out in front of the body, as when kicking; it also assists in raising the knee. It originates in the anterior inferior iliac spine and in the superior margin of the acetabulum. The vastus lateralis moves the lower leg out in front of the body, as when kicking. It originates in the greater trochanter, the intertrochanteric line, and the linea aspera. The vastus medialis moves the lower leg out in front of the body, as when kicking. It originates in the linea aspera and the intertrochanteric line. The vastus intermedius moves the lower leg out in front of the body, as when kicking. It originates in the proximal femur shaft. The sartorius moves the back of the lower legs up and back toward the buttocks, as when kneeling; it also assists in moving the thigh diagonally upward and outward as when mounting a bike. It originates in the anterior superior iliac spine. These muscles, the hamstring group, make up the posterior compartment of the thigh. The biceps femoris moves the back of the lower leg up and back toward the buttocks, as when kneeling; it also moves the thigh down and back and twists the thigh (and lower leg) outward. It originates in the ischial tuberosity, linea aspera, and distal femur. The semitendinosus moves the back of the lower legs up toward the buttocks, as when kneeling; it also moves the thigh down and back and twists the thigh (and lower leg) inward. It originates in the ischial tuberosity. The semi-membranosus moves the back of the lower legs up and back toward the buttocks, as when kneeling; it also moves the thigh down and back and twists the thigh (and lower leg) inward. It originates in the ischial tuberosity. Figure 3. Thigh Muscles That Move the Femur, Tibia, and Fibula[/caption]

The muscles of the anterior compartment of the thigh flex the thigh and extend the leg. This compartment contains the quadriceps femoris group, which actually comprises four muscles that extend and stabilize the knee. The rectus femoris is on the anterior aspect of the thigh, the vastus lateralis is on the lateral aspect of the thigh, the vastus medialis is on the medial aspect of the thigh, and the vastus intermedius is between the vastus lateralis and vastus medialis and deep to the rectus femoris. The tendon common to all four is the quadriceps tendon (patellar tendon), which inserts into the patella and continues below it as the patellar ligament. The patellar ligament attaches to the tibial tuberosity. In addition to the quadriceps femoris, the sartorius is a band-like muscle that extends from the anterior superior iliac spine to the medial side of the proximal tibia. This versatile muscle flexes the leg at the knee and flexes, abducts, and laterally rotates the leg at the hip. This muscle allows us to sit cross-legged.

The posterior compartment of the thigh includes muscles that flex the leg and extend the thigh. The three long muscles on the back of the knee are the hamstring group, which flexes the knee. These are the biceps femoris, semitendinosus, and semimembranosus. The tendons of these muscles form the popliteal fossa, the diamond-shaped space at the back of the knee.

Muscles That Move the Feet and Toes

Similar to the thigh muscles, the muscles of the leg are divided by deep fascia into compartments, although the leg has three: anterior, lateral, and posterior (Figure 4 and Figure 5).

[caption id="" align="aligncenter" width="530"]The left panel shows the superficial muscles that move the feet and the center panel shows the posterior view of the same muscles. The right panel shows the deep muscles of the right lower leg. Figure 4. Muscles of the Lower Leg The muscles of the anterior compartment of the lower leg are generally responsible for dorsiflexion, and the muscles of the posterior compartment of the lower leg are generally responsible for plantar flexion. The lateral and medial muscles in both compartments invert, evert, and rotate the foot.[/caption]
[caption id="" align="aligncenter" width="550"]This tables describes the muscles that move the feet and toes. These muscles make up the anterior compartment of the leg. The tibialis anterior raises the sole of the foot off the ground, as when preparing to foot-tap; it also bends the inside of the foot upwards, as when catching your balance while falling laterally toward the opposite side as the balancing foot. It originates in the lateral condyle and upper tibial shaft and in the interosseous membrane. The extensor hallucis longus raises the sole of the foot off the ground, as when preparing to foot-tap; it also extends the big toe. It originates in the anteromedial fibula shaft and interosseous membrane. The extensor digitorum longus raises the sole of the foot off the ground, as when preparing to foot-tap; it also extends the toes. It originates in the lateral condyle of the tibia, the proximal portion of the fibula, and the interosseous membrane. These muscles make up the lateral compartment of the leg. The fibularis longus lowers the sole of the foot to the ground, as when foot-tapping or jumping; it also bends the inside of the foot downwards, as when catching your balance while falling laterally toward the same side as the balancing foot. It originates in the upper portion of the lateral fibula. The fibularis (peroneus) brevis lowers the side of the foot to the ground, as when foot-tapping or jumping; it also bends the inside of the foot downward, as when catching your balance while falling laterally toward the same side as the balancing foot. It originates in the distal fibula shaft. These superficial muscles make up the posterior compartment of the leg. The gastrocnemius lowers the sole of the foot to the ground, as when foot-tapping or jumping; it also assists in moving the back of the lower legs up and back toward the buttocks. It originates in the medial and lateral condyles of the femur. The soleus lowers the sole of the foot the ground, as when foot-tapping or jumping; it also maintains posture while walking. It originates in the superior tibia, fibula, and interosseous membrane. The plantaris lowers the sole of the foot to the ground, as when foot-tapping or jumping; it also assists in moving the back of the lower legs up and back toward the buttocks. It originates in the posterior femur above the lateral condyle. The tibialis posterior lowers the sole of the foot to the ground, as when foot-tapping or jumping. It originates in the superior tibia and fibula and in the interosseous membrane. These deep muscles also make up the posterior compartment of the leg. The popliteus moves the back of the lower legs up and back toward the buttocks; it also assists in rotation of the leg at the knee and thigh. It originates in the lateral condyle of the femur and the lateral meniscus. The flexor digitorum longus lowers the sole of the foot to the ground, as when foot-tapping or jumping; it also bends the inside of the foot upward and flexes the toes. It originates in the posterior tibia. The flexor hallicis longus flexes the big toe. It originates in the midshaft of the fibula and in the interosseous membrane. Figure 5. Muscles That Move the Feet and Toes[/caption]

The muscles in the anterior compartment of the leg: the tibialis anterior, a long and thick muscle on the lateral surface of the tibia, the extensor hallucis longus, deep under it, and the extensor digitorum longus, lateral to it, all contribute to raising the front of the foot when they contract. The fibularis tertius, a small muscle that originates on the anterior surface of the fibula, is associated with the extensor digitorum longus and sometimes fused to it, but is not present in all people. Thick bands of connective tissue called the superior extensor retinaculum (transverse ligament of the ankle) and the inferior extensor retinaculum, hold the tendons of these muscles in place during dorsiflexion.

The lateral compartment of the leg includes two muscles: the fibularis longus (peroneus longus) and the fibularis brevis (peroneus brevis). The superficial muscles in the posterior compartment of the leg all insert onto the calcaneal tendon (Achilles tendon), a strong tendon that inserts into the calcaneal bone of the ankle. The muscles in this compartment are large and strong and keep humans upright. The most superficial and visible muscle of the calf is the gastrocnemius. Deep to the gastrocnemius is the wide, flat soleus. The plantaris runs obliquely between the two; some people may have two of these muscles, whereas no plantaris is observed in about seven percent of other cadaver dissections. The plantaris tendon is a desirable substitute for the fascia lata in hernia repair, tendon transplants, and repair of ligaments. There are four deep muscles in the posterior compartment of the leg as well: the popliteus, flexor digitorum longus, flexor hallucis longus, and tibialis posterior.

]]> 2193 2162 7 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2196 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2196 [caption id="" align="aligncenter" width="550"]This photo shows a red blood cell and a white blood cell. Blood Cells A single drop of blood contains millions of red blood cells, white blood cells, and platelets. One of each type is shown here, isolated from a scanning electron micrograph.[/caption]

Chapter Objectives

After studying this chapter, you will be able to:
  • Identify the primary functions of blood, its fluid and cellular components, and its physical characteristics
  • Identify the most important proteins and other solutes present in blood plasma
  • Describe the formation of the formed element components of blood
  • Discuss the structure and function of red blood cells and hemoglobin
  • Classify and characterize white blood cells
  • Describe the structure of platelets and explain the process of hemostasis
  • Explain the significance of AB and Rh blood groups in blood transfusions
  • Discuss a variety of blood disorders
Single-celled organisms do not need blood. They obtain nutrients directly from and excrete wastes directly into their environment. The human organism cannot do that. Our large, complex bodies need blood to deliver nutrients to and remove wastes from our trillions of cells. The heart pumps blood throughout the body in a network of blood vessels. Together, these three components—blood, heart, and vessels—makes up the cardiovascular system. This chapter focuses on the medium of transport: blood.]]> 2196 2194 1 0 18.2 Production of the Formed Elements https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2202 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2202

Learning Objectives

By the end of this section, you will be able to:
  • Trace the generation of the formed elements of blood from bone marrow stem cells
  • Discuss the role of hemopoietic growth factors in promoting the production of the formed elements

The lifespan of the formed elements is very brief. Although one type of leukocyte called memory cells can survive for years, most erythrocytes, leukocytes, and platelets normally live only a few hours to a few weeks. Thus, the body must form new blood cells and platelets quickly and continuously. When you donate a unit of blood during a blood drive (approximately 475 mL, or about 1 pint), your body typically replaces the donated plasma within 24 hours, but it takes about 4 to 6 weeks to replace the blood cells. This restricts the frequency with which donors can contribute their blood. The process by which this replacement occurs is called hemopoiesis, or hematopoiesis (from the Greek root haima- = “blood”; -poiesis = “production”).

Sites of Hemopoiesis

Prior to birth, hemopoiesis occurs in a number of tissues, beginning with the yolk sac of the developing embryo, and continuing in the fetal liver, spleen, lymphatic tissue, and eventually the red bone marrow. Following birth, most hemopoiesis occurs in the red marrow, a connective tissue within the spaces of spongy (cancellous) bone tissue. In children, hemopoiesis can occur in the medullary cavity of long bones; in adults, the process is largely restricted to the cranial and pelvic bones, the vertebrae, the sternum, and the proximal epiphyses of the femur and humerus.

Throughout adulthood, the liver and spleen maintain their ability to generate the formed elements. This process is referred to as extramedullary hemopoiesis (meaning hemopoiesis outside the medullary cavity of adult bones). When a disease such as bone cancer destroys the bone marrow, causing hemopoiesis to fail, extramedullary hemopoiesis may be initiated.

Differentiation of Formed Elements from Stem Cells

All formed elements arise from stem cells of the red bone marrow. Recall that stem cells undergo mitosis plus cytokinesis (cellular division) to give rise to new daughter cells: One of these remains a stem cell and the other differentiates into one of any number of diverse cell types. Stem cells may be viewed as occupying a hierarchal system, with some loss of the ability to diversify at each step. The totipotent stem cell is the zygote, or fertilized egg. The totipotent (toti- = “all”) stem cell gives rise to all cells of the human body. The next level is the pluripotent stem cell, which gives rise to multiple types of cells of the body and some of the supporting fetal membranes. Beneath this level, the mesenchymal cell is a stem cell that develops only into types of connective tissue, including fibrous connective tissue, bone, cartilage, and blood, but not epithelium, muscle, and nervous tissue. One step lower on the hierarchy of stem cells is the hemopoietic stem cell, or hemocytoblast. All of the formed elements of blood originate from this specific type of cell.

Hemopoiesis begins when the hemopoietic stem cell is exposed to appropriate chemical stimuli collectively called hemopoietic growth factors, which prompt it to divide and differentiate. One daughter cell remains a hemopoietic stem cell, allowing hemopoiesis to continue. The other daughter cell becomes either of two types of more specialized stem cells (Figure 1):

  • Lymphoid stem cells give rise to a class of leukocytes known as lymphocytes, which include the various T cells, B cells, and natural killer (NK) cells, all of which function in immunity. However, hemopoiesis of lymphocytes progresses somewhat differently from the process for the other formed elements. In brief, lymphoid stem cells quickly migrate from the bone marrow to lymphatic tissues, including the lymph nodes, spleen, and thymus, where their production and differentiation continues. B cells are so named since they mature in the bone marrow, while T cells mature in the thymus.
  • Myeloid stem cells give rise to all the other formed elements, including the erythrocytes; megakaryocytes that produce platelets; and a myeloblast lineage that gives rise to monocytes and three forms of granular leukocytes: neutrophils, eosinophils, and basophils.
[caption id="" align="aligncenter" width="600"]This flowchart shows the pathways in which a multipotent hemotopoietic stem cell differentiates into the different cell types found in blood. Figure 1. Hematopoietic System of Bone Marrow. Hemopoiesis is the proliferation and differentiation of the formed elements of blood.[/caption]

Lymphoid and myeloid stem cells do not immediately divide and differentiate into mature formed elements. As you can see in the figure above, there are several intermediate stages of precursor cells (literally, forerunner cells), many of which can be recognized by their names, which have the suffix -blast. For instance, megakaryoblasts are the precursors of megakaryocytes, and proerythroblasts become reticulocytes, which eject their nucleus and most other organelles before maturing into erythrocytes.

Hemopoietic Growth Factors

Development from stem cells to precursor cells to mature cells is again initiated by hemopoietic growth factors. These include the following:

  • Erythropoietin (EPO) is a glycoprotein hormone secreted by the interstitial fibroblast cells of the kidneys in response to low oxygen levels. It prompts the production of erythrocytes. Some athletes use synthetic EPO as a performance-enhancing drug (called blood doping) to increase RBC counts and subsequently increase oxygen delivery to tissues throughout the body. EPO is a banned substance in most organized sports, but it is also used medically in the treatment of certain anemia, specifically those triggered by certain types of cancer, and other disorders in which increased erythrocyte counts and oxygen levels are desirable.
  • Thrombopoietin, another glycoprotein hormone, is produced by the liver and kidneys. It triggers the development of megakaryocytes into platelets.
  • Cytokines are glycoproteins secreted by a wide variety of cells, including red bone marrow, leukocytes, macrophages, fibroblasts, and endothelial cells. They act locally as autocrine or paracrine factors, stimulating the proliferation of progenitor cells and helping to stimulate both nonspecific and specific resistance to disease. There are two major subtypes of cytokines known as colony-stimulating factors and interleukins.
    • Colony-stimulating factors (CSFs) are glycoproteins that act locally, as autocrine or paracrine factors. Some trigger the differentiation of myeloblasts into granular leukocytes, namely, neutrophils, eosinophils, and basophils. These are referred to as granulocyte CSFs. A different CSF induces the production of monocytes, called monocyte CSFs. Both granulocytes and monocytes are stimulated by GM-CSF; granulocytes, monocytes, platelets, and erythrocytes are stimulated by multi-CSF. Synthetic forms of these hormones are often administered to patients with various forms of cancer who are receiving chemotherapy to revive their WBC counts.
    • Interleukins are another class of cytokine signaling molecules important in hemopoiesis. They were initially thought to be secreted uniquely by leukocytes and to communicate only with other leukocytes, and were named accordingly, but are now known to be produced by a variety of cells including bone marrow and endothelium. Researchers now suspect that interleukins may play other roles in body functioning, including differentiation and maturation of cells, producing immunity and inflammation. To date, more than a dozen interleukins have been identified, with others likely to follow. They are generally numbered IL-1, IL-2, IL-3, etc.
]]> 2202 2194 3 0 18.3 Erythrocytes https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2208 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2208

Learning Objectives

By the end of this section, you will be able to:
  • Describe the anatomy of erythrocytes
  • Discuss the various steps in the lifecycle of an erythrocyte
  • Explain the composition and function of hemoglobin

The erythrocyte, commonly known as a red blood cell (or RBC), is by far the most common formed element: A single drop of blood contains millions of erythrocytes and just thousands of leukocytes. Specifically, males have about 5.4 million erythrocytes per microliter (µL) of blood, and females have approximately 4.8 million per µL. In fact, erythrocytes are estimated to make up about 25 percent of the total cells in the body. As you can imagine, they are quite small cells, with a mean diameter of only about 7–8 micrometers (µm) (Figure 1). The primary functions of erythrocytes are to pick up inhaled oxygen from the lungs and transport it to the body’s tissues, and to pick up some (about 24 percent) carbon dioxide waste at the tissues and transport it to the lungs for exhalation. Erythrocytes remain within the vascular network. Although leukocytes typically leave the blood vessels to perform their defensive functions, movement of erythrocytes from the blood vessels is abnormal.

[caption id="" align="aligncenter" width="600"]This table shows the different types of cells present in blood, the number of cells, their appearance, and a summary of their function. Figure 1. Summary of Formed Elements in Blood[/caption]

Shape and Structure of Erythrocytes

As an erythrocyte matures in the red bone marrow, it extrudes its nucleus and most of its other organelles. During the first day or two that it is in the circulation, an immature erythrocyte, known as a reticulocyte, will still typically contain remnants of organelles. Reticulocytes should comprise approximately 1–2 percent of the erythrocyte count and provide a rough estimate of the rate of RBC production, with abnormally low or high rates indicating deviations in the production of these cells. These remnants, primarily of networks (reticulum) of ribosomes, are quickly shed, however, and mature, circulating erythrocytes have few internal cellular structural components. Lacking mitochondria, for example, they rely on anaerobic respiration. This means that they do not utilize any of the oxygen they are transporting, so they can deliver it all to the tissues. They also lack endoplasmic reticula and do not synthesize proteins. Erythrocytes do, however, contain some structural proteins that help the blood cells maintain their unique structure and enable them to change their shape to squeeze through capillaries. This includes the protein spectrin, a cytoskeletal protein element.

Erythrocytes are biconcave disks; that is, they are plump at their periphery and very thin in the center (Figure 2). Since they lack most organelles, there is more interior space for the presence of the hemoglobin molecules that, as you will see shortly, transport gases. The biconcave shape also provides a greater surface area across which gas exchange can occur, relative to its volume; a sphere of a similar diameter would have a lower surface area-to-volume ratio. In the capillaries, the oxygen carried by the erythrocytes can diffuse into the plasma and then through the capillary walls to reach the cells, whereas some of the carbon dioxide produced by the cells as a waste product diffuses into the capillaries to be picked up by the erythrocytes. Capillary beds are extremely narrow, slowing the passage of the erythrocytes and providing an extended opportunity for gas exchange to occur. However, the space within capillaries can be so minute that, despite their own small size, erythrocytes may have to fold in on themselves if they are to make their way through. Fortunately, their structural proteins like spectrin are flexible, allowing them to bend over themselves to a surprising degree, then spring back again when they enter a wider vessel. In wider vessels, erythrocytes may stack up much like a roll of coins, forming a rouleaux, from the French word for “roll.”

[caption id="" align="aligncenter" width="300"]This photograph shows a few red blood cells. Figure 2. Shape of Red Blood Cells. Erythrocytes are biconcave discs with very shallow centers. This shape optimizes the ratio of surface area to volume, facilitating gas exchange. It also enables them to fold up as they move through narrow blood vessels.[/caption]

Hemoglobin

Hemoglobin is a large molecule made up of proteins and iron. It consists of four folded chains of a protein called globin, designated alpha 1 and 2, and beta 1 and 2 (Figure 3a). Each of these globin molecules is bound to a red pigment molecule called heme, which contains an ion of iron (Fe2+) (Figure 3b).

[caption id="" align="aligncenter" width="500"]This figure shows the structure of hemoglobin. The left panel shows the protein structure and the right panel shows the chemical formula. Figure 3. Hemoglobin. (a) A molecule of hemoglobin contains four globin proteins, each of which is bound to one molecule of the iron-containing pigment heme. (b) A single erythrocyte can contain 300 million hemoglobin molecules, and thus more than 1 billion oxygen molecules.[/caption]

Each iron ion in the heme can bind to one oxygen molecule; therefore, each hemoglobin molecule can transport four oxygen molecules. An individual erythrocyte may contain about 300 million hemoglobin molecules, and therefore can bind to and transport up to 1.2 billion oxygen molecules (see Figure 3b).

In the lungs, hemoglobin picks up oxygen, which binds to the iron ions, forming oxyhemoglobin. The bright red, oxygenated hemoglobin travels to the body tissues, where it releases some of the oxygen molecules, becoming darker red deoxyhemoglobin, sometimes referred to as reduced hemoglobin. Oxygen release depends on the need for oxygen in the surrounding tissues, so hemoglobin rarely if ever leaves all of its oxygen behind. In the capillaries, carbon dioxide enters the bloodstream. About 76 percent dissolves in the plasma, some of it remaining as dissolved CO2, and the remainder forming bicarbonate ion. About 23–24 percent of it binds to the amino acids in hemoglobin, forming a molecule known as carbaminohemoglobin. From the capillaries, the hemoglobin carries carbon dioxide back to the lungs, where it releases it for exchange of oxygen.

Changes in the levels of RBCs can have significant effects on the body’s ability to effectively deliver oxygen to the tissues. Ineffective hematopoiesis results in insufficient numbers of RBCs and results in one of several forms of anemia. An overproduction of RBCs produces a condition called polycythemia. The primary drawback with polycythemia is not a failure to directly deliver enough oxygen to the tissues, but rather the increased viscosity of the blood, which makes it more difficult for the heart to circulate the blood.

In patients with insufficient hemoglobin, the tissues may not receive sufficient oxygen, resulting in another form of anemia. In determining oxygenation of tissues, the value of greatest interest in healthcare is the percent saturation; that is, the percentage of hemoglobin sites occupied by oxygen in a patient’s blood. Clinically this value is commonly referred to simply as “percent sat.”

Percent saturation is normally monitored using a device known as a pulse oximeter, which is applied to a thin part of the body, typically the tip of the patient’s finger. The device works by sending two different wavelengths of light (one red, the other infrared) through the finger and measuring the light with a photodetector as it exits. Hemoglobin absorbs light differentially depending upon its saturation with oxygen. The machine calibrates the amount of light received by the photodetector against the amount absorbed by the partially oxygenated hemoglobin and presents the data as percent saturation. Normal pulse oximeter readings range from 95–100 percent. Lower percentages reflect hypoxemia, or low blood oxygen. The term hypoxia is more generic and simply refers to low oxygen levels. Oxygen levels are also directly monitored from free oxygen in the plasma typically following an arterial stick. When this method is applied, the amount of oxygen present is expressed in terms of partial pressure of oxygen or simply pO2 and is typically recorded in units of millimeters of mercury, mm Hg.

The kidneys filter about 180 liters (~380 pints) of blood in an average adult each day, or about 20 percent of the total resting volume, and thus serve as ideal sites for receptors that determine oxygen saturation. In response to hypoxemia, less oxygen will exit the vessels supplying the kidney, resulting in hypoxia (low oxygen concentration) in the tissue fluid of the kidney where oxygen concentration is actually monitored. Interstitial fibroblasts within the kidney secrete EPO, thereby increasing erythrocyte production and restoring oxygen levels. In a classic negative-feedback loop, as oxygen saturation rises, EPO secretion falls, and vice versa, thereby maintaining homeostasis. Populations dwelling at high elevations, with inherently lower levels of oxygen in the atmosphere, naturally maintain a hematocrit higher than people living at sea level. Consequently, people traveling to high elevations may experience symptoms of hypoxemia, such as fatigue, headache, and shortness of breath, for a few days after their arrival. In response to the hypoxemia, the kidneys secrete EPO to step up the production of erythrocytes until homeostasis is achieved once again. To avoid the symptoms of hypoxemia, or altitude sickness, mountain climbers typically rest for several days to a week or more at a series of camps situated at increasing elevations to allow EPO levels and, consequently, erythrocyte counts to rise. When climbing the tallest peaks, such as Mt. Everest and K2 in the Himalayas, many mountain climbers rely upon bottled oxygen as they near the summit.

Lifecycle of Erythrocytes

Production of erythrocytes in the marrow occurs at the staggering rate of more than 2 million cells per second. For this production to occur, a number of raw materials must be present in adequate amounts. These include the same nutrients that are essential to the production and maintenance of any cell, such as glucose, lipids, and amino acids. However, erythrocyte production also requires several trace elements:

  • Iron. We have said that each heme group in a hemoglobin molecule contains an ion of the trace mineral iron. On average, less than 20 percent of the iron we consume is absorbed. Heme iron, from animal foods such as meat, poultry, and fish, is absorbed more efficiently than non-heme iron from plant foods. Upon absorption, iron becomes part of the body’s total iron pool. The bone marrow, liver, and spleen can store iron in the protein compounds ferritin and hemosiderin. Ferroportin transports the iron across the intestinal cell plasma membranes and from its storage sites into tissue fluid where it enters the blood. When EPO stimulates the production of erythrocytes, iron is released from storage, bound to transferrin, and carried to the red marrow where it attaches to erythrocyte precursors.
  • Copper. A trace mineral, copper is a component of two plasma proteins, hephaestin and ceruloplasmin. Without these, hemoglobin could not be adequately produced. Located in intestinal villi, hephaestin enables iron to be absorbed by intestinal cells. Ceruloplasmin transports copper. Both enable the oxidation of iron from Fe2+ to Fe3+, a form in which it can be bound to its transport protein, transferrin, for transport to body cells. In a state of copper deficiency, the transport of iron for heme synthesis decreases, and iron can accumulate in tissues, where it can eventually lead to organ damage.
  • Zinc. The trace mineral zinc functions as a co-enzyme that facilitates the synthesis of the heme portion of hemoglobin.
  • B vitamins. The B vitamins folate and vitamin B12 function as co-enzymes that facilitate DNA synthesis. Thus, both are critical for the synthesis of new cells, including erythrocytes.

Erythrocytes live up to 120 days in the circulation, after which the worn-out cells are removed by a type of myeloid phagocytic cell called a macrophage, located primarily within the bone marrow, liver, and spleen. The components of the degraded erythrocytes’ hemoglobin are further processed as follows:

  • Globin, the protein portion of hemoglobin, is broken down into amino acids, which can be sent back to the bone marrow to be used in the production of new erythrocytes. Hemoglobin that is not phagocytized is broken down in the circulation, releasing alpha and beta chains that are removed from circulation by the kidneys.
  • The iron contained in the heme portion of hemoglobin may be stored in the liver or spleen, primarily in the form of ferritin or hemosiderin, or carried through the bloodstream by transferrin to the red bone marrow for recycling into new erythrocytes.
  • The non-iron portion of heme is degraded into the waste product biliverdin, a green pigment, and then into another waste product, bilirubin, a yellow pigment. Bilirubin binds to albumin and travels in the blood to the liver, which uses it in the manufacture of bile, a compound released into the intestines to help emulsify dietary fats. In the large intestine, bacteria breaks the bilirubin apart from the bile and converts it to urobilinogen and then into stercobilin. It is then eliminated from the body in the feces. Broad-spectrum antibiotics typically eliminate these bacteria as well and may alter the color of feces. The kidneys also remove any circulating bilirubin and other related metabolic byproducts such as urobilins and secrete them into the urine.

The breakdown pigments formed from the destruction of hemoglobin can be seen in a variety of situations. At the site of an injury, biliverdin from damaged RBCs produces some of the dramatic colors associated with bruising. With a failing liver, bilirubin cannot be removed effectively from circulation and causes the body to assume a yellowish tinge associated with jaundice. Stercobilins within the feces produce the typical brown color associated with this waste. And the yellow of urine is associated with the urobilins.

The erythrocyte lifecycle is summarized in Figure 4.

[caption id="" align="aligncenter" width="600"]This flow chart shows the life cycle of a red blood cell. The first step is the hemopoeisis of erythrocytes in the bone marrow. Further steps in this diagram show the passage of erythrocytes through the blood stream, the breakdown of heme protein, and liver function. Figure 4. Erythrocyte Lifecycle. Erythrocytes are produced in the bone marrow and sent into the circulation. At the end of their lifecycle, they are destroyed by macrophages, and their components are recycled.[/caption]

Disorders of Erythrocytes

The size, shape, and number of erythrocytes, and the number of hemoglobin molecules can have a major impact on a person’s health. When the number of RBCs or hemoglobin is deficient, the general condition is called anemia. There are more than 400 types of anemia and more than 3.5 million Americans suffer from this condition. Anemia can be broken down into three major groups: those caused by blood loss, those caused by faulty or decreased RBC production, and those caused by excessive destruction of RBCs. Clinicians often use two groupings in diagnosis: The kinetic approach focuses on evaluating the production, destruction, and removal of RBCs, whereas the morphological approach examines the RBCs themselves, paying particular emphasis to their size. A common test is the mean corpuscle volume (MCV), which measures size. Normal-sized cells are referred to as normocytic, smaller-than-normal cells are referred to as microcytic, and larger-than-normal cells are referred to as macrocytic. Reticulocyte counts are also important and may reveal inadequate production of RBCs. The effects of the various anemias are widespread, because reduced numbers of RBCs or hemoglobin will result in lower levels of oxygen being delivered to body tissues. Since oxygen is required for tissue functioning, anemia produces fatigue, lethargy, and an increased risk for infection. An oxygen deficit in the brain impairs the ability to think clearly, and may prompt headaches and irritability. Lack of oxygen leaves the patient short of breath, even as the heart and lungs work harder in response to the deficit.

Blood loss anemias are fairly straightforward. In addition to bleeding from wounds or other lesions, these forms of anemia may be due to ulcers, hemorrhoids, inflammation of the stomach (gastritis), and some cancers of the gastrointestinal tract. The excessive use of aspirin or other nonsteroidal anti-inflammatory drugs such as ibuprofen can trigger ulceration and gastritis. Excessive menstruation and loss of blood during childbirth are also potential causes.

Anemias caused by faulty or decreased RBC production include sickle cell anemia, iron deficiency anemia, vitamin deficiency anemia, and diseases of the bone marrow and stem cells.

  • A characteristic change in the shape of erythrocytes is seen in sickle cell disease (also referred to as sickle cell anemia). A genetic disorder, it is caused by production of an abnormal type of hemoglobin, called hemoglobin S, which delivers less oxygen to tissues and causes erythrocytes to assume a sickle (or crescent) shape, especially at low oxygen concentrations (Figure 5). These abnormally shaped cells can then become lodged in narrow capillaries because they are unable to fold in on themselves to squeeze through, blocking blood flow to tissues and causing a variety of serious problems from painful joints to delayed growth and even blindness and cerebrovascular accidents (strokes). Sickle cell anemia is a genetic condition particularly found in individuals of African descent.
[caption id="" align="aligncenter" width="300"]This photograph shows red blood cells of a person suffering from sickle cell anemia. Instead of being discoid shaped like healthy blood cells, sickle red blood cells are shaped like a sickle. Figure 5. Sickle Cells. Sickle cell anemia is caused by a mutation in one of the hemoglobin genes. Erythrocytes produce an abnormal type of hemoglobin, which causes the cell to take on a sickle or crescent shape. (credit: Janice Haney Carr)[/caption]
  • Iron deficiency anemia is the most common type and results when the amount of available iron is insufficient to allow production of sufficient heme. This condition can occur in individuals with a deficiency of iron in the diet and is especially common in teens and children as well as in vegans and vegetarians. Additionally, iron deficiency anemia may be caused by either an inability to absorb and transport iron or slow, chronic bleeding.
  • Vitamin-deficient anemias generally involve insufficient vitamin B12 and folate.
    • Megaloblastic anemia involves a deficiency of vitamin B12 and/or folate, and often involves diets deficient in these essential nutrients. Lack of meat or a viable alternate source, and overcooking or eating insufficient amounts of vegetables may lead to a lack of folate.
    • Pernicious anemia is caused by poor absorption of vitamin B12 and is often seen in patients with Crohn’s disease (a severe intestinal disorder often treated by surgery), surgical removal of the intestines or stomach (common in some weight loss surgeries), intestinal parasites, and AIDS.
    • Pregnancies, some medications, excessive alcohol consumption, and some diseases such as celiac disease are also associated with vitamin deficiencies. It is essential to provide sufficient folic acid during the early stages of pregnancy to reduce the risk of neurological defects, including spina bifida, a failure of the neural tube to close.
  • Assorted disease processes can also interfere with the production and formation of RBCs and hemoglobin. If myeloid stem cells are defective or replaced by cancer cells, there will be insufficient quantities of RBCs produced.
    • Aplastic anemia is the condition in which there are deficient numbers of RBC stem cells. Aplastic anemia is often inherited, or it may be triggered by radiation, medication, chemotherapy, or infection.
    • Thalassemia is an inherited condition typically occurring in individuals from the Middle East, the Mediterranean, African, and Southeast Asia, in which maturation of the RBCs does not proceed normally. The most severe form is called Cooley’s anemia.
    • Lead exposure from industrial sources or even dust from paint chips of iron-containing paints or pottery that has not been properly glazed may also lead to destruction of the red marrow.
  • Various disease processes also can lead to anemias. These include chronic kidney diseases often associated with a decreased production of EPO, hypothyroidism, some forms of cancer, lupus, and rheumatoid arthritis.

In contrast to anemia, an elevated RBC count is called polycythemia and is detected in a patient’s elevated hematocrit. It can occur transiently in a person who is dehydrated; when water intake is inadequate or water losses are excessive, the plasma volume falls. As a result, the hematocrit rises. For reasons mentioned earlier, a mild form of polycythemia is chronic but normal in people living at high altitudes. Some elite athletes train at high elevations specifically to induce this phenomenon. Finally, a type of bone marrow disease called polycythemia vera (from the Greek vera = “true”) causes an excessive production of immature erythrocytes. Polycythemia vera can dangerously elevate the viscosity of blood, raising blood pressure and making it more difficult for the heart to pump blood throughout the body. It is a relatively rare disease that occurs more often in men than women, and is more likely to be present in elderly patients those over 60 years of age.

]]> 2208 2194 4 0 18.4 Leukocytes and Platelets https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2214 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2214

Learning Objectives

By the end of this section, you will be able to:
  • Describe the general characteristics of leukocytes
  • Classify leukocytes according to their lineage, their main structural features, and their primary functions
  • Discuss the most common malignancies involving leukocytes
  • Identify the lineage, basic structure, and function of platelets

The leukocyte, commonly known as a white blood cell (or WBC), is a major component of the body’s defenses against disease. Leukocytes protect the body against invading microorganisms and body cells with mutated DNA, and they clean up debris. Platelets are essential for the repair of blood vessels when damage to them has occurred; they also provide growth factors for healing and repair.

Characteristics of Leukocytes

Although leukocytes and erythrocytes both originate from hematopoietic stem cells in the bone marrow, they are very different from each other in many significant ways. For instance, leukocytes are far less numerous than erythrocytes: Typically there are only 5000 to 10,000 per µL. They are also larger than erythrocytes and are the only formed elements that are complete cells, possessing a nucleus and organelles. And although there is just one type of erythrocyte, there are many types of leukocytes. Most of these types have a much shorter lifespan than that of erythrocytes, some as short as a few hours or even a few minutes in the case of acute infection.

One of the most distinctive characteristics of leukocytes is their movement. Whereas erythrocytes spend their days circulating within the blood vessels, leukocytes routinely leave the bloodstream to perform their defensive functions in the body’s tissues. For leukocytes, the vascular network is simply a highway they travel and soon exit to reach their true destination. When they arrive, they are often given distinct names, such as macrophage or microglia, depending on their function. As shown in Figure 1, they leave the capillaries—the smallest blood vessels—or other small vessels through a process known as emigration (from the Latin for “removal”) or diapedesis (dia- = “through”; -pedan = “to leap”) in which they squeeze through adjacent cells in a blood vessel wall.

Once they have exited the capillaries, some leukocytes will take up fixed positions in lymphatic tissue, bone marrow, the spleen, the thymus, or other organs. Others will move about through the tissue spaces very much like amoebas, continuously extending their plasma membranes, sometimes wandering freely, and sometimes moving toward the direction in which they are drawn by chemical signals. This attracting of leukocytes occurs because of positive chemotaxis (literally “movement in response to chemicals”), a phenomenon in which injured or infected cells and nearby leukocytes emit the equivalent of a chemical “911” call, attracting more leukocytes to the site. In clinical medicine, the differential counts of the types and percentages of leukocytes present are often key indicators in making a diagnosis and selecting a treatment.

[caption id="" align="aligncenter" width="600"]This figure shows how leukocytes respond to chemical signals from injured cells. The top panel shows chemical signals sent out by the injured cells. The middle panel shows leukocytes migrating to the injured cells. The bottom panel shows macrophages phagocytosing the pathogens. Figure 1. Emigration. Leukocytes exit the blood vessel and then move through the connective tissue of the dermis toward the site of a wound. Some leukocytes, such as the eosinophil and neutrophil, are characterized as granular leukocytes. They release chemicals from their granules that destroy pathogens; they are also capable of phagocytosis. The monocyte, an agranular leukocyte, differentiates into a macrophage that then phagocytizes the pathogens.[/caption]

Classification of Leukocytes

When scientists first began to observe stained blood slides, it quickly became evident that leukocytes could be divided into two groups, according to whether their cytoplasm contained highly visible granules:

  • Granular leukocytes contain abundant granules within the cytoplasm. They include neutrophils, eosinophils, and basophils (you can view their lineage from myeloid stem cells in Chapter 18.2 Production of Formed Elements).
  • While granules are not totally lacking in agranular leukocytes, they are far fewer and less obvious. Agranular leukocytes include monocytes, which mature into macrophages that are phagocytic, and lymphocytes, which arise from the lymphoid stem cell line.

Granular Leukocytes

We will consider the granular leukocytes in order from most common to least common. All of these are produced in the red bone marrow and have a short lifespan of hours to days. They typically have a lobed nucleus and are classified according to which type of stain best highlights their granules (Figure 2).

[caption id="" align="aligncenter" width="380"]The left image shows a neutrophil, the middle image shows an eosinophil, and the right image shows a basophil. Figure 2. Granular Leukocytes. A neutrophil has small granules that stain light lilac and a nucleus with two to five lobes. An eosinophil’s granules are slightly larger and stain reddish-orange, and its nucleus has two to three lobes. A basophil has large granules that stain dark blue to purple and a two-lobed nucleus.[/caption]

The most common of all the leukocytes, neutrophils will normally comprise 50–70 percent of total leukocyte count. They are 10–12 µm in diameter, significantly larger than erythrocytes. They are called neutrophils because their granules show up most clearly with stains that are chemically neutral (neither acidic nor basic). The granules are numerous but quite fine and normally appear light lilac. The nucleus has a distinct lobed appearance and may have two to five lobes, the number increasing with the age of the cell. Older neutrophils have increasing numbers of lobes and are often referred to as polymorphonuclear (a nucleus with many forms), or simply “polys.” Younger and immature neutrophils begin to develop lobes and are known as “bands.”

Neutrophils are rapid responders to the site of infection and are efficient phagocytes with a preference for bacteria. Their granules include lysozyme, an enzyme capable of lysing, or breaking down, bacterial cell walls; oxidants such as hydrogen peroxide; and defensins, proteins that bind to and puncture bacterial and fungal plasma membranes, so that the cell contents leak out. Abnormally high counts of neutrophils indicate infection and/or inflammation, particularly triggered by bacteria, but are also found in burn patients and others experiencing unusual stress. A burn injury increases the proliferation of neutrophils in order to fight off infection that can result from the destruction of the barrier of the skin. Low counts may be caused by drug toxicity and other disorders, and may increase an individual’s susceptibility to infection.

Eosinophils typically represent 2–4 percent of total leukocyte count. They are also 10–12 µm in diameter. The granules of eosinophils stain best with an acidic stain known as eosin. The nucleus of the eosinophil will typically have two to three lobes and, if stained properly, the granules will have a distinct red to orange color.

The granules of eosinophils include antihistamine molecules, which counteract the activities of histamines, inflammatory chemicals produced by basophils and mast cells. Some eosinophil granules contain molecules toxic to parasitic worms, which can enter the body through the integument, or when an individual consumes raw or undercooked fish or meat. Eosinophils are also capable of phagocytosis and are particularly effective when antibodies bind to the target and form an antigen-antibody complex. High counts of eosinophils are typical of patients experiencing allergies, parasitic worm infestations, and some autoimmune diseases. Low counts may be due to drug toxicity and stress.

Basophils are the least common leukocytes, typically comprising less than one percent of the total leukocyte count. They are slightly smaller than neutrophils and eosinophils at 8–10 µm in diameter. The granules of basophils stain best with basic (alkaline) stains. Basophils contain large granules that pick up a dark blue stain and are so common they may make it difficult to see the two-lobed nucleus.

In general, basophils intensify the inflammatory response. They share this trait with mast cells. In the past, mast cells were considered to be basophils that left the circulation. However, this appears not to be the case, as the two cell types develop from different lineages.

The granules of basophils release histamines, which contribute to inflammation, and heparin, which opposes blood clotting. High counts of basophils are associated with allergies, parasitic infections, and hypothyroidism. Low counts are associated with pregnancy, stress, and hyperthyroidism.

Agranular Leukocytes

Agranular leukocytes contain smaller, less-visible granules in their cytoplasm than do granular leukocytes. The nucleus is simple in shape, sometimes with an indentation but without distinct lobes. There are two major types of agranulocytes: lymphocytes and monocytes.

Lymphocytes are the only formed element of blood that arises from lymphoid stem cells. Although they form initially in the bone marrow, much of their subsequent development and reproduction occurs in the lymphatic tissues. Lymphocytes are the second most common type of leukocyte, accounting for about 20–30 percent of all leukocytes, and are essential for the immune response. The size range of lymphocytes is quite extensive, with some authorities recognizing two size classes and others three. Typically, the large cells are 10–14 µm and have a smaller nucleus-to-cytoplasm ratio and more granules. The smaller cells are typically 6–9 µm with a larger volume of nucleus to cytoplasm, creating a “halo” effect. A few cells may fall outside these ranges, at 14–17 µm. This finding has led to the three size range classification.

The three major groups of lymphocytes include natural killer cells, B cells, and T cells. Natural killer (NK) cells are capable of recognizing cells that do not express “self” proteins on their plasma membrane or that contain foreign or abnormal markers. These “nonself” cells include cancer cells, cells infected with a virus, and other cells with atypical surface proteins. Thus, they provide generalized, nonspecific immunity. The larger lymphocytes are typically NK cells.

B cells and T cells, also called B lymphocytes and T lymphocytes, play prominent roles in defending the body against specific pathogens (disease-causing microorganisms) and are involved in specific immunity. One form of B cells (plasma cells) produces the antibodies or immunoglobulins that bind to specific foreign or abnormal components of plasma membranes. This is also referred to as humoral (body fluid) immunity. T cells provide cellular-level immunity by physically attacking foreign or diseased cells. A memory cell is a variety of both B and T cells that forms after exposure to a pathogen and mounts rapid responses upon subsequent exposures. Unlike other leukocytes, memory cells live for many years. B cells undergo a maturation process in the bone marrow, whereas T cells undergo maturation in the thymus. This site of the maturation process gives rise to the name B and T cells. The functions of lymphocytes are complex and will be covered in detail in the chapter covering the lymphatic system and immunity. Smaller lymphocytes are either B or T cells, although they cannot be differentiated in a normal blood smear.

Abnormally high lymphocyte counts are characteristic of viral infections as well as some types of cancer. Abnormally low lymphocyte counts are characteristic of prolonged (chronic) illness or immunosuppression, including that caused by HIV infection and drug therapies that often involve steroids.

Monocytes originate from myeloid stem cells. They normally represent 2–8 percent of the total leukocyte count. They are typically easily recognized by their large size of 12–20 µm and indented or horseshoe-shaped nuclei. Macrophages are monocytes that have left the circulation and phagocytize debris, foreign pathogens, worn-out erythrocytes, and many other dead, worn out, or damaged cells. Macrophages also release antimicrobial defensins and chemotactic chemicals that attract other leukocytes to the site of an infection. Some macrophages occupy fixed locations, whereas others wander through the tissue fluid.

Abnormally high counts of monocytes are associated with viral or fungal infections, tuberculosis, and some forms of leukemia and other chronic diseases. Abnormally low counts are typically caused by suppression of the bone marrow.

Lifecycle of Leukocytes

Most leukocytes have a relatively short lifespan, typically measured in hours or days. Production of all leukocytes begins in the bone marrow under the influence of CSFs and interleukins. Secondary production and maturation of lymphocytes occurs in specific regions of lymphatic tissue known as germinal centers. Lymphocytes are fully capable of mitosis and may produce clones of cells with identical properties. This capacity enables an individual to maintain immunity throughout life to many threats that have been encountered in the past.

Disorders of Leukocytes

Leukopenia is a condition in which too few leukocytes are produced. If this condition is pronounced, the individual may be unable to ward off disease. Excessive leukocyte proliferation is known as leukocytosis. Although leukocyte counts are high, the cells themselves are often nonfunctional, leaving the individual at increased risk for disease.

Leukemia is a cancer involving an abundance of leukocytes. It may involve only one specific type of leukocyte from either the myeloid line (myelocytic leukemia) or the lymphoid line (lymphocytic leukemia). In chronic leukemia, mature leukocytes accumulate and fail to die. In acute leukemia, there is an overproduction of young, immature leukocytes. In both conditions the cells do not function properly.

Lymphoma is a form of cancer in which masses of malignant T and/or B lymphocytes collect in lymph nodes, the spleen, the liver, and other tissues. As in leukemia, the malignant leukocytes do not function properly, and the patient is vulnerable to infection. Some forms of lymphoma tend to progress slowly and respond well to treatment. Others tend to progress quickly and require aggressive treatment, without which they are rapidly fatal.

Platelets

You may occasionally see platelets referred to as thrombocytes, but because this name suggests they are a type of cell, it is not accurate. A platelet is not a cell but rather a fragment of the cytoplasm of a cell called a megakaryocyte that is surrounded by a plasma membrane. Megakaryocytes are descended from myeloid stem cells (see Chapter 18.2 Production of the Formed Elements) and are large, typically 50–100 µm in diameter, and contain an enlarged, lobed nucleus. As noted earlier, thrombopoietin, a glycoprotein secreted by the kidneys and liver, stimulates the proliferation of megakaryoblasts, which mature into megakaryocytes. These remain within bone marrow tissue (Figure 3) and ultimately form platelet-precursor extensions that extend through the walls of bone marrow capillaries to release into the circulation thousands of cytoplasmic fragments, each enclosed by a bit of plasma membrane. These enclosed fragments are platelets. Each megakarocyte releases 2000–3000 platelets during its lifespan. Following platelet release, megakaryocyte remnants, which are little more than a cell nucleus, are consumed by macrophages.

Platelets are relatively small, 2–4 µm in diameter, but numerous, with typically 150,000–160,000 per µL of blood. After entering the circulation, approximately one-third migrate to the spleen for storage for later release in response to any rupture in a blood vessel. They then become activated to perform their primary function, which is to limit blood loss. Platelets remain only about 10 days, then are phagocytized by macrophages.

Platelets are critical to hemostasis, the stoppage of blood flow following damage to a vessel. They also secrete a variety of growth factors essential for growth and repair of tissue, particularly connective tissue. Infusions of concentrated platelets are now being used in some therapies to stimulate healing.

Disorders of Platelets

Thrombocytosis is a condition in which there are too many platelets. This may trigger formation of unwanted blood clots (thrombosis), a potentially fatal disorder. If there is an insufficient number of platelets, called thrombocytopenia, blood may not clot properly, and excessive bleeding may result.
[caption id="" align="aligncenter" width="200"]This flowchart shows a myeloid stem cell differentiating into platelets. Figure 3. Platelets. Platelets are derived from cells called megakaryocytes.[/caption]
[caption id="" align="aligncenter" width="120"]QR Code representing a URL View University of Michigan Webscopes at http://virtualslides.med.umich.edu/Histology/Cardiovascular%20System/081-2_HISTO_40X.svs/view.apml?cwidth=860&cheight=733&chost=virtualslides.med.umich.edu&listview=1&title=&csis=1 and explore the blood slides in greater detail.[/caption]
[caption id="" align="aligncenter" width="380"]This figure shows micrographs of the different types of leukocytes. From left to right, the order of leukocytes shown are: basophil, eosinophil, neutrophil, monocyte, and lymphocyte. Figure 4. Leukocytes. (Micrographs provided by the Regents of University of Michigan Medical School © 2012)[/caption]
]]> 2214 2194 5 0 18.5 Hemostasis https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2217 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2217

Learning Objectives

By the end of this section, you will be able to:
  • Describe the three mechanisms involved in hemostasis
  • Explain how the extrinsic and intrinsic coagulation pathways lead to the common pathway, and the coagulation factors involved in each
  • Discuss disorders affecting hemostasis

Platelets are key players in hemostasis, the process by which the body seals a ruptured blood vessel and prevents further loss of blood. Although rupture of larger vessels usually requires medical intervention, hemostasis is quite effective in dealing with small, simple wounds. There are three steps to the process: vascular spasm, the formation of a platelet plug, and coagulation (blood clotting). Failure of any of these steps will result in hemorrhage—excessive bleeding.

Vascular Spasm

When a vessel is severed or punctured, or when the wall of a vessel is damaged, vascular spasm occurs. In vascular spasm, the smooth muscle in the walls of the vessel contracts dramatically. This smooth muscle has both circular layers; larger vessels also have longitudinal layers. The circular layers tend to constrict the flow of blood, whereas the longitudinal layers, when present, draw the vessel back into the surrounding tissue, often making it more difficult for a surgeon to locate, clamp, and tie off a severed vessel. The vascular spasm response is believed to be triggered by several chemicals called endothelins that are released by vessel-lining cells and by pain receptors in response to vessel injury. This phenomenon typically lasts for up to 30 minutes, although it can last for hours.

Formation of the Platelet Plug

In the second step, platelets, which normally float free in the plasma, encounter the area of vessel rupture with the exposed underlying connective tissue and collagenous fibers. The platelets begin to clump together, become spiked and sticky, and bind to the exposed collagen and endothelial lining. This process is assisted by a glycoprotein in the blood plasma called von Willebrand factor, which helps stabilize the growing platelet plug. As platelets collect, they simultaneously release chemicals from their granules into the plasma that further contribute to hemostasis. Among the substances released by the platelets are:

  • adenosine diphosphate (ADP), which helps additional platelets to adhere to the injury site, reinforcing and expanding the platelet plug
  • serotonin, which maintains vasoconstriction
  • prostaglandins and phospholipids, which also maintain vasoconstriction and help to activate further clotting chemicals, as discussed next

A platelet plug can temporarily seal a small opening in a blood vessel. Plug formation, in essence, buys the body time while more sophisticated and durable repairs are being made. In a similar manner, even modern naval warships still carry an assortment of wooden plugs to temporarily repair small breaches in their hulls until permanent repairs can be made.

Coagulation

Those more sophisticated and more durable repairs are collectively called coagulation, the formation of a blood clot. The process is sometimes characterized as a cascade, because one event prompts the next as in a multi-level waterfall. The result is the production of a gelatinous but robust clot made up of a mesh of fibrin—an insoluble filamentous protein derived from fibrinogen, the plasma protein introduced earlier—in which platelets and blood cells are trapped. Figure 1 summarizes the three steps of hemostasis.

[caption id="" align="aligncenter" width="600"]This figure details the steps in the clotting of blood. Each step is shown along with a detailed text box describing the steps on the left. On the right, a signaling pathway shows the different chemical signals involved in the clotting process. Figure 1. Hemostasis. (a) An injury to a blood vessel initiates the process of hemostasis. Blood clotting involves three steps. First, vascular spasm constricts the flow of blood. Next, a platelet plug forms to temporarily seal small openings in the vessel. Coagulation then enables the repair of the vessel wall once the leakage of blood has stopped. (b) The synthesis of fibrin in blood clots involves either an intrinsic pathway or an extrinsic pathway, both of which lead to a common pathway. (credit a: Kevin MacKenzie)[/caption]

Clotting Factors Involved in Coagulation

In the coagulation cascade, chemicals called clotting factors (or coagulation factors) prompt reactions that activate still more coagulation factors. The process is complex, but is initiated along two basic pathways:

  • The extrinsic pathway, which normally is triggered by trauma.
  • The intrinsic pathway, which begins in the bloodstream and is triggered by internal damage to the wall of the vessel.

Both of these merge into a third pathway, referred to as the common pathway (see Figure 1b). All three pathways are dependent upon the 12 known clotting factors, including Ca2+ and vitamin K (Table 1). Clotting factors are secreted primarily by the liver and the platelets. The liver requires the fat-soluble vitamin K to produce many of them. Vitamin K (along with biotin and folate) is somewhat unusual among vitamins in that it is not only consumed in the diet but is also synthesized by bacteria residing in the large intestine. The calcium ion, considered factor IV, is derived from the diet and from the breakdown of bone. Some recent evidence indicates that activation of various clotting factors occurs on specific receptor sites on the surfaces of platelets.

The 12 clotting factors are numbered I through XIII according to the order of their discovery. Factor VI was once believed to be a distinct clotting factor, but is now thought to be identical to factor V. Rather than renumber the other factors, factor VI was allowed to remain as a placeholder and also a reminder that knowledge changes over time.

Extrinsic Pathway

The quicker responding and more direct extrinsic pathway (also known as the tissue factor pathway) begins when damage occurs to the surrounding tissues, such as in a traumatic injury. Upon contact with blood plasma, the damaged extravascular cells, which are extrinsic to the bloodstream, release factor III (thromboplastin). Sequentially, Ca2+ then factor VII (proconvertin), which is activated by factor III, are added, forming an enzyme complex. This enzyme complex leads to activation of factor X (Stuart–Prower factor), which activates the common pathway discussed below. The events in the extrinsic pathway are completed in a matter of seconds.

Intrinsic Pathway

The intrinsic pathway (also known as the contact activation pathway) is longer and more complex. In this case, the factors involved are intrinsic to (present within) the bloodstream. The pathway can be prompted by damage to the tissues, resulting from internal factors such as arterial disease; however, it is most often initiated when factor XII (Hageman factor) comes into contact with foreign materials, such as when a blood sample is put into a glass test tube. Within the body, factor XII is typically activated when it encounters negatively charged molecules, such as inorganic polymers and phosphate produced earlier in the series of intrinsic pathway reactions. Factor XII sets off a series of reactions that in turn activates factor XI (antihemolytic factor C or plasma thromboplastin antecedent) then factor IX (antihemolytic factor B or plasma thromboplasmin). In the meantime, chemicals released by the platelets increase the rate of these activation reactions. Finally, factor VIII (antihemolytic factor A) from the platelets and endothelial cells combines with factor IX (antihemolytic factor B or plasma thromboplasmin) to form an enzyme complex that activates factor X (Stuart–Prower factor or thrombokinase), leading to the common pathway. The events in the intrinsic pathway are completed in a few minutes.

Common Pathway

Both the intrinsic and extrinsic pathways lead to the common pathway, in which fibrin is produced to seal off the vessel. Once factor X has been activated by either the intrinsic or extrinsic pathway, the enzyme prothrombinase converts factor II, the inactive enzyme prothrombin, into the active enzyme thrombin. (Note that if the enzyme thrombin were not normally in an inactive form, clots would form spontaneously, a condition not consistent with life.) Then, thrombin converts factor I, the insoluble fibrinogen, into the soluble fibrin protein strands. Factor XIII then stabilizes the fibrin clot.
*Vitamin K required.
Clotting Factors (Table 1
Factor number Name Type of molecule Source Pathway(s)
I Fibrinogen Plasma protein Liver Common; converted into fibrin
II Prothrombin Plasma protein Liver* Common; converted into thrombin
III Tissue thromboplastin or tissue factor Lipoprotein mixture Damaged cells and platelets Extrinsic
IV Calcium ions Inorganic ions in plasma Diet, platelets, bone matrix Entire process
V Proaccelerin Plasma protein Liver, platelets Extrinsic and intrinsic
VI Not used Not used Not used Not used
VII Proconvertin Plasma protein Liver * Extrinsic
VIII Antihemolytic factor A Plasma protein factor Platelets and endothelial cells Intrinsic; deficiency results in hemophilia A
IX Antihemolytic factor B (plasma thromboplastin component) Plasma protein Liver* Intrinsic; deficiency results in hemophilia B
X Stuart–Prower factor (thrombokinase) Protein Liver* Extrinsic and intrinsic
XI Antihemolytic factor C (plasma thromboplastin antecedent) Plasma protein Liver Intrinsic; deficiency results in hemophilia C
XII Hageman factor Plasma protein Liver Intrinsic; initiates clotting in vitro also activates plasmin
XIII Fibrin-stabilizing factor Plasma protein Liver, platelets Stabilizes fibrin; slows fibrinolysis

Fibrinolysis

The stabilized clot is acted upon by contractile proteins within the platelets. As these proteins contract, they pull on the fibrin threads, bringing the edges of the clot more tightly together, somewhat as we do when tightening loose shoelaces (see Figure 1a). This process also wrings out of the clot a small amount of fluid called serum, which is blood plasma without its clotting factors.

To restore normal blood flow as the vessel heals, the clot must eventually be removed. Fibrinolysis is the gradual degradation of the clot. Again, there is a fairly complicated series of reactions that involves factor XII and protein-catabolizing enzymes. During this process, the inactive protein plasminogen is converted into the active plasmin, which gradually breaks down the fibrin of the clot. Additionally, bradykinin, a vasodilator, is released, reversing the effects of the serotonin and prostaglandins from the platelets. This allows the smooth muscle in the walls of the vessels to relax and helps to restore the circulation.

Plasma Anticoagulants

An anticoagulant is any substance that opposes coagulation. Several circulating plasma anticoagulants play a role in limiting the coagulation process to the region of injury and restoring a normal, clot-free condition of blood. For instance, a cluster of proteins collectively referred to as the protein C system inactivates clotting factors involved in the intrinsic pathway. TFPI (tissue factor pathway inhibitor) inhibits the conversion of the inactive factor VII to the active form in the extrinsic pathway. Antithrombin inactivates factor X and opposes the conversion of prothrombin (factor II) to thrombin in the common pathway. And as noted earlier, basophils release heparin, a short-acting anticoagulant that also opposes prothrombin. Heparin is also found on the surfaces of cells lining the blood vessels. A pharmaceutical form of heparin is often administered therapeutically, for example, in surgical patients at risk for blood clots.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL View these animations to explore the intrinsic, extrinsic, and common pathways that are involved the process of coagulation.[/caption]

Disorders of Clotting

Either an insufficient or an excessive production of platelets can lead to severe disease or death. As discussed earlier, an insufficient number of platelets, called thrombocytopenia, typically results in the inability of blood to form clots. This can lead to excessive bleeding, even from minor wounds.

Another reason for failure of the blood to clot is the inadequate production of functional amounts of one or more clotting factors. This is the case in the genetic disorder hemophilia, which is actually a group of related disorders, the most common of which is hemophilia A, accounting for approximately 80 percent of cases. This disorder results in the inability to synthesize sufficient quantities of factor VIII. Hemophilia B is the second most common form, accounting for approximately 20 percent of cases. In this case, there is a deficiency of factor IX. Both of these defects are linked to the X chromosome and are typically passed from a healthy (carrier) mother to her male offspring, since males are XY. Females would need to inherit a defective gene from each parent to manifest the disease, since they are XX. Patients with hemophilia bleed from even minor internal and external wounds, and leak blood into joint spaces after exercise and into urine and stool. Hemophilia C is a rare condition that is triggered by an autosomal (not sex) chromosome that renders factor XI nonfunctional. It is not a true recessive condition, since even individuals with a single copy of the mutant gene show a tendency to bleed. Regular infusions of clotting factors isolated from healthy donors can help prevent bleeding in hemophiliac patients. At some point, genetic therapy will become a viable option.

In contrast to the disorders characterized by coagulation failure is thrombocytosis, also mentioned earlier, a condition characterized by excessive numbers of platelets that increases the risk for excessive clot formation, a condition known as thrombosis. A thrombus (plural = thrombi) is an aggregation of platelets, erythrocytes, and even WBCs typically trapped within a mass of fibrin strands. While the formation of a clot is normal following the hemostatic mechanism just described, thrombi can form within an intact or only slightly damaged blood vessel. In a large vessel, a thrombus will adhere to the vessel wall and decrease the flow of blood, and is referred to as a mural thrombus. In a small vessel, it may actually totally block the flow of blood and is termed an occlusive thrombus. Thrombi are most commonly caused by vessel damage to the endothelial lining, which activates the clotting mechanism. These may include venous stasis, when blood in the veins, particularly in the legs, remains stationary for long periods. This is one of the dangers of long airplane flights in crowded conditions and may lead to deep vein thrombosis or atherosclerosis, an accumulation of debris in arteries. Thrombophilia, also called hypercoagulation, is a condition in which there is a tendency to form thrombosis. This may be familial (genetic) or acquired. Acquired forms include the autoimmune disease lupus, immune reactions to heparin, polycythemia vera, thrombocytosis, sickle cell disease, pregnancy, and even obesity. A thrombus can seriously impede blood flow to or from a region and will cause a local increase in blood pressure. If flow is to be maintained, the heart will need to generate a greater pressure to overcome the resistance.

When a portion of a thrombus breaks free from the vessel wall and enters the circulation, it is referred to as an embolus. An embolus that is carried through the bloodstream can be large enough to block a vessel critical to a major organ. When it becomes trapped, an embolus is called an embolism. In the heart, brain, or lungs, an embolism may accordingly cause a heart attack, a stroke, or a pulmonary embolism. These are medical emergencies.

Among the many known biochemical activities of aspirin is its role as an anticoagulant. Aspirin (acetylsalicylic acid) is very effective at inhibiting the aggregation of platelets. It is routinely administered during a heart attack or stroke to reduce the adverse effects. Physicians sometimes recommend that patients at risk for cardiovascular disease take a low dose of aspirin on a daily basis as a preventive measure. However, aspirin can also lead to serious side effects, including increasing the risk of ulcers. A patient is well advised to consult a physician before beginning any aspirin regimen.

A class of drugs collectively known as thrombolytic agents can help speed up the degradation of an abnormal clot. If a thrombolytic agent is administered to a patient within 3 hours following a thrombotic stroke, the patient’s prognosis improves significantly. However, some strokes are not caused by thrombi, but by hemorrhage. Thus, the cause must be determined before treatment begins. Tissue plasminogen activator is an enzyme that catalyzes the conversion of plasminogen to plasmin, the primary enzyme that breaks down clots. It is released naturally by endothelial cells but is also used in clinical medicine. New research is progressing using compounds isolated from the venom of some species of snakes, particularly vipers and cobras, which may eventually have therapeutic value as thrombolytic agents.

]]> 2217 2194 6 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2224 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2224 [caption id="" align="aligncenter" width="500"]This photo shows a human heart. Human Heart This artist’s conception of the human heart suggests a powerful engine—not inappropriate for a muscular pump that keeps the body continually supplied with blood. (credit: Patrick J. Lynch)[/caption]

Learning Objectives

After studying this chapter, you will be able to:
  • Identify and describe the interior and exterior parts of the human heart
  • Describe the path of blood through the cardiac circuits
  • Describe the size, shape, and location of the heart
  • Compare cardiac muscle to skeletal and smooth muscle
  • Explain the cardiac conduction system
  • Describe the process and purpose of an electrocardiogram
  • Explain the cardiac cycle
  • Calculate cardiac output
  • Describe the effects of exercise on cardiac output and heart rate
  • Name the centers of the brain that control heart rate and describe their function
  • Identify other factors affecting heart rate
  • Describe fetal heart development
In this chapter, you will explore the remarkable pump that propels the blood into the vessels. There is no single better word to describe the function of the heart other than “pump,” since its contraction develops the pressure that ejects blood into the major vessels: the aorta and pulmonary trunk. From these vessels, the blood is distributed to the remainder of the body. Although the connotation of the term “pump” suggests a mechanical device made of steel and plastic, the anatomical structure is a living, sophisticated muscle. As you read this chapter, try to keep these twin concepts in mind: pump and muscle. Although the term “heart” is an English word, cardiac (heart-related) terminology can be traced back to the Latin term, “kardia.” Cardiology is the study of the heart, and cardiologists are the physicians who deal primarily with the heart.]]> 2224 2222 1 0 19.2 Cardiac Muscle and Electrical Activity https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2249 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2249

Learning Objectives

By the end of this section, you will be able to:
  • Describe the structure of cardiac muscle
  • Identify and describe the components of the conducting system that distributes electrical impulses through the heart
  • Compare the effect of ion movement on membrane potential of cardiac conductive and contractile cells
  • Relate characteristics of an electrocardiogram to events in the cardiac cycle
  • Identify blocks that can interrupt the cardiac cycle

Recall that cardiac muscle shares a few characteristics with both skeletal muscle and smooth muscle, but it has some unique properties of its own. Not the least of these exceptional properties is its ability to initiate an electrical potential at a fixed rate that spreads rapidly from cell to cell to trigger the contractile mechanism. This property is known as autorhythmicity. Neither smooth nor skeletal muscle can do this. Even though cardiac muscle has autorhythmicity, heart rate is modulated by the endocrine and nervous systems.

There are two major types of cardiac muscle cells: myocardial contractile cells and myocardial conducting cells. The myocardial contractile cells constitute the bulk (99 percent) of the cells in the atria and ventricles. Contractile cells conduct impulses and are responsible for contractions that pump blood through the body. The myocardial conducting cells (1 percent of the cells) form the conduction system of the heart. Except for Purkinje cells, they are generally much smaller than the contractile cells and have few of the myofibrils or filaments needed for contraction. Their function is similar in many respects to neurons, although they are specialized muscle cells. Myocardial conduction cells initiate and propagate the action potential (the electrical impulse) that travels throughout the heart and triggers the contractions that propel the blood.

Structure of Cardiac Muscle

Compared to the giant cylinders of skeletal muscle, cardiac muscle cells, or cardiomyocytes, are considerably shorter with much smaller diameters. Cardiac muscle also demonstrates striations, the alternating pattern of dark A bands and light I bands attributed to the precise arrangement of the myofilaments and fibrils that are organized in sarcomeres along the length of the cell (Figure 1a). These contractile elements are virtually identical to skeletal muscle. T (transverse) tubules penetrate from the surface plasma membrane, the sarcolemma, to the interior of the cell, allowing the electrical impulse to reach the interior. The T tubules are only found at the Z discs, whereas in skeletal muscle, they are found at the junction of the A and I bands. Therefore, there are one-half as many T tubules in cardiac muscle as in skeletal muscle. In addition, the sarcoplasmic reticulum stores few calcium ions, so most of the calcium ions must come from outside the cells. The result is a slower onset of contraction. Mitochondria are plentiful, providing energy for the contractions of the heart. Typically, cardiomyocytes have a single, central nucleus, but two or more nuclei may be found in some cells.

Cardiac muscle cells branch freely. A junction between two adjoining cells is marked by a critical structure called an intercalated disc, which helps support the synchronized contraction of the muscle (Figure 1b). The sarcolemmas from adjacent cells bind together at the intercalated discs. They consist of desmosomes, specialized linking proteoglycans, tight junctions, and large numbers of gap junctions that allow the passage of ions between the cells and help to synchronize the contraction (Figure 1c). Intercellular connective tissue also helps to bind the cells together. The importance of strongly binding these cells together is necessitated by the forces exerted by contraction.

[caption id="" align="aligncenter" width="520"]The top left panel of this figure shows the cross structure of cardiac muscle with the major parts labeled. The top right panel shows a micrograph of cardiac muscle. The bottom panel shows the structure of intercalated discs. Figure 1. Cardiac Muscle. (a) Cardiac muscle cells have myofibrils composed of myofilaments arranged in sarcomeres, T tubules to transmit the impulse from the sarcolemma to the interior of the cell, numerous mitochondria for energy, and intercalated discs that are found at the junction of different cardiac muscle cells. (b) A photomicrograph of cardiac muscle cells shows the nuclei and intercalated discs. (c) An intercalated disc connects cardiac muscle cells and consists of desmosomes and gap junctions. LM × 1600. (Micrograph provided by the Regents of the University of Michigan Medical School © 2012)[/caption]

Cardiac muscle undergoes aerobic respiration patterns, primarily metabolizing lipids and carbohydrates. Myoglobin, lipids, and glycogen are all stored within the cytoplasm. Cardiac muscle cells undergo twitch-type contractions with long refractory periods followed by brief relaxation periods. The relaxation is essential so the heart can fill with blood for the next cycle. The refractory period is very long to prevent the possibility of tetany, a condition in which muscle remains involuntarily contracted. In the heart, tetany is not compatible with life, since it would prevent the heart from pumping blood.

Repair and Replacement

Damaged cardiac muscle cells have extremely limited abilities to repair themselves or to replace dead cells via mitosis. Recent evidence indicates that at least some stem cells remain within the heart that continue to divide and at least potentially replace these dead cells. However, newly formed or repaired cells are rarely as functional as the original cells, and cardiac function is reduced. In the event of a heart attack or MI, dead cells are often replaced by patches of scar tissue. Autopsies performed on individuals who had successfully received heart transplants show some proliferation of original cells. If researchers can unlock the mechanism that generates new cells and restore full mitotic capabilities to heart muscle, the prognosis for heart attack survivors will be greatly enhanced. To date, myocardial cells produced within the patient (in situ) by cardiac stem cells seem to be nonfunctional, although those grown in Petri dishes (in vitro) do beat. Perhaps soon this mystery will be solved, and new advances in treatment will be commonplace.

Conduction System of the Heart

If embryonic heart cells are separated into a Petri dish and kept alive, each is capable of generating its own electrical impulse followed by contraction. When two independently beating embryonic cardiac muscle cells are placed together, the cell with the higher inherent rate sets the pace, and the impulse spreads from the faster to the slower cell to trigger a contraction. As more cells are joined together, the fastest cell continues to assume control of the rate. A fully developed adult heart maintains the capability of generating its own electrical impulse, triggered by the fastest cells, as part of the cardiac conduction system. The components of the cardiac conduction system include the sinoatrial node, the atrioventricular node, the atrioventricular bundle, the atrioventricular bundle branches, and the Purkinje cells (Figure 2).

[caption id="" align="aligncenter" width="500"]This image shows the anterior view of the frontal section of the heart with the major parts labeled. Figure 2. Conduction System of the Heart. Specialized conducting components of the heart include the sinoatrial node, the internodal pathways, the atrioventricular node, the atrioventricular bundle, the right and left bundle branches, and the Purkinje fibers.[/caption]

Sinoatrial (SA) Node

Normal cardiac rhythm is established by the sinoatrial (SA) node, a specialized clump of myocardial conducting cells located in the superior and posterior walls of the right atrium in close proximity to the orifice of the superior vena cava. The SA node has the highest inherent rate of depolarization and is known as the pacemaker of the heart. It initiates the sinus rhythm, or normal electrical pattern followed by contraction of the heart.

This impulse spreads from its initiation in the SA node throughout the atria through specialized internodal pathways, to the atrial myocardial contractile cells and the atrioventricular node. The internodal pathways consist of three bands (anterior, middle, and posterior) that lead directly from the SA node to the next node in the conduction system, the atrioventricular node (see Figure 2). The impulse takes approximately 50 ms (milliseconds) to travel between these two nodes. The relative importance of this pathway has been debated since the impulse would reach the atrioventricular node simply following the cell-by-cell pathway through the contractile cells of the myocardium in the atria. In addition, there is a specialized pathway called Bachmann’s bundle or the interatrial band that conducts the impulse directly from the right atrium to the left atrium. Regardless of the pathway, as the impulse reaches the atrioventricular septum, the connective tissue of the cardiac skeleton prevents the impulse from spreading into the myocardial cells in the ventricles except at the atrioventricular node. Figure 3 illustrates the initiation of the impulse in the SA node that then spreads the impulse throughout the atria to the atrioventricular node.

[caption id="" align="aligncenter" width="400"]This image shows the different stages in the conduction cycle of the heart. Figure 3. Cardiac Conduction. (1) The sinoatrial (SA) node and the remainder of the conduction system are at rest. (2) The SA node initiates the action potential, which sweeps across the atria. (3) After reaching the atrioventricular node, there is a delay of approximately 100 ms that allows the atria to complete pumping blood before the impulse is transmitted to the atrioventricular bundle. (4) Following the delay, the impulse travels through the atrioventricular bundle and bundle branches to the Purkinje fibers, and also reaches the right papillary muscle via the moderator band. (5) The impulse spreads to the contractile fibers of the ventricle. (6) Ventricular contraction begins.[/caption]

The electrical event, the wave of depolarization, is the trigger for muscular contraction. The wave of depolarization begins in the right atrium, and the impulse spreads across the superior portions of both atria and then down through the contractile cells. The contractile cells then begin contraction from the superior to the inferior portions of the atria, efficiently pumping blood into the ventricles.

Atrioventricular (AV) Node

The atrioventricular (AV) node is a second clump of specialized myocardial conductive cells, located in the inferior portion of the right atrium within the atrioventricular septum. The septum prevents the impulse from spreading directly to the ventricles without passing through the AV node. There is a critical pause before the AV node depolarizes and transmits the impulse to the atrioventricular bundle (see Figure 3, step 3). This delay in transmission is partially attributable to the small diameter of the cells of the node, which slow the impulse. Also, conduction between nodal cells is less efficient than between conducting cells. These factors mean that it takes the impulse approximately 100 ms to pass through the node. This pause is critical to heart function, as it allows the atrial cardiomyocytes to complete their contraction that pumps blood into the ventricles before the impulse is transmitted to the cells of the ventricle itself. With extreme stimulation by the SA node, the AV node can transmit impulses maximally at 220 per minute. This establishes the typical maximum heart rate in a healthy young individual. Damaged hearts or those stimulated by drugs can contract at higher rates, but at these rates, the heart can no longer effectively pump blood.

Atrioventricular Bundle (Bundle of His), Bundle Branches, and Purkinje Fibers

Arising from the AV node, the atrioventricular bundle, or bundle of His, proceeds through the interventricular septum before dividing into two atrioventricular bundle branches, commonly called the left and right bundle branches. The left bundle branch has two fascicles. The left bundle branch supplies the left ventricle, and the right bundle branch the right ventricle. Since the left ventricle is much larger than the right, the left bundle branch is also considerably larger than the right. Portions of the right bundle branch are found in the moderator band and supply the right papillary muscles. Because of this connection, each papillary muscle receives the impulse at approximately the same time, so they begin to contract simultaneously just prior to the remainder of the myocardial contractile cells of the ventricles. This is believed to allow tension to develop on the chordae tendineae prior to right ventricular contraction. There is no corresponding moderator band on the left. Both bundle branches descend and reach the apex of the heart where they connect with the Purkinje fibers (see Figure 3, step 4). This passage takes approximately 25 ms.

The Purkinje fibers are additional myocardial conductive fibers that spread the impulse to the myocardial contractile cells in the ventricles. They extend throughout the myocardium from the apex of the heart toward the atrioventricular septum and the base of the heart. The Purkinje fibers have a fast inherent conduction rate, and the electrical impulse reaches all of the ventricular muscle cells in about 75 ms (see Figure 3, step 5). Since the electrical stimulus begins at the apex, the contraction also begins at the apex and travels toward the base of the heart, similar to squeezing a tube of toothpaste from the bottom. This allows the blood to be pumped out of the ventricles and into the aorta and pulmonary trunk. The total time elapsed from the initiation of the impulse in the SA node until depolarization of the ventricles is approximately 225 ms.

Membrane Potentials and Ion Movement in Cardiac Conductive Cells

Action potentials are considerably different between cardiac conductive cells and cardiac contractive cells. While Na+ and K+ play essential roles, Ca2+ is also critical for both types of cells. Unlike skeletal muscles and neurons, cardiac conductive cells do not have a stable resting potential. Conductive cells contain a series of sodium ion channels that allow a normal and slow influx of sodium ions that causes the membrane potential to rise slowly from an initial value of −60 mV up to about –40 mV. The resulting movement of sodium ions creates spontaneous depolarization (or prepotential depolarization). At this point, calcium ion channels open and Ca2+ enters the cell, further depolarizing it at a more rapid rate until it reaches a value of approximately +5 mV. At this point, the calcium ion channels close and K+ channels open, allowing outflux of K+ and resulting in repolarization. When the membrane potential reaches approximately −60 mV, the K+ channels close and Na+ channels open, and the prepotential phase begins again. This phenomenon explains the autorhythmicity properties of cardiac muscle (Figure 4).

[caption id="" align="aligncenter" width="430"]This graph shows the change in membrane potential as a function of time. Figure 4. Action Potential at the SA Node. The prepotential is due to a slow influx of sodium ions until the threshold is reached followed by a rapid depolarization and repolarization. The prepotential accounts for the membrane reaching threshold and initiates the spontaneous depolarization and contraction of the cell. Note the lack of a resting potential.[/caption]

Membrane Potentials and Ion Movement in Cardiac Contractile Cells

There is a distinctly different electrical pattern involving the contractile cells. In this case, there is a rapid depolarization, followed by a plateau phase and then repolarization. This phenomenon accounts for the long refractory periods required for the cardiac muscle cells to pump blood effectively before they are capable of firing for a second time. These cardiac myocytes normally do not initiate their own electrical potential, although they are capable of doing so, but rather wait for an impulse to reach them.

Contractile cells demonstrate a much more stable resting phase than conductive cells at approximately −80 mV for cells in the atria and −90 mV for cells in the ventricles. Despite this initial difference, the other components of their action potentials are virtually identical. In both cases, when stimulated by an action potential, voltage-gated channels rapidly open, beginning the positive-feedback mechanism of depolarization. This rapid influx of positively charged ions raises the membrane potential to approximately +30 mV, at which point the sodium channels close. The rapid depolarization period typically lasts 3–5 ms. Depolarization is followed by the plateau phase, in which membrane potential declines relatively slowly. This is due in large part to the opening of the slow Ca2+ channels, allowing Ca2+ to enter the cell while few K+ channels are open, allowing K+ to exit the cell. The relatively long plateau phase lasts approximately 175 ms. Once the membrane potential reaches approximately zero, the Ca2+ channels close and K+ channels open, allowing K+ to exit the cell. The repolarization lasts approximately 75 ms. At this point, membrane potential drops until it reaches resting levels once more and the cycle repeats. The entire event lasts between 250 and 300 ms (Figure 5).

The absolute refractory period for cardiac contractile muscle lasts approximately 200 ms, and the relative refractory period lasts approximately 50 ms, for a total of 250 ms. This extended period is critical, since the heart muscle must contract to pump blood effectively and the contraction must follow the electrical events. Without extended refractory periods, premature contractions would occur in the heart and would not be compatible with life.

[caption id="" align="aligncenter" width="480"]The top panel of this figure shows millivolts as a function of time with the various stages labeled. The bottom left panel shows action potential and tension as a function of time for skeletal muscle, and the bottom right panel shows the action potential and tension as a function of time for cardiac muscle. Figure 5. Action Potential in Cardiac Contractile Cells. (a) Note the long plateau phase due to the influx of calcium ions. The extended refractory period allows the cell to fully contract before another electrical event can occur. (b) The action potential for heart muscle is compared to that of skeletal muscle.[/caption]

Calcium Ions

Calcium ions play two critical roles in the physiology of cardiac muscle. Their influx through slow calcium channels accounts for the prolonged plateau phase and absolute refractory period that enable cardiac muscle to function properly. Calcium ions also combine with the regulatory protein troponin in the troponin-tropomyosin complex; this complex removes the inhibition that prevents the heads of the myosin molecules from forming cross bridges with the active sites on actin that provide the power stroke of contraction. This mechanism is virtually identical to that of skeletal muscle. Approximately 20 percent of the calcium required for contraction is supplied by the influx of Ca2+ during the plateau phase. The remaining Ca2+ for contraction is released from storage in the sarcoplasmic reticulum.

Comparative Rates of Conduction System Firing

The pattern of prepotential or spontaneous depolarization, followed by rapid depolarization and repolarization just described, are seen in the SA node and a few other conductive cells in the heart. Since the SA node is the pacemaker, it reaches threshold faster than any other component of the conduction system. It will initiate the impulses spreading to the other conducting cells. The SA node, without nervous or endocrine control, would initiate a heart impulse approximately 80–100 times per minute. Although each component of the conduction system is capable of generating its own impulse, the rate progressively slows as you proceed from the SA node to the Purkinje fibers. Without the SA node, the AV node would generate a heart rate of 40–60 beats per minute. If the AV node were blocked, the atrioventricular bundle would fire at a rate of approximately 30–40 impulses per minute. The bundle branches would have an inherent rate of 20–30 impulses per minute, and the Purkinje fibers would fire at 15–20 impulses per minute. While a few exceptionally trained aerobic athletes demonstrate resting heart rates in the range of 30–40 beats per minute (the lowest recorded figure is 28 beats per minute for Miguel Indurain, a cyclist), for most individuals, rates lower than 50 beats per minute would indicate a condition called bradycardia. Depending upon the specific individual, as rates fall much below this level, the heart would be unable to maintain adequate flow of blood to vital tissues, initially resulting in decreasing loss of function across the systems, unconsciousness, and ultimately death.

Electrocardiogram

By careful placement of surface electrodes on the body, it is possible to record the complex, compound electrical signal of the heart. This tracing of the electrical signal is the electrocardiogram (ECG), also commonly abbreviated EKG (K coming kardiology, from the German term for cardiology). Careful analysis of the ECG reveals a detailed picture of both normal and abnormal heart function, and is an indispensable clinical diagnostic tool. The standard electrocardiograph (the instrument that generates an ECG) uses 3, 5, or 12 leads. The greater the number of leads an electrocardiograph uses, the more information the ECG provides. The term “lead” may be used to refer to the cable from the electrode to the electrical recorder, but it typically describes the voltage difference between two of the electrodes. The 12-lead electrocardiograph uses 10 electrodes placed in standard locations on the patient’s skin (Figure 6). In continuous ambulatory electrocardiographs, the patient wears a small, portable, battery-operated device known as a Holter monitor, or simply a Holter, that continuously monitors heart electrical activity, typically for a period of 24 hours during the patient’s normal routine.
[caption id="" align="aligncenter" width="350"]This diagram shows the points where electrodes are placed on the body for an ECG. Figure 6. Standard Placement of ECG Leads. In a 12-lead ECG, six electrodes are placed on the chest, and four electrodes are placed on the limbs.[/caption]

A normal ECG tracing is presented in Figure 7. Each component, segment, and interval is labeled and corresponds to important electrical events, demonstrating the relationship between these events and contraction in the heart.

There are five prominent points on the ECG: the P wave, the QRS complex, and the T wave. The small P wave represents the depolarization of the atria. The atria begin contracting approximately 25 ms after the start of the P wave. The large QRS complex represents the depolarization of the ventricles, which requires a much stronger electrical signal because of the larger size of the ventricular cardiac muscle. The ventricles begin to contract as the QRS reaches the peak of the R wave. Lastly, the T wave represents the repolarization of the ventricles. The repolarization of the atria occurs during the QRS complex, which masks it on an ECG.

The major segments and intervals of an ECG tracing are indicated in Figure 7. Segments are defined as the regions between two waves. Intervals include one segment plus one or more waves. For example, the PR segment begins at the end of the P wave and ends at the beginning of the QRS complex. The PR interval starts at the beginning of the P wave and ends with the beginning of the QRS complex. The PR interval is more clinically relevant, as it measures the duration from the beginning of atrial depolarization (the P wave) to the initiation of the QRS complex. Since the Q wave may be difficult to view in some tracings, the measurement is often extended to the R that is more easily visible. Should there be a delay in passage of the impulse from the SA node to the AV node, it would be visible in the PR interval. Figure 8 correlates events of heart contraction to the corresponding segments and intervals of an ECG.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit this site for a more detailed analysis of ECGs.[/caption]
[caption id="" align="aligncenter" width="430"]This figure shows a graph of millivolts over time and the heart cycles during an ECG. Figure 7. Electrocardiogram. A normal tracing shows the P wave, QRS complex, and T wave. Also indicated are the PR, QT, QRS, and ST intervals, plus the P-R and S-T segments.[/caption]
[caption id="" align="aligncenter" width="480"]This diagram shows the different stages of heart contraction and relaxation along with the stages in the QT cycle. Figure 8. ECG Tracing Correlated to the Cardiac Cycle. This diagram correlates an ECG tracing with the electrical and mechanical events of a heart contraction. Each segment of an ECG tracing corresponds to one event in the cardiac cycle.[/caption]
Occassionally, an area of the heart other than the SA node will initiate an impulse that will be followed by a premature contraction. Such an area, which may actually be a component of the conduction system or some other contractile cells, is known as an ectopic focus or ectopic pacemaker. An ectopic focus may be stimulated by localized ischemia; exposure to certain drugs, including caffeine, digitalis, or acetylcholine; elevated stimulation by both sympathetic or parasympathetic divisions of the autonomic nervous system; or a number of disease or pathological conditions. Occasional occurances are generally transitory and nonlife threatening, but if the condition becomes chronic, it may lead to either an arrhythmia, a deviation from the normal pattern of impulse conduction and contraction, or to fibrillation, an uncoordinated beating of the heart.

While interpretation of an ECG is possible and extremely valuable after some training, a full understanding of the complexities and intricacies generally requires several years of experience. In general, the size of the electrical variations, the duration of the events, and detailed vector analysis provide the most comprehensive picture of cardiac function. For example, an amplified P wave may indicate enlargement of the atria, an enlarged Q wave may indicate a MI, and an enlarged suppressed or inverted Q wave often indicates enlarged ventricles. T waves often appear flatter when insufficient oxygen is being delivered to the myocardium. An elevation of the ST segment above baseline is often seen in patients with an acute MI, and may appear depressed below the baseline when hypoxia is occurring.

As useful as analyzing these electrical recordings may be, there are limitations. For example, not all areas suffering a MI may be obvious on the ECG. Additionally, it will not reveal the effectiveness of the pumping, which requires further testing, such as an ultrasound test called an echocardiogram or nuclear medicine imaging. It is also possible for there to be pulseless electrical activity, which will show up on an ECG tracing, although there is no corresponding pumping action. Common abnormalities that may be detected by the ECGs are shown in Figure 9.

[caption id="" align="aligncenter" width="520"]In this image the QT cycle for different heart conditions are shown. From top to bottom, the arrhythmias shown are second-degree partial block, atrial fibrillation, ventricular tachycardia, ventricular fibrillation and third degree block. Figure 9. Common ECG Abnormalities. (a) In a second-degree or partial block, one-half of the P waves are not followed by the QRS complex and T waves while the other half are. (b) In atrial fibrillation, the electrical pattern is abnormal prior to the QRS complex, and the frequency between the QRS complexes has increased. (c) In ventricular tachycardia, the shape of the QRS complex is abnormal. (d) In ventricular fibrillation, there is no normal electrical activity. (e) In a third-degree block, there is no correlation between atrial activity (the P wave) and ventricular activity (the QRS complex).[/caption]
[caption id="attachment_1737" align="aligncenter" width="150"]QR Code representing a URL Visit this site for a more complete library of abnormal ECGs.[/caption]
]]> 2249 2222 3 0 19.3 Cardiac Cycle https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2254 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2254

Learning Objectives

By the end of this section, you will be able to:
  • Describe the relationship between blood pressure and blood flow
  • Summarize the events of the cardiac cycle
  • Compare atrial and ventricular systole and diastole
  • Relate heart sounds detected by auscultation to action of heart’s valves
The period of time that begins with contraction of the atria and ends with ventricular relaxation is known as the cardiac cycle (Figure 1). The period of contraction that the heart undergoes while it pumps blood into circulation is called systole. The period of relaxation that occurs as the chambers fill with blood is called diastole. Both the atria and ventricles undergo systole and diastole, and it is essential that these components be carefully regulated and coordinated to ensure blood is pumped efficiently to the body.
[caption id="" align="aligncenter" width="420"]This pie chart shows the different phases of the cardiac cycle and details the atrial and ventricular stages. Figure 1. Overview of the Cardiac Cycle. The cardiac cycle begins with atrial systole and progresses to ventricular systole, atrial diastole, and ventricular diastole, when the cycle begins again. Correlations to the ECG are highlighted.[/caption]

Pressures and Flow

Fluids, whether gases or liquids, are materials that flow according to pressure gradients—that is, they move from regions that are higher in pressure to regions that are lower in pressure. Accordingly, when the heart chambers are relaxed (diastole), blood will flow into the atria from the veins, which are higher in pressure. As blood flows into the atria, the pressure will rise, so the blood will initially move passively from the atria into the ventricles. When the action potential triggers the muscles in the atria to contract (atrial systole), the pressure within the atria rises further, pumping blood into the ventricles. During ventricular systole, pressure rises in the ventricles, pumping blood into the pulmonary trunk from the right ventricle and into the aorta from the left ventricle. Again, as you consider this flow and relate it to the conduction pathway, the elegance of the system should become apparent.

Phases of the Cardiac Cycle

At the beginning of the cardiac cycle, both the atria and ventricles are relaxed (diastole). Blood is flowing into the right atrium from the superior and inferior venae cavae and the coronary sinus. Blood flows into the left atrium from the four pulmonary veins. The two atrioventricular valves, the tricuspid and mitral valves, are both open, so blood flows unimpeded from the atria and into the ventricles. Approximately 70–80 percent of ventricular filling occurs by this method. The two semilunar valves, the pulmonary and aortic valves, are closed, preventing backflow of blood into the right and left ventricles from the pulmonary trunk on the right and the aorta on the left.

Atrial Systole and Diastole

Contraction of the atria follows depolarization, represented by the P wave of the ECG. As the atrial muscles contract from the superior portion of the atria toward the atrioventricular septum, pressure rises within the atria and blood is pumped into the ventricles through the open atrioventricular (tricuspid, and mitral or bicuspid) valves. At the start of atrial systole, the ventricles are normally filled with approximately 70–80 percent of their capacity due to inflow during diastole. Atrial contraction, also referred to as the “atrial kick,” contributes the remaining 20–30 percent of filling (see Figure 1). Atrial systole lasts approximately 100 ms and ends prior to ventricular systole, as the atrial muscle returns to diastole.

Ventricular Systole

Ventricular systole (see Figure 1) follows the depolarization of the ventricles and is represented by the QRS complex in the ECG. It may be conveniently divided into two phases, lasting a total of 270 ms. At the end of atrial systole and just prior to atrial contraction, the ventricles contain approximately 130 mL blood in a resting adult in a standing position. This volume is known as the end diastolic volume (EDV) or preload.

Initially, as the muscles in the ventricle contract, the pressure of the blood within the chamber rises, but it is not yet high enough to open the semilunar (pulmonary and aortic) valves and be ejected from the heart. However, blood pressure quickly rises above that of the atria that are now relaxed and in diastole. This increase in pressure causes blood to flow back toward the atria, closing the tricuspid and mitral valves. Since blood is not being ejected from the ventricles at this early stage, the volume of blood within the chamber remains constant. Consequently, this initial phase of ventricular systole is known as isovolumic contraction, also called isovolumetric contraction (see Figure 1).

In the second phase of ventricular systole, the ventricular ejection phase, the contraction of the ventricular muscle has raised the pressure within the ventricle to the point that it is greater than the pressures in the pulmonary trunk and the aorta. Blood is pumped from the heart, pushing open the pulmonary and aortic semilunar valves. Pressure generated by the left ventricle will be appreciably greater than the pressure generated by the right ventricle, since the existing pressure in the aorta will be so much higher. Nevertheless, both ventricles pump the same amount of blood. This quantity is referred to as stroke volume. Stroke volume will normally be in the range of 70–80 mL. Since ventricular systole began with an EDV of approximately 130 mL of blood, this means that there is still 50–60 mL of blood remaining in the ventricle following contraction. This volume of blood is known as the end systolic volume (ESV).

Ventricular Diastole

Ventricular relaxation, or diastole, follows repolarization of the ventricles and is represented by the T wave of the ECG. It too is divided into two distinct phases and lasts approximately 430 ms.

During the early phase of ventricular diastole, as the ventricular muscle relaxes, pressure on the remaining blood within the ventricle begins to fall. When pressure within the ventricles drops below pressure in both the pulmonary trunk and aorta, blood flows back toward the heart, producing the dicrotic notch (small dip) seen in blood pressure tracings. The semilunar valves close to prevent backflow into the heart. Since the atrioventricular valves remain closed at this point, there is no change in the volume of blood in the ventricle, so the early phase of ventricular diastole is called the isovolumic ventricular relaxation phase, also called isovolumetric ventricular relaxation phase (see Figure 1).

In the second phase of ventricular diastole, called late ventricular diastole, as the ventricular muscle relaxes, pressure on the blood within the ventricles drops even further. Eventually, it drops below the pressure in the atria. When this occurs, blood flows from the atria into the ventricles, pushing open the tricuspid and mitral valves. As pressure drops within the ventricles, blood flows from the major veins into the relaxed atria and from there into the ventricles. Both chambers are in diastole, the atrioventricular valves are open, and the semilunar valves remain closed (see Figure 1). The cardiac cycle is complete.

Figure 2 illustrates the relationship between the cardiac cycle and the ECG.

[caption id="" align="aligncenter" width="480"]This image shows the correlation between the cardiac cycle and the different stages in a electrocardiogram. Figure 2. Relationship between the Cardiac Cycle and ECG. Initially, both the atria and ventricles are relaxed (diastole). The P wave represents depolarization of the atria and is followed by atrial contraction (systole). Atrial systole extends until the QRS complex, at which point, the atria relax. The QRS complex represents depolarization of the ventricles and is followed by ventricular contraction. The T wave represents the repolarization of the ventricles and marks the beginning of ventricular relaxation.[/caption]

Heart Sounds

One of the simplest, yet effective, diagnostic techniques applied to assess the state of a patient’s heart is auscultation using a stethoscope.

In a normal, healthy heart, there are only two audible heart sounds: S1 and S2. S1 is the sound created by the closing of the atrioventricular valves during ventricular contraction and is normally described as a “lub,” or first heart sound. The second heart sound, S2, is the sound of the closing of the semilunar valves during ventricular diastole and is described as a “dub” (Figure 3). In both cases, as the valves close, the openings within the atrioventricular septum guarded by the valves will become reduced, and blood flow through the opening will become more turbulent until the valves are fully closed. There is a third heart sound, S3, but it is rarely heard in healthy individuals. It may be the sound of blood flowing into the atria, or blood sloshing back and forth in the ventricle, or even tensing of the chordae tendineae. S3 may be heard in youth, some athletes, and pregnant women. If the sound is heard later in life, it may indicate congestive heart failure, warranting further tests. Some cardiologists refer to the collective S1, S2, and S3 sounds as the “Kentucky gallop,” because they mimic those produced by a galloping horse. The fourth heart sound, S4, results from the contraction of the atria pushing blood into a stiff or hypertrophic ventricle, indicating failure of the left ventricle. S4 occurs prior to S1 and the collective sounds S4, S1, and S2 are referred to by some cardiologists as the “Tennessee gallop,” because of their similarity to the sound produced by a galloping horse with a different gait. A few individuals may have both S3 and S4, and this combined sound is referred to as S7.

[caption id="" align="aligncenter" width="480"]This image shows a graph of the blood pressure with the different stages labeled. Under the graph, a line shows the different sounds made by the beating heart. Figure 3. Heart Sounds and the Cardiac Cycle. In this illustration, the x-axis reflects time with a recording of the heart sounds. The y-axis represents pressure.[/caption]

The term murmur is used to describe an unusual sound coming from the heart that is caused by the turbulent flow of blood. Murmurs are graded on a scale of 1 to 6, with 1 being the most common, the most difficult sound to detect, and the least serious. The most severe is a 6. Phonocardiograms or auscultograms can be used to record both normal and abnormal sounds using specialized electronic stethoscopes.

During auscultation, it is common practice for the clinician to ask the patient to breathe deeply. This procedure not only allows for listening to airflow, but it may also amplify heart murmurs. Inhalation increases blood flow into the right side of the heart and may increase the amplitude of right-sided heart murmurs. Expiration partially restricts blood flow into the left side of the heart and may amplify left-sided heart murmurs. Figure 4 indicates proper placement of the bell of the stethoscope to facilitate auscultation.
[caption id="" align="aligncenter" width="480"]This image shows the points on the human chest where the stethoscope can be placed to hear the heart beat. Figure 4. Stethoscope Placement for Auscultation. Proper placement of the bell of the stethoscope facilitates auscultation. At each of the four locations on the chest, a different valve can be heard.[/caption]
]]> 2254 2222 4 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2263 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2263 [caption id="" align="aligncenter" width="500"]This photo shows a forearm with the veins bulging. Figure 1. Blood Vessels. While most blood vessels are located deep from the surface and are not visible, the superficial veins of the upper limb provide an indication of the extent, prominence, and importance of these structures to the body. (credit: Colin Davis)[/caption]

Chapter Objectives

After studying this chapter, you will be able to:
  • Compare and contrast the anatomical structure of arteries, arterioles, capillaries, venules, and veins
  • Accurately describe the forces that account for capillary exchange
  • List the major factors affecting blood flow, blood pressure, and resistance
  • Describe how blood flow, blood pressure, and resistance interrelate
  • Discuss how the neural and endocrine mechanisms maintain homeostasis within the blood vessels
  • Describe the interaction of the cardiovascular system with other body systems
  • Label the major blood vessels of the pulmonary and systemic circulations
  • Identify and describe the hepatic portal system
  • Describe the development of blood vessels and fetal circulation
  • Compare fetal circulation to that of an individual after birth
In this chapter, you will learn about the vascular part of the cardiovascular system, that is, the vessels that transport blood throughout the body and provide the physical site where gases, nutrients, and other substances are exchanged with body cells. When vessel functioning is reduced, blood-borne substances do not circulate effectively throughout the body. As a result, tissue injury occurs, metabolism is impaired, and the functions of every bodily system are threatened.]]> 2263 2261 1 0 20.1 Structure and Function of Blood Vessels https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2271 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2271

Learning Objectives

By the end of this section, you will be able to:
  • Compare and contrast the three tunics that make up the walls of most blood vessels
  • Distinguish between elastic arteries, muscular arteries, and arterioles on the basis of structure, location, and function
  • Describe the basic structure of a capillary bed, from the supplying metarteriole to the venule into which it drains
  • Explain the structure and function of venous valves in the large veins of the extremities

Blood is carried through the body via blood vessels. An artery is a blood vessel that carries blood away from the heart, where it branches into ever-smaller vessels. Eventually, the smallest arteries, vessels called arterioles, further branch into tiny capillaries, where nutrients and wastes are exchanged, and then combine with other vessels that exit capillaries to form venules, small blood vessels that carry blood to a vein, a larger blood vessel that returns blood to the heart.

Arteries and veins transport blood in two distinct circuits: the systemic circuit and the pulmonary circuit (Figure 1). Systemic arteries provide blood rich in oxygen to the body’s tissues. The blood returned to the heart through systemic veins has less oxygen, since much of the oxygen carried by the arteries has been delivered to the cells. In contrast, in the pulmonary circuit, arteries carry blood low in oxygen exclusively to the lungs for gas exchange. Pulmonary veins then return freshly oxygenated blood from the lungs to the heart to be pumped back out into systemic circulation. Although arteries and veins differ structurally and functionally, they share certain features.

[caption id="" align="aligncenter" width="520"]This diagram shows how oxygenated and deoxygenated blood flow through the major organs in the body. Figure 1. Cardiovascular Circulation. The pulmonary circuit moves blood from the right side of the heart to the lungs and back to the heart. The systemic circuit moves blood from the left side of the heart to the head and body and returns it to the right side of the heart to repeat the cycle. The arrows indicate the direction of blood flow, and the colors show the relative levels of oxygen concentration.[/caption]

Shared Structures

Different types of blood vessels vary slightly in their structures, but they share the same general features. Arteries and arterioles have thicker walls than veins and venules because they are closer to the heart and receive blood that is surging at a far greater pressure (Figure 2). Each type of vessel has a lumen—a hollow passageway through which blood flows. Arteries have smaller lumens than veins, a characteristic that helps to maintain the pressure of blood moving through the system. Together, their thicker walls and smaller diameters give arterial lumens a more rounded appearance in cross section than the lumens of veins.

[caption id="" align="aligncenter" width="425"]The top left panel of this figure shows the ultrastructure of an artery, and the top right panel shows the ultrastructure of a vein. The bottom panel shows a micrograph with the cross sections of an artery and a vein. Figure 2. Structure of Blood Vessels. (a) Arteries and (b) veins share the same general features, but the walls of arteries are much thicker because of the higher pressure of the blood that flows through them. (c) A micrograph shows the relative differences in thickness. LM × 160. (Micrograph provided by the Regents of the University of Michigan Medical School © 2012)[/caption]

By the time blood has passed through capillaries and entered venules, the pressure initially exerted upon it by heart contractions has diminished. In other words, in comparison to arteries, venules and veins withstand a much lower pressure from the blood that flows through them. Their walls are considerably thinner and their lumens are correspondingly larger in diameter, allowing more blood to flow with less vessel resistance. In addition, many veins of the body, particularly those of the limbs, contain valves that assist the unidirectional flow of blood toward the heart. This is critical because blood flow becomes sluggish in the extremities, as a result of the lower pressure and the effects of gravity.

The walls of arteries and veins are largely composed of living cells and their products (including collagenous and elastic fibers); the cells require nourishment and produce waste. Since blood passes through the larger vessels relatively quickly, there is limited opportunity for blood in the lumen of the vessel to provide nourishment to or remove waste from the vessel’s cells. Further, the walls of the larger vessels are too thick for nutrients to diffuse through to all of the cells. Larger arteries and veins contain small blood vessels within their walls known as the vasa vasorum—literally “vessels of the vessel”—to provide them with this critical exchange. Since the pressure within arteries is relatively high, the vasa vasorum must function in the outer layers of the vessel (see Figure 2) or the pressure exerted by the blood passing through the vessel would collapse it, preventing any exchange from occurring. The lower pressure within veins allows the vasa vasorum to be located closer to the lumen. The restriction of the vasa vasorum to the outer layers of arteries is thought to be one reason that arterial diseases are more common than venous diseases, since its location makes it more difficult to nourish the cells of the arteries and remove waste products. There are also minute nerves within the walls of both types of vessels that control the contraction and dilation of smooth muscle. These minute nerves are known as the nervi vasorum.

Both arteries and veins have the same three distinct tissue layers, called tunics (from the Latin term tunica), for the garments first worn by ancient Romans; the term tunic is also used for some modern garments. From the most interior layer to the outer, these tunics are the tunica intima, the tunica media, and the tunica externa (see Figure 2). Table 1 compares and contrasts the tunics of the arteries and veins.

Comparison of Tunics in Arteries and Veins (Table 1)
Arteries Veins
General appearance Thick walls with small lumens
Generally appear rounded
 Thin walls with large lumens
Generally appear flattened
Tunica intima Endothelium usually appears wavy due to constriction of smooth muscle
Internal elastic membrane present in larger vessels
 Endothelium appears smooth
Internal elastic membrane absent
Tunica media Normally the thickest layer in arteries
Smooth muscle cells and elastic fibers predominate (the proportions of these vary with distance from the heart)
External elastic membrane present in larger vessels
 Normally thinner than the tunica externa
Smooth muscle cells and collagenous fibers predominate
Nervi vasorum and vasa vasorum present
External elastic membrane absent
Tunica externa Normally thinner than the tunica media in all but the largest arteries
Collagenous and elastic fibers
Nervi vasorum and vasa vasorum present
 Normally the thickest layer in veins
Collagenous and smooth fibers predominate
Some smooth muscle fibers
Nervi vasorum and vasa vasorum present

Tunica Intima

The tunica intima (also called the tunica interna) is composed of epithelial and connective tissue layers. Lining the tunica intima is the specialized simple squamous epithelium called the endothelium, which is continuous throughout the entire vascular system, including the lining of the chambers of the heart. Damage to this endothelial lining and exposure of blood to the collagenous fibers beneath is one of the primary causes of clot formation. Until recently, the endothelium was viewed simply as the boundary between the blood in the lumen and the walls of the vessels. Recent studies, however, have shown that it is physiologically critical to such activities as helping to regulate capillary exchange and altering blood flow. The endothelium releases local chemicals called endothelins that can constrict the smooth muscle within the walls of the vessel to increase blood pressure. Uncompensated overproduction of endothelins may contribute to hypertension (high blood pressure) and cardiovascular disease.

Next to the endothelium is the basement membrane, or basal lamina, that effectively binds the endothelium to the connective tissue. The basement membrane provides strength while maintaining flexibility, and it is permeable, allowing materials to pass through it. The thin outer layer of the tunica intima contains a small amount of areolar connective tissue that consists primarily of elastic fibers to provide the vessel with additional flexibility; it also contains some collagenous fibers to provide additional strength.

In larger arteries, there is also a thick, distinct layer of elastic fibers known as the internal elastic membrane (also called the internal elastic lamina) at the boundary with the tunica media. Like the other components of the tunica intima, the internal elastic membrane provides structure while allowing the vessel to stretch. It is permeated with small openings that allow exchange of materials between the tunics. The internal elastic membrane is not apparent in veins. In addition, many veins, particularly in the lower limbs, contain valves formed by sections of thickened endothelium that are reinforced with connective tissue, extending into the lumen.

Under the microscope, the lumen and the entire tunica intima of a vein will appear smooth, whereas those of an artery will normally appear wavy because of the partial constriction of the smooth muscle in the tunica media, the next layer of blood vessel walls.

Tunica Media

The tunica media is the substantial middle layer of the vessel wall (see Figure 2). It is generally the thickest layer in arteries, and it is much thicker in arteries than it is in veins. The tunica media consists of layers of smooth muscle supported by connective tissue that is primarily made up of elastic fibers, most of which are arranged in circular sheets. Toward the outer portion of the tunic, there are also layers of longitudinal muscle. Contraction and relaxation of the circular muscles decrease and increase the diameter of the vessel lumen, respectively. Specifically in arteries, vasoconstriction decreases blood flow as the smooth muscle in the walls of the tunica media contracts, making the lumen narrower and increasing blood pressure. Similarly, vasodilation increases blood flow as the smooth muscle relaxes, allowing the lumen to widen and blood pressure to drop. Both vasoconstriction and vasodilation are regulated in part by small vascular nerves, known as nervi vasorum, or “nerves of the vessel,” that run within the walls of blood vessels. These are generally all sympathetic fibers, although some trigger vasodilation and others induce vasoconstriction, depending upon the nature of the neurotransmitter and receptors located on the target cell. Parasympathetic stimulation does trigger vasodilation as well as erection during sexual arousal in the external genitalia of both sexes. Nervous control over vessels tends to be more generalized than the specific targeting of individual blood vessels. Local controls, discussed later, account for this phenomenon. (Seek additional content for more information on these dynamic aspects of the autonomic nervous system.) Hormones and local chemicals also control blood vessels. Together, these neural and chemical mechanisms reduce or increase blood flow in response to changing body conditions, from exercise to hydration. Regulation of both blood flow and blood pressure is discussed in detail later in this chapter.

The smooth muscle layers of the tunica media are supported by a framework of collagenous fibers that also binds the tunica media to the inner and outer tunics. Along with the collagenous fibers are large numbers of elastic fibers that appear as wavy lines in prepared slides. Separating the tunica media from the outer tunica externa in larger arteries is the external elastic membrane (also called the external elastic lamina), which also appears wavy in slides. This structure is not usually seen in smaller arteries, nor is it seen in veins.

Tunica Externa

The outer tunic, the tunica externa (also called the tunica adventitia), is a substantial sheath of connective tissue composed primarily of collagenous fibers. Some bands of elastic fibers are found here as well. The tunica externa in veins also contains groups of smooth muscle fibers. This is normally the thickest tunic in veins and may be thicker than the tunica media in some larger arteries. The outer layers of the tunica externa are not distinct but rather blend with the surrounding connective tissue outside the vessel, helping to hold the vessel in relative position. If you are able to palpate some of the superficial veins on your upper limbs and try to move them, you will find that the tunica externa prevents this. If the tunica externa did not hold the vessel in place, any movement would likely result in disruption of blood flow.

Arteries

An artery is a blood vessel that conducts blood away from the heart. All arteries have relatively thick walls that can withstand the high pressure of blood ejected from the heart. However, those close to the heart have the thickest walls, containing a high percentage of elastic fibers in all three of their tunics. This type of artery is known as an elastic artery (Figure 3). Vessels larger than 10 mm in diameter are typically elastic. Their abundant elastic fibers allow them to expand, as blood pumped from the ventricles passes through them, and then to recoil after the surge has passed. If artery walls were rigid and unable to expand and recoil, their resistance to blood flow would greatly increase and blood pressure would rise to even higher levels, which would in turn require the heart to pump harder to increase the volume of blood expelled by each pump (the stroke volume) and maintain adequate pressure and flow. Artery walls would have to become even thicker in response to this increased pressure. The elastic recoil of the vascular wall helps to maintain the pressure gradient that drives the blood through the arterial system. An elastic artery is also known as a conducting artery, because the large diameter of the lumen enables it to accept a large volume of blood from the heart and conduct it to smaller branches.

[caption id="" align="aligncenter" width="550"]The left panel shows the cross-section of an elastic artery, the middle panel shows the cross section of a muscular artery, and the right panel shows the cross-section of an arteriole. Figure 3. Types of Arteries and Arterioles. Comparison of the walls of an elastic artery, a muscular artery, and an arteriole is shown. In terms of scale, the diameter of an arteriole is measured in micrometers compared to millimeters for elastic and muscular arteries.[/caption]

Farther from the heart, where the surge of blood has dampened, the percentage of elastic fibers in an artery’s tunica intima decreases and the amount of smooth muscle in its tunica media increases. The artery at this point is described as a muscular artery. The diameter of muscular arteries typically ranges from 0.1 mm to 10 mm. Their thick tunica media allows muscular arteries to play a leading role in vasoconstriction. In contrast, their decreased quantity of elastic fibers limits their ability to expand. Fortunately, because the blood pressure has eased by the time it reaches these more distant vessels, elasticity has become less important.

Notice that although the distinctions between elastic and muscular arteries are important, there is no “line of demarcation” where an elastic artery suddenly becomes muscular. Rather, there is a gradual transition as the vascular tree repeatedly branches. In turn, muscular arteries branch to distribute blood to the vast network of arterioles. For this reason, a muscular artery is also known as a distributing artery.

Arterioles

An arteriole is a very small artery that leads to a capillary. Arterioles have the same three tunics as the larger vessels, but the thickness of each is greatly diminished. The critical endothelial lining of the tunica intima is intact. The tunica media is restricted to one or two smooth muscle cell layers in thickness. The tunica externa remains but is very thin (see Figure 3).

With a lumen averaging 30 micrometers or less in diameter, arterioles are critical in slowing down—or resisting—blood flow and, thus, causing a substantial drop in blood pressure. Because of this, you may see them referred to as resistance vessels. The muscle fibers in arterioles are normally slightly contracted, causing arterioles to maintain a consistent muscle tone—in this case referred to as vascular tone—in a similar manner to the muscular tone of skeletal muscle. In reality, all blood vessels exhibit vascular tone due to the partial contraction of smooth muscle. The importance of the arterioles is that they will be the primary site of both resistance and regulation of blood pressure. The precise diameter of the lumen of an arteriole at any given moment is determined by neural and chemical controls, and vasoconstriction and vasodilation in the arterioles are the primary mechanisms for distribution of blood flow.

Capillaries

A capillary is a microscopic channel that supplies blood to the tissues themselves, a process called perfusion. Exchange of gases and other substances occurs in the capillaries between the blood and the surrounding cells and their tissue fluid (interstitial fluid). The diameter of a capillary lumen ranges from 5–10 micrometers; the smallest are just barely wide enough for an erythrocyte to squeeze through. Flow through capillaries is often described as microcirculation.

The wall of a capillary consists of the endothelial layer surrounded by a basement membrane with occasional smooth muscle fibers. There is some variation in wall structure: In a large capillary, several endothelial cells bordering each other may line the lumen; in a small capillary, there may be only a single cell layer that wraps around to contact itself.

For capillaries to function, their walls must be leaky, allowing substances to pass through. There are three major types of capillaries, which differ according to their degree of “leakiness:” continuous, fenestrated, and sinusoid capillaries (Figure 4).

Continuous Capillaries

The most common type of capillary, the continuous capillary, is found in almost all vascularized tissues. Continuous capillaries are characterized by a complete endothelial lining with tight junctions between endothelial cells. Although a tight junction is usually impermeable and only allows for the passage of water and ions, they are often incomplete in capillaries, leaving intercellular clefts that allow for exchange of water and other very small molecules between the blood plasma and the interstitial fluid. Substances that can pass between cells include metabolic products, such as glucose, water, and small hydrophobic molecules like gases and hormones, as well as various leukocytes. Continuous capillaries not associated with the brain are rich in transport vesicles, contributing to either endocytosis or exocytosis. Those in the brain are part of the blood-brain barrier. Here, there are tight junctions and no intercellular clefts, plus a thick basement membrane and astrocyte extensions called end feet; these structures combine to prevent the movement of nearly all substances.
[caption id="" align="aligncenter" width="500"]The left panel shows the structure of a continuous capillary, the middle panel shows a fenestrated capillary, and the right panel shows a sinusoid capillary. Figure 4. Types of Capillaries. The three major types of capillaries: continuous, fenestrated, and sinusoid.[/caption]

Fenestrated Capillaries

A fenestrated capillary is one that has pores (or fenestrations) in addition to tight junctions in the endothelial lining. These make the capillary permeable to larger molecules. The number of fenestrations and their degree of permeability vary, however, according to their location. Fenestrated capillaries are common in the small intestine, which is the primary site of nutrient absorption, as well as in the kidneys, which filter the blood. They are also found in the choroid plexus of the brain and many endocrine structures, including the hypothalamus, pituitary, pineal, and thyroid glands.

Sinusoid Capillaries

A sinusoid capillary (or sinusoid) is the least common type of capillary. Sinusoid capillaries are flattened, and they have extensive intercellular gaps and incomplete basement membranes, in addition to intercellular clefts and fenestrations. This gives them an appearance not unlike Swiss cheese. These very large openings allow for the passage of the largest molecules, including plasma proteins and even cells. Blood flow through sinusoids is very slow, allowing more time for exchange of gases, nutrients, and wastes. Sinusoids are found in the liver and spleen, bone marrow, lymph nodes (where they carry lymph, not blood), and many endocrine glands including the pituitary and adrenal glands. Without these specialized capillaries, these organs would not be able to provide their myriad of functions. For example, when bone marrow forms new blood cells, the cells must enter the blood supply and can only do so through the large openings of a sinusoid capillary; they cannot pass through the small openings of continuous or fenestrated capillaries. The liver also requires extensive specialized sinusoid capillaries in order to process the materials brought to it by the hepatic portal vein from both the digestive tract and spleen, and to release plasma proteins into circulation.

Metarterioles and Capillary Beds

A metarteriole is a type of vessel that has structural characteristics of both an arteriole and a capillary. Slightly larger than the typical capillary, the smooth muscle of the tunica media of the metarteriole is not continuous but forms rings of smooth muscle (sphincters) prior to the entrance to the capillaries. Each metarteriole arises from a terminal arteriole and branches to supply blood to a capillary bed that may consist of 10–100 capillaries.

The precapillary sphincters, circular smooth muscle cells that surround the capillary at its origin with the metarteriole, tightly regulate the flow of blood from a metarteriole to the capillaries it supplies. Their function is critical: If all of the capillary beds in the body were to open simultaneously, they would collectively hold every drop of blood in the body and there would be none in the arteries, arterioles, venules, veins, or the heart itself. Normally, the precapillary sphincters are closed. When the surrounding tissues need oxygen and have excess waste products, the precapillary sphincters open, allowing blood to flow through and exchange to occur before closing once more (Figure 5). If all of the precapillary sphincters in a capillary bed are closed, blood will flow from the metarteriole directly into a thoroughfare channel and then into the venous circulation, bypassing the capillary bed entirely. This creates what is known as a vascular shunt. In addition, an arteriovenous anastomosis may bypass the capillary bed and lead directly to the venous system. Although you might expect blood flow through a capillary bed to be smooth, in reality, it moves with an irregular, pulsating flow. This pattern is called vasomotion and is regulated by chemical signals that are triggered in response to changes in internal conditions, such as oxygen, carbon dioxide, hydrogen ion, and lactic acid levels. For example, during strenuous exercise when oxygen levels decrease and carbon dioxide, hydrogen ion, and lactic acid levels all increase, the capillary beds in skeletal muscle are open, as they would be in the digestive system when nutrients are present in the digestive tract. During sleep or rest periods, vessels in both areas are largely closed; they open only occasionally to allow oxygen and nutrient supplies to travel to the tissues to maintain basic life processes.
[caption id="" align="aligncenter" width="480"]This diagram shows a capillary bed connecting an arteriole and a venule. Figure 5. Capillary Bed. In a capillary bed, arterioles give rise to metarterioles. Precapillary sphincters located at the junction of a metarteriole with a capillary regulate blood flow. A thoroughfare channel connects the metarteriole to a venule. An arteriovenous anastomosis, which directly connects the arteriole with the venule, is shown at the bottom.[/caption]

Venules

A venule is an extremely small vein, generally 8–100 micrometers in diameter. Postcapillary venules join multiple capillaries exiting from a capillary bed. Multiple venules join to form veins. The walls of venules consist of endothelium, a thin middle layer with a few muscle cells and elastic fibers, plus an outer layer of connective tissue fibers that constitute a very thin tunica externa (Figure 6). Venules as well as capillaries are the primary sites of emigration or diapedesis, in which the white blood cells adhere to the endothelial lining of the vessels and then squeeze through adjacent cells to enter the tissue fluid.

Veins

A vein is a blood vessel that conducts blood toward the heart. Compared to arteries, veins are thin-walled vessels with large and irregular lumens (see Figure 6). Because they are low-pressure vessels, larger veins are commonly equipped with valves that promote the unidirectional flow of blood toward the heart and prevent backflow toward the capillaries caused by the inherent low blood pressure in veins as well as the pull of gravity. Table 2 compares the features of arteries and veins.

[caption id="" align="aligncenter" width="390"]The top panel shows the cross-section of a large vein, the middle panel shows the cross-section of a medium sized vein, and the bottom panel shows the cross-section of a venule. Figure 6. Comparison of Veins and Venules. Many veins have valves to prevent back flow of blood, whereas venules do not. In terms of scale, the diameter of a venule is measured in micrometers compared to millimeters for veins.[/caption]
Comparison of Arteries and Veins (Table 2)
Arteries Veins
Direction of blood flow Conducts blood away from the heart Conducts blood toward the heart
General appearance Rounded Irregular, often collapsed
Pressure High Low
Wall thickness Thick Thin
Relative oxygen concentration Higher in systemic arteries
Lower in pulmonary arteries
 Lower in systemic veins
Higher in pulmonary veins
Valves Not present Present most commonly in limbs and in veins inferior to the heart

Veins as Blood Reservoirs

In addition to their primary function of returning blood to the heart, veins may be considered blood reservoirs, since systemic veins contain approximately 64 percent of the blood volume at any given time (Figure 8). Their ability to hold this much blood is due to their high capacitance, that is, their capacity to distend (expand) readily to store a high volume of blood, even at a low pressure. The large lumens and relatively thin walls of veins make them far more distensible than arteries; thus, they are said to be capacitance vessels.

[caption id="" align="aligncenter" width="450"]This table describes the distribution of blood flow. 84 percent of blood flow is systemic circulation, of which 64 percent happens in systemic veins (18 percent in large veins, 21 percent in large venous networks such as liver, bone marrow, and integument, and 25 percent in venules and medium-sized veins); 13 percent happens in systemic arteries (2 percent in arterioles, 5 percent in muscular arteries, 4 percent in elastic arteries, and 2 percent in the aorta); and 7 percent happens in systemic capillaries. 9 percent of blood flow is pulmonary circulation, of which 4 percent happens in pulmonary veins, 2 percent happens in pulmonary capillaries, and 3 percent happens in pulmonary arteries. The remaining 7 percent of blood flow is in the heart. Figure 8. Distribution of Blood Flow[/caption]

When blood flow needs to be redistributed to other portions of the body, the vasomotor center located in the medulla oblongata sends sympathetic stimulation to the smooth muscles in the walls of the veins, causing constriction—or in this case, venoconstriction. Less dramatic than the vasoconstriction seen in smaller arteries and arterioles, venoconstriction may be likened to a “stiffening” of the vessel wall. This increases pressure on the blood within the veins, speeding its return to the heart. As you will note in Figure 8, approximately 21 percent of the venous blood is located in venous networks within the liver, bone marrow, and integument. This volume of blood is referred to as venous reserve. Through venoconstriction, this “reserve” volume of blood can get back to the heart more quickly for redistribution to other parts of the circulation.

]]> 2271 2261 2 0 20.2 Blood Flow, Blood Pressure, and Resistance https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2277 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2277

Learning Objectives

By the end of this section, you will be able to:
  • Distinguish between systolic pressure, diastolic pressure, pulse pressure, and mean arterial pressure
  • Describe the clinical measurement of pulse and blood pressure
  • Identify and discuss five variables affecting arterial blood flow and blood pressure
  • Discuss several factors affecting blood flow in the venous system
Blood flow refers to the movement of blood through a vessel, tissue, or organ, and is usually expressed in terms of volume of blood per unit of time. It is initiated by the contraction of the ventricles of the heart. Ventricular contraction ejects blood into the major arteries, resulting in flow from regions of higher pressure to regions of lower pressure, as blood encounters smaller arteries and arterioles, then capillaries, then the venules and veins of the venous system. This section discusses a number of critical variables that contribute to blood flow throughout the body. It also discusses the factors that impede or slow blood flow, a phenomenon known as resistance.

As noted earlier, hydrostatic pressure is the force exerted by a fluid due to gravitational pull, usually against the wall of the container in which it is located. One form of hydrostatic pressure is blood pressure, the force exerted by blood upon the walls of the blood vessels or the chambers of the heart. Blood pressure may be measured in capillaries and veins, as well as the vessels of the pulmonary circulation; however, the term blood pressure without any specific descriptors typically refers to systemic arterial blood pressure—that is, the pressure of blood flowing in the arteries of the systemic circulation. In clinical practice, this pressure is measured in mm Hg and is usually obtained using the brachial artery of the arm.

Components of Arterial Blood Pressure

Arterial blood pressure in the larger vessels consists of several distinct components (Figure 1): systolic and diastolic pressures, pulse pressure, and mean arterial pressure.

Systolic and Diastolic Pressures

When systemic arterial blood pressure is measured, it is recorded as a ratio of two numbers (e.g., 120/80 is a normal adult blood pressure), expressed as systolic pressure over diastolic pressure. The systolic pressure is the higher value (typically around 120 mm Hg) and reflects the arterial pressure resulting from the ejection of blood during ventricular contraction, or systole. The diastolic pressure is the lower value (usually about 80 mm Hg) and represents the arterial pressure of blood during ventricular relaxation, or diastole.

[caption id="" align="aligncenter" width="500"]This graph shows the value of pulse pressure in different types of blood vessels. Figure 1. Systemic Blood Pressure. The graph shows the components of blood pressure throughout the blood vessels, including systolic, diastolic, mean arterial, and pulse pressures.[/caption]

Pulse Pressure

As shown in Figure 1, the difference between the systolic pressure and the diastolic pressure is the pulse pressure. For example, an individual with a systolic pressure of 120 mm Hg and a diastolic pressure of 80 mm Hg would have a pulse pressure of 40 mmHg.

Generally, a pulse pressure should be at least 25 percent of the systolic pressure. A pulse pressure below this level is described as low or narrow. This may occur, for example, in patients with a low stroke volume, which may be seen in congestive heart failure, stenosis of the aortic valve, or significant blood loss following trauma. In contrast, a high or wide pulse pressure is common in healthy people following strenuous exercise, when their resting pulse pressure of 30–40 mm Hg may increase temporarily to 100 mm Hg as stroke volume increases. A persistently high pulse pressure at or above 100 mm Hg may indicate excessive resistance in the arteries and can be caused by a variety of disorders. Chronic high resting pulse pressures can degrade the heart, brain, and kidneys, and warrant medical treatment.

Mean Arterial Pressure

Mean arterial pressure (MAP) represents the “average” pressure of blood in the arteries, that is, the average force driving blood into vessels that serve the tissues. Mean is a statistical concept and is calculated by taking the sum of the values divided by the number of values. Although complicated to measure directly and complicated to calculate, MAP can be approximated by adding the diastolic pressure to one-third of the pulse pressure or systolic pressure minus the diastolic pressure:

MAP = diastolic BP + ((systolic-diastolic BP) / 3)

In Figure 1, this value is approximately 80 + (120 − 80) / 3, or 93.33. Normally, the MAP falls within the range of 70–110 mm Hg. If the value falls below 60 mm Hg for an extended time, blood pressure will not be high enough to ensure circulation to and through the tissues, which results in ischemia, or insufficient blood flow. A condition called hypoxia, inadequate oxygenation of tissues, commonly accompanies ischemia. The term hypoxemia refers to low levels of oxygen in systemic arterial blood. Neurons are especially sensitive to hypoxia and may die or be damaged if blood flow and oxygen supplies are not quickly restored.

Pulse

After blood is ejected from the heart, elastic fibers in the arteries help maintain a high-pressure gradient as they expand to accommodate the blood, then recoil. This expansion and recoiling effect, known as the pulse, can be palpated manually or measured electronically. Although the effect diminishes over distance from the heart, elements of the systolic and diastolic components of the pulse are still evident down to the level of the arterioles.

Because pulse indicates heart rate, it is measured clinically to provide clues to a patient’s state of health. It is recorded as beats per minute. Both the rate and the strength of the pulse are important clinically. A high or irregular pulse rate can be caused by physical activity or other temporary factors, but it may also indicate a heart condition. The pulse strength indicates the strength of ventricular contraction and cardiac output. If the pulse is strong, then systolic pressure is high. If it is weak, systolic pressure has fallen, and medical intervention may be warranted.

Pulse can be palpated manually by placing the tips of the fingers across an artery that runs close to the body surface and pressing lightly. While this procedure is normally performed using the radial artery in the wrist or the common carotid artery in the neck, any superficial artery that can be palpated may be used (Figure 2). Common sites to find a pulse include temporal and facial arteries in the head, brachial arteries in the upper arm, femoral arteries in the thigh, popliteal arteries behind the knees, posterior tibial arteries near the medial tarsal regions, and dorsalis pedis arteries in the feet. A variety of commercial electronic devices are also available to measure pulse.
[caption id="" align="aligncenter" width="320"]This image shows the pulse points in a woman’s body. Figure 2. Pulse Sites. The pulse is most readily measured at the radial artery, but can be measured at any of the pulse points shown.[/caption]

Measurement of Blood Pressure

Blood pressure is one of the critical parameters measured on virtually every patient in every healthcare setting. The technique used today was developed more than 100 years ago by a pioneering Russian physician, Dr. Nikolai Korotkoff. Turbulent blood flow through the vessels can be heard as a soft ticking while measuring blood pressure; these sounds are known as Korotkoff sounds. The technique of measuring blood pressure requires the use of a sphygmomanometer (a blood pressure cuff attached to a measuring device) and a stethoscope. The technique is as follows:

  • The clinician wraps an inflatable cuff tightly around the patient’s arm at about the level of the heart.
  • The clinician squeezes a rubber pump to inject air into the cuff, raising pressure around the artery and temporarily cutting off blood flow into the patient’s arm.
  • The clinician places the stethoscope on the patient’s antecubital region and, while gradually allowing air within the cuff to escape, listens for the Korotkoff sounds.
Although there are five recognized Korotkoff sounds, only two are normally recorded. Initially, no sounds are heard since there is no blood flow through the vessels, but as air pressure drops, the cuff relaxes, and blood flow returns to the arm. As shown in Figure 3, the first sound heard through the stethoscope—the first Korotkoff sound—indicates systolic pressure. As more air is released from the cuff, blood is able to flow freely through the brachial artery and all sounds disappear. The point at which the last sound is heard is recorded as the patient’s diastolic pressure.
[caption id="" align="aligncenter" width="450"]This image shows blood pressure as a function of time. Figure 3. Blood Pressure Measurement. When pressure in a sphygmomanometer cuff is released, a clinician can hear the Korotkoff sounds. In this graph, a blood pressure tracing is aligned to a measurement of systolic and diastolic pressures.[/caption]

The majority of hospitals and clinics have automated equipment for measuring blood pressure that work on the same principles. An even more recent innovation is a small instrument that wraps around a patient’s wrist. The patient then holds the wrist over the heart while the device measures blood flow and records pressure.

Variables Affecting Blood Flow and Blood Pressure

Five variables influence blood flow and blood pressure:

  • Cardiac output
  • Compliance
  • Volume of the blood
  • Viscosity of the blood
  • Blood vessel length and diameter
Recall that blood moves from higher pressure to lower pressure. It is pumped from the heart into the arteries at high pressure. If you increase pressure in the arteries (afterload), and cardiac function does not compensate, blood flow will actually decrease. In the venous system, the opposite relationship is true. Increased pressure in the veins does not decrease flow as it does in arteries, but actually increases flow. Since pressure in the veins is normally relatively low, for blood to flow back into the heart, the pressure in the atria during atrial diastole must be even lower. It normally approaches zero, except when the atria contract (see Figure 1).

Cardiac Output

Cardiac output is the measurement of blood flow from the heart through the ventricles, and is usually measured in liters per minute. Any factor that causes cardiac output to increase, by elevating heart rate or stroke volume or both, will elevate blood pressure and promote blood flow. These factors include sympathetic stimulation, the catecholamines epinephrine and norepinephrine, thyroid hormones, and increased calcium ion levels. Conversely, any factor that decreases cardiac output, by decreasing heart rate or stroke volume or both, will decrease arterial pressure and blood flow. These factors include parasympathetic stimulation, elevated or decreased potassium ion levels, decreased calcium levels, anoxia, and acidosis.

Compliance

Compliance is the ability of any compartment to expand to accommodate increased content. A metal pipe, for example, is not compliant, whereas a balloon is. The greater the compliance of an artery, the more effectively it is able to expand to accommodate surges in blood flow without increased resistance or blood pressure. Veins are more compliant than arteries and can expand to hold more blood. When vascular disease causes stiffening of arteries, compliance is reduced and resistance to blood flow is increased. The result is more turbulence, higher pressure within the vessel, and reduced blood flow. This increases the work of the heart.

A Mathematical Approach to Factors Affecting Blood Flow

Jean Louis Marie Poiseuille was a French physician and physiologist who devised a mathematical equation describing blood flow and its relationship to known parameters. The same equation also applies to engineering studies of the flow of fluids. Although understanding the math behind the relationships among the factors affecting blood flow is not necessary to understand blood flow, it can help solidify an understanding of their relationships. Please note that even if the equation looks intimidating, breaking it down into its components and following the relationships will make these relationships clearer, even if you are weak in math. Focus on the three critical variables: radius (r), vessel length (λ), and viscosity (η).

Poiseuille’s equation:
Blood flow = π ΔP r48ηλ
  • π is the Greek letter pi, used to represent the mathematical constant that is the ratio of a circle’s circumference to its diameter. It may commonly be represented as 3.14, although the actual number extends to infinity.
  • ΔP represents the difference in pressure.
  • r4 is the radius (one-half of the diameter) of the vessel to the fourth power.
  • η is the Greek letter eta and represents the viscosity of the blood.
  • λ is the Greek letter lambda and represents the length of a blood vessel.

One of several things this equation allows us to do is calculate the resistance in the vascular system. Normally this value is extremely difficult to measure, but it can be calculated from this known relationship:

Blood flow = ΔP/Resistance
If we rearrange this slightly,
Resistance = ΔP/Blood flow

Then by substituting Pouseille’s equation for blood flow:

Resistance =8ηλ/πr4
By examining this equation, you can see that there are only three variables: viscosity, vessel length, and radius, since 8 and π are both constants. The important thing to remember is this: Two of these variables, viscosity and vessel length, will change slowly in the body. Only one of these factors, the radius, can be changed rapidly by vasoconstriction and vasodilation, thus dramatically impacting resistance and flow. Further, small changes in the radius will greatly affect flow, since it is raised to the fourth power in the equation.

We have briefly considered how cardiac output and blood volume impact blood flow and pressure; the next step is to see how the other variables (contraction, vessel length, and viscosity) articulate with Pouseille’s equation and what they can teach us about the impact on blood flow.

Blood Volume

The relationship between blood volume, blood pressure, and blood flow is intuitively obvious. Water may merely trickle along a creek bed in a dry season, but rush quickly and under great pressure after a heavy rain. Similarly, as blood volume decreases, pressure and flow decrease. As blood volume increases, pressure and flow increase.

Under normal circumstances, blood volume varies little. Low blood volume, called hypovolemia, may be caused by bleeding, dehydration, vomiting, severe burns, or some medications used to treat hypertension. It is important to recognize that other regulatory mechanisms in the body are so effective at maintaining blood pressure that an individual may be asymptomatic until 10–20 percent of the blood volume has been lost. Treatment typically includes intravenous fluid replacement.

Hypervolemia, excessive fluid volume, may be caused by retention of water and sodium, as seen in patients with heart failure, liver cirrhosis, some forms of kidney disease, hyperaldosteronism, and some glucocorticoid steroid treatments. Restoring homeostasis in these patients depends upon reversing the condition that triggered the hypervolemia.

Blood Viscosity

Viscosity is the thickness of fluids that affects their ability to flow. Clean water, for example, is less viscous than mud. The viscosity of blood is directly proportional to resistance and inversely proportional to flow; therefore, any condition that causes viscosity to increase will also increase resistance and decrease flow. For example, imagine sipping milk, then a milkshake, through the same size straw. You experience more resistance and therefore less flow from the milkshake. Conversely, any condition that causes viscosity to decrease (such as when the milkshake melts) will decrease resistance and increase flow.

Normally the viscosity of blood does not change over short periods of time. The two primary determinants of blood viscosity are the formed elements and plasma proteins. Since the vast majority of formed elements are erythrocytes, any condition affecting erythropoiesis, such as polycythemia or anemia, can alter viscosity. Since most plasma proteins are produced by the liver, any condition affecting liver function can also change the viscosity slightly and therefore decrease blood flow. Liver abnormalities include hepatitis, cirrhosis, alcohol damage, and drug toxicities. While leukocytes and platelets are normally a small component of the formed elements, there are some rare conditions in which severe overproduction can impact viscosity as well.

Vessel Length and Diameter

The length of a vessel is directly proportional to its resistance: the longer the vessel, the greater the resistance and the lower the flow. As with blood volume, this makes intuitive sense, since the increased surface area of the vessel will impede the flow of blood. Likewise, if the vessel is shortened, the resistance will decrease and flow will increase.

The length of our blood vessels increases throughout childhood as we grow, of course, but is unchanging in adults under normal physiological circumstances. Further, the distribution of vessels is not the same in all tissues. Adipose tissue does not have an extensive vascular supply. One pound of adipose tissue contains approximately 200 miles of vessels, whereas skeletal muscle contains more than twice that. Overall, vessels decrease in length only during loss of mass or amputation. An individual weighing 150 pounds has approximately 60,000 miles of vessels in the body. Gaining about 10 pounds adds from 2000 to 4000 miles of vessels, depending upon the nature of the gained tissue. One of the great benefits of weight reduction is the reduced stress to the heart, which does not have to overcome the resistance of as many miles of vessels.

In contrast to length, the diameter of blood vessels changes throughout the body, according to the type of vessel, as we discussed earlier. The diameter of any given vessel may also change frequently throughout the day in response to neural and chemical signals that trigger vasodilation and vasoconstriction. The vascular tone of the vessel is the contractile state of the smooth muscle and the primary determinant of diameter, and thus of resistance and flow. The effect of vessel diameter on resistance is inverse: Given the same volume of blood, an increased diameter means there is less blood contacting the vessel wall, thus lower friction and lower resistance, subsequently increasing flow. A decreased diameter means more of the blood contacts the vessel wall, and resistance increases, subsequently decreasing flow.

The influence of lumen diameter on resistance is dramatic: A slight increase or decrease in diameter causes a huge decrease or increase in resistance. This is because resistance is inversely proportional to the radius of the blood vessel (one-half of the vessel’s diameter) raised to the fourth power (R = 1/r4). This means, for example, that if an artery or arteriole constricts to one-half of its original radius, the resistance to flow will increase 16 times. And if an artery or arteriole dilates to twice its initial radius, then resistance in the vessel will decrease to 1/16 of its original value and flow will increase 16 times.

The Roles of Vessel Diameter and Total Area in Blood Flow and Blood Pressure

Recall that we classified arterioles as resistance vessels, because given their small lumen, they dramatically slow the flow of blood from arteries. In fact, arterioles are the site of greatest resistance in the entire vascular network. This may seem surprising, given that capillaries have a smaller size. How can this phenomenon be explained?

Figure 4 compares vessel diameter, total cross-sectional area, average blood pressure, and blood velocity through the systemic vessels. Notice in parts (a) and (b) that the total cross-sectional area of the body’s capillary beds is far greater than any other type of vessel. Although the diameter of an individual capillary is significantly smaller than the diameter of an arteriole, there are vastly more capillaries in the body than there are other types of blood vessels. Part (c) shows that blood pressure drops unevenly as blood travels from arteries to arterioles, capillaries, venules, and veins, and encounters greater resistance. However, the site of the most precipitous drop, and the site of greatest resistance, is the arterioles. This explains why vasodilation and vasoconstriction of arterioles play more significant roles in regulating blood pressure than do the vasodilation and vasoconstriction of other vessels.

Part (d) shows that the velocity (speed) of blood flow decreases dramatically as the blood moves from arteries to arterioles to capillaries. This slow flow rate allows more time for exchange processes to occur. As blood flows through the veins, the rate of velocity increases, as blood is returned to the heart.
[caption id="" align="aligncenter" width="520"]This figure shows four graphs. The top left graph shows the vessel diameter for different types of blood vessels. The top right panel shows cross-sectional area for different blood vessels. The bottom left panel shows the average blood pressure for different blood vessels, and the bottom right panel shows the velocity of blood flow in different blood vessels. Figure 4. Relationships among Vessels in the Systemic Circuit. The relationships among blood vessels that can be compared include (a) vessel diameter, (b) total cross-sectional area, (c) average blood pressure, and (d) velocity of blood flow.[/caption]

Venous System

The pumping action of the heart propels the blood into the arteries, from an area of higher pressure toward an area of lower pressure. If blood is to flow from the veins back into the heart, the pressure in the veins must be greater than the pressure in the atria of the heart. Two factors help maintain this pressure gradient between the veins and the heart. First, the pressure in the atria during diastole is very low, often approaching zero when the atria are relaxed (atrial diastole). Second, two physiologic “pumps” increase pressure in the venous system. The use of the term “pump” implies a physical device that speeds flow. These physiological pumps are less obvious.

Skeletal Muscle Pump

In many body regions, the pressure within the veins can be increased by the contraction of the surrounding skeletal muscle. This mechanism, known as the skeletal muscle pump (Figure 6), helps the lower-pressure veins counteract the force of gravity, increasing pressure to move blood back to the heart. As leg muscles contract, for example during walking or running, they exert pressure on nearby veins with their numerous one-way valves. This increased pressure causes blood to flow upward, opening valves superior to the contracting muscles so blood flows through. Simultaneously, valves inferior to the contracting muscles close; thus, blood should not seep back downward toward the feet. Military recruits are trained to flex their legs slightly while standing at attention for prolonged periods. Failure to do so may allow blood to pool in the lower limbs rather than returning to the heart. Consequently, the brain will not receive enough oxygenated blood, and the individual may lose consciousness.

[caption id="" align="aligncenter" width="380"]The left panel shows the structure of a skeletal muscle vein pump when the muscle is relaxed, and the right panel shows the structure of a skeletal muscle vein pump when the muscle is contracted. Figure 6. Skeletal Muscle Pump. The contraction of skeletal muscles surrounding a vein compresses the blood and increases the pressure in that area. This action forces blood closer to the heart where venous pressure is lower. Note the importance of the one-way valves to assure that blood flows only in the proper direction.[/caption]

Respiratory Pump

The respiratory pump aids blood flow through the veins of the thorax and abdomen. During inhalation, the volume of the thorax increases, largely through the contraction of the diaphragm, which moves downward and compresses the abdominal cavity. The elevation of the chest caused by the contraction of the external intercostal muscles also contributes to the increased volume of the thorax. The volume increase causes air pressure within the thorax to decrease, allowing us to inhale. Additionally, as air pressure within the thorax drops, blood pressure in the thoracic veins also decreases, falling below the pressure in the abdominal veins. This causes blood to flow along its pressure gradient from veins outside the thorax, where pressure is higher, into the thoracic region, where pressure is now lower. This in turn promotes the return of blood from the thoracic veins to the atria. During exhalation, when air pressure increases within the thoracic cavity, pressure in the thoracic veins increases, speeding blood flow into the heart while valves in the veins prevent blood from flowing backward from the thoracic and abdominal veins.

Pressure Relationships in the Venous System

Although vessel diameter increases from the smaller venules to the larger veins and eventually to the venae cavae (singular = vena cava), the total cross-sectional area actually decreases (see Figure 6a and b). The individual veins are larger in diameter than the venules, but their total number is much lower, so their total cross-sectional area is also lower.

Also notice that, as blood moves from venules to veins, the average blood pressure drops (see Figure 6c), but the blood velocity actually increases (see Figure 6). This pressure gradient drives blood back toward the heart. Again, the presence of one-way valves and the skeletal muscle and respiratory pumps contribute to this increased flow. Since approximately 64 percent of the total blood volume resides in systemic veins, any action that increases the flow of blood through the veins will increase venous return to the heart. Maintaining vascular tone within the veins prevents the veins from merely distending, dampening the flow of blood, and as you will see, vasoconstriction actually enhances the flow.

The Role of Venoconstriction in Resistance, Blood Pressure, and Flow

As previously discussed, vasoconstriction of an artery or arteriole decreases the radius, increasing resistance and pressure, but decreasing flow. Venoconstriction, on the other hand, has a very different outcome. The walls of veins are thin but irregular; thus, when the smooth muscle in those walls constricts, the lumen becomes more rounded. The more rounded the lumen, the less surface area the blood encounters, and the less resistance the vessel offers. Vasoconstriction increases pressure within a vein as it does in an artery, but in veins, the increased pressure increases flow. Recall that the pressure in the atria, into which the venous blood will flow, is very low, approaching zero for at least part of the relaxation phase of the cardiac cycle. Thus, venoconstriction increases the return of blood to the heart. Another way of stating this is that venoconstriction increases the preload or stretch of the cardiac muscle and increases contraction.

]]> 2277 2261 3 0 20.3 Capillary Exchange https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2279 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2279

Learning Objectives

By the end of this section, you will be able to:
  • Identify the primary mechanisms of capillary exchange
  • Distinguish between capillary hydrostatic pressure and blood colloid osmotic pressure, explaining the contribution of each to net filtration pressure
  • Compare filtration and reabsorption
  • Explain the fate of fluid that is not reabsorbed from the tissues into the vascular capillaries

The primary purpose of the cardiovascular system is to circulate gases, nutrients, wastes, and other substances to and from the cells of the body. Small molecules, such as gases, lipids, and lipid-soluble molecules, can diffuse directly through the membranes of the endothelial cells of the capillary wall. Glucose, amino acids, and ions—including sodium, potassium, calcium, and chloride—use transporters to move through specific channels in the membrane by facilitated diffusion. Glucose, ions, and larger molecules may also leave the blood through intercellular clefts. Larger molecules can pass through the pores of fenestrated capillaries, and even large plasma proteins can pass through the great gaps in the sinusoids. Some large proteins in blood plasma can move into and out of the endothelial cells packaged within vesicles by endocytosis and exocytosis. Water moves by osmosis.

Bulk Flow

The mass movement of fluids into and out of capillary beds requires a transport mechanism far more efficient than mere diffusion. This movement, often referred to as bulk flow, involves two pressure-driven mechanisms: Volumes of fluid move from an area of higher pressure in a capillary bed to an area of lower pressure in the tissues via filtration. In contrast, the movement of fluid from an area of higher pressure in the tissues into an area of lower pressure in the capillaries is reabsorption. Two types of pressure interact to drive each of these movements: hydrostatic pressure and osmotic pressure.

Hydrostatic Pressure

The primary force driving fluid transport between the capillaries and tissues is hydrostatic pressure, which can be defined as the pressure of any fluid enclosed in a space. Blood hydrostatic pressure is the force exerted by the blood confined within blood vessels or heart chambers. Even more specifically, the pressure exerted by blood against the wall of a capillary is called capillary hydrostatic pressure (CHP), and is the same as capillary blood pressure. CHP is the force that drives fluid out of capillaries and into the tissues.

As fluid exits a capillary and moves into tissues, the hydrostatic pressure in the interstitial fluid correspondingly rises. This opposing hydrostatic pressure is called the interstitial fluid hydrostatic pressure (IFHP). Generally, the CHP originating from the arterial pathways is considerably higher than the IFHP, because lymphatic vessels are continually absorbing excess fluid from the tissues. Thus, fluid generally moves out of the capillary and into the interstitial fluid. This process is called filtration.

Osmotic Pressure

The net pressure that drives reabsorption—the movement of fluid from the interstitial fluid back into the capillaries—is called osmotic pressure (sometimes referred to as oncotic pressure). Whereas hydrostatic pressure forces fluid out of the capillary, osmotic pressure draws fluid back in. Osmotic pressure is determined by osmotic concentration gradients, that is, the difference in the solute-to-water concentrations in the blood and tissue fluid. A region higher in solute concentration (and lower in water concentration) draws water across a semipermeable membrane from a region higher in water concentration (and lower in solute concentration).

As we discuss osmotic pressure in blood and tissue fluid, it is important to recognize that the formed elements of blood do not contribute to osmotic concentration gradients. Rather, it is the plasma proteins that play the key role. Solutes also move across the capillary wall according to their concentration gradient, but overall, the concentrations should be similar and not have a significant impact on osmosis. Because of their large size and chemical structure, plasma proteins are not truly solutes, that is, they do not dissolve but are dispersed or suspended in their fluid medium, forming a colloid rather than a solution.

The pressure created by the concentration of colloidal proteins in the blood is called the blood colloidal osmotic pressure (BCOP). Its effect on capillary exchange accounts for the reabsorption of water. The plasma proteins suspended in blood cannot move across the semipermeable capillary cell membrane, and so they remain in the plasma. As a result, blood has a higher colloidal concentration and lower water concentration than tissue fluid. It therefore attracts water. We can also say that the BCOP is higher than the interstitial fluid colloidal osmotic pressure (IFCOP), which is always very low because interstitial fluid contains few proteins. Thus, water is drawn from the tissue fluid back into the capillary, carrying dissolved molecules with it. This difference in colloidal osmotic pressure accounts for reabsorption.

Interaction of Hydrostatic and Osmotic Pressures

The normal unit used to express pressures within the cardiovascular system is millimeters of mercury (mm Hg). When blood leaving an arteriole first enters a capillary bed, the CHP is quite high—about 35 mm Hg. Gradually, this initial CHP declines as the blood moves through the capillary so that by the time the blood has reached the venous end, the CHP has dropped to approximately 18 mm Hg. In comparison, the plasma proteins remain suspended in the blood, so the BCOP remains fairly constant at about 25 mm Hg throughout the length of the capillary and considerably below the osmotic pressure in the interstitial fluid.

The net filtration pressure (NFP) represents the interaction of the hydrostatic and osmotic pressures, driving fluid out of the capillary. It is equal to the difference between the CHP and the BCOP. Since filtration is, by definition, the movement of fluid out of the capillary, when reabsorption is occurring, the NFP is a negative number.

NFP changes at different points in a capillary bed (Figure 1). Close to the arterial end of the capillary, it is approximately 10 mm Hg, because the CHP of 35 mm Hg minus the BCOP of 25 mm Hg equals 10 mm Hg. Recall that the hydrostatic and osmotic pressures of the interstitial fluid are essentially negligible. Thus, the NFP of 10 mm Hg drives a net movement of fluid out of the capillary at the arterial end. At approximately the middle of the capillary, the CHP is about the same as the BCOP of 25 mm Hg, so the NFP drops to zero. At this point, there is no net change of volume: Fluid moves out of the capillary at the same rate as it moves into the capillary. Near the venous end of the capillary, the CHP has dwindled to about 18 mm Hg due to loss of fluid. Because the BCOP remains steady at 25 mm Hg, water is drawn into the capillary, that is, reabsorption occurs. Another way of expressing this is to say that at the venous end of the capillary, there is an NFP of −7 mm Hg.

[caption id="" align="aligncenter" width="520"]This diagram shows the process of fluid exchange in a capillary from the arterial end to the venous end. Figure 1. Capillary Exchange. Net filtration occurs near the arterial end of the capillary since capillary hydrostatic pressure (CHP) is greater than blood colloidal osmotic pressure (BCOP). There is no net movement of fluid near the midpoint since CHP = BCOP. Net reabsorption occurs near the venous end since BCOP is greater than CHP.[/caption]
]]> 2279 2261 4 0 20.4 Homeostatic Regulation of the Vascular System https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2286 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2286

Learning Objectives

By the end of this section, you will be able to:
  • Discuss the mechanisms involved in the neural regulation of vascular homeostasis
  • Describe the contribution of a variety of hormones to the renal regulation of blood pressure
  • Identify the effects of exercise on vascular homeostasis
  • Discuss how hypertension, hemorrhage, and circulatory shock affect vascular health

In order to maintain homeostasis in the cardiovascular system and provide adequate blood to the tissues, blood flow must be redirected continually to the tissues as they become more active. In a very real sense, the cardiovascular system engages in resource allocation, because there is not enough blood flow to distribute blood equally to all tissues simultaneously. For example, when an individual is exercising, more blood will be directed to skeletal muscles, the heart, and the lungs. Following a meal, more blood is directed to the digestive system. Only the brain receives a more or less constant supply of blood whether you are active, resting, thinking, or engaged in any other activity.

Table 3 provides the distribution of systemic blood at rest and during exercise. Although most of the data appears logical, the values for the distribution of blood to the integument may seem surprising. During exercise, the body distributes more blood to the body surface where it can dissipate the excess heat generated by increased activity into the environment.
Systemic Blood Flow During Rest, Mild Exercise, and Maximal Exercise in a Healthy Young Individual (Table 3)
Organ Resting
(mL/min)
 Mild exercise
(mL/min)
 Maximal exercise
(mL/min)
Skeletal muscle 1200 4500 12,500
Heart 250 350 750
Brain 750 750 750
Integument 500 1500 1900
Kidney 1100 900 600
Gastrointestinal 1400 1100 600
Others
(i.e., liver, spleen)
 600 400 400
Total 5800 9500 17,500
Three homeostatic mechanisms ensure adequate blood flow, blood pressure, distribution, and ultimately perfusion: neural, endocrine, and autoregulatory mechanisms. They are summarized in Figure 1.
[caption id="" align="aligncenter" width="580"]This flowchart shows the various factors that control the flow of blood. The top panel focuses on autoregulation, and the bottom panel focuses on neural and endocrine mechanisms. Figure 1. Summary of Factors Maintaining Vascular Homeostasis. Adequate blood flow, blood pressure, distribution, and perfusion involve autoregulatory, neural, and endocrine mechanisms.[/caption]

Neural Regulation

The nervous system plays a critical role in the regulation of vascular homeostasis. The primary regulatory sites include the cardiovascular centers in the brain that control both cardiac and vascular functions. In addition, more generalized neural responses from the limbic system and the autonomic nervous system are factors.

The Cardiovascular Centers in the Brain

Neurological regulation of blood pressure and flow depends on the cardiovascular centers located in the medulla oblongata. This cluster of neurons responds to changes in blood pressure as well as blood concentrations of oxygen, carbon dioxide, and hydrogen ions. The cardiovascular center contains three distinct paired components:

  • The cardioaccelerator centers stimulate cardiac function by regulating heart rate and stroke volume via sympathetic stimulation from the cardiac accelerator nerve.
  • The cardioinhibitor centers slow cardiac function by decreasing heart rate and stroke volume via parasympathetic stimulation from the vagus nerve.
  • The vasomotor centers control vessel tone or contraction of the smooth muscle in the tunica media. Changes in diameter affect peripheral resistance, pressure, and flow, which affect cardiac output. The majority of these neurons act via the release of the neurotransmitter norepinephrine from sympathetic neurons.

Although each center functions independently, they are not anatomically distinct.

There is also a small population of neurons that control vasodilation in the vessels of the brain and skeletal muscles by relaxing the smooth muscle fibers in the vessel tunics. Many of these are cholinergic neurons, that is, they release acetylcholine, which in turn stimulates the vessels’ endothelial cells to release nitric oxide (NO), which causes vasodilation. Others release norepinephrine that binds to β2 receptors. A few neurons release NO directly as a neurotransmitter.

Recall that mild stimulation of the skeletal muscles maintains muscle tone. A similar phenomenon occurs with vascular tone in vessels. As noted earlier, arterioles are normally partially constricted: With maximal stimulation, their radius may be reduced to one-half of the resting state. Full dilation of most arterioles requires that this sympathetic stimulation be suppressed. When it is, an arteriole can expand by as much as 150 percent. Such a significant increase can dramatically affect resistance, pressure, and flow.

Baroreceptor Reflexes

Baroreceptors are specialized stretch receptors located within thin areas of blood vessels and heart chambers that respond to the degree of stretch caused by the presence of blood. They send impulses to the cardiovascular center to regulate blood pressure. Vascular baroreceptors are found primarily in sinuses (small cavities) within the aorta and carotid arteries: The aortic sinuses are found in the walls of the ascending aorta just superior to the aortic valve, whereas the carotid sinuses are in the base of the internal carotid arteries. There are also low-pressure baroreceptors located in the walls of the venae cavae and right atrium. When blood pressure increases, the baroreceptors are stretched more tightly and initiate action potentials at a higher rate. At lower blood pressures, the degree of stretch is lower and the rate of firing is slower. When the cardiovascular center in the medulla oblongata receives this input, it triggers a reflex that maintains homeostasis (Figure 2):
  • When blood pressure rises too high, the baroreceptors fire at a higher rate and trigger parasympathetic stimulation of the heart. As a result, cardiac output falls. Sympathetic stimulation of the peripheral arterioles will also decrease, resulting in vasodilation. Combined, these activities cause blood pressure to fall.
  • When blood pressure drops too low, the rate of baroreceptor firing decreases. This will trigger an increase in sympathetic stimulation of the heart, causing cardiac output to increase. It will also trigger sympathetic stimulation of the peripheral vessels, resulting in vasoconstriction. Combined, these activities cause blood pressure to rise.
[caption id="" align="aligncenter" width="530"]This flow chart shows what happens when blood pressure is increased or decreased. The top panel shows the events that take place when blood pressure is increased, and the bottom panel shows the events that take place when blood pressure is decreased. Figure 2. Baroreceptor Reflexes for Maintaining Vascular Homeostasis. Increased blood pressure results in increased rates of baroreceptor firing, whereas decreased blood pressure results in slower rates of fire, both initiating the homeostatic mechanism to restore blood pressure.[/caption]

The baroreceptors in the venae cavae and right atrium monitor blood pressure as the blood returns to the heart from the systemic circulation. Normally, blood flow into the aorta is the same as blood flow back into the right atrium. If blood is returning to the right atrium more rapidly than it is being ejected from the left ventricle, the atrial receptors will stimulate the cardiovascular centers to increase sympathetic firing and increase cardiac output until homeostasis is achieved. The opposite is also true. This mechanism is referred to as the atrial reflex.

Chemoreceptor Reflexes

In addition to the baroreceptors are chemoreceptors that monitor levels of oxygen, carbon dioxide, and hydrogen ions (pH), and thereby contribute to vascular homeostasis. Chemoreceptors monitoring the blood are located in close proximity to the baroreceptors in the aortic and carotid sinuses. They signal the cardiovascular center as well as the respiratory centers in the medulla oblongata.

Since tissues consume oxygen and produce carbon dioxide and acids as waste products, when the body is more active, oxygen levels fall and carbon dioxide levels rise as cells undergo cellular respiration to meet the energy needs of activities. This causes more hydrogen ions to be produced, causing the blood pH to drop. When the body is resting, oxygen levels are higher, carbon dioxide levels are lower, more hydrogen is bound, and pH rises. (Seek additional content for more detail about pH.)

The chemoreceptors respond to increasing carbon dioxide and hydrogen ion levels (falling pH) by stimulating the cardioaccelerator and vasomotor centers, increasing cardiac output and constricting peripheral vessels. The cardioinhibitor centers are suppressed. With falling carbon dioxide and hydrogen ion levels (increasing pH), the cardioinhibitor centers are stimulated, and the cardioaccelerator and vasomotor centers are suppressed, decreasing cardiac output and causing peripheral vasodilation. In order to maintain adequate supplies of oxygen to the cells and remove waste products such as carbon dioxide, it is essential that the respiratory system respond to changing metabolic demands. In turn, the cardiovascular system will transport these gases to the lungs for exchange, again in accordance with metabolic demands. This interrelationship of cardiovascular and respiratory control cannot be overemphasized.

Other neural mechanisms can also have a significant impact on cardiovascular function. These include the limbic system that links physiological responses to psychological stimuli, as well as generalized sympathetic and parasympathetic stimulation.

Endocrine Regulation

Endocrine control over the cardiovascular system involves the catecholamines, epinephrine and norepinephrine, as well as several hormones that interact with the kidneys in the regulation of blood volume.

Epinephrine and Norepinephrine

The catecholamines epinephrine and norepinephrine are released by the adrenal medulla, and enhance and extend the body’s sympathetic or “fight-or-flight” response (see Figure 1). They increase heart rate and force of contraction, while temporarily constricting blood vessels to organs not essential for flight-or-fight responses and redirecting blood flow to the liver, muscles, and heart.

Antidiuretic Hormone

Antidiuretic hormone (ADH), also known as vasopressin, is secreted by the cells in the hypothalamus and transported via the hypothalamic-hypophyseal tracts to the posterior pituitary where it is stored until released upon nervous stimulation. The primary trigger prompting the hypothalamus to release ADH is increasing osmolarity of tissue fluid, usually in response to significant loss of blood volume. ADH signals its target cells in the kidneys to reabsorb more water, thus preventing the loss of additional fluid in the urine. This will increase overall fluid levels and help restore blood volume and pressure. In addition, ADH constricts peripheral vessels.

Renin-Angiotensin-Aldosterone Mechanism

The renin-angiotensin-aldosterone mechanism has a major effect upon the cardiovascular system (Figure 3). Renin is an enzyme, although because of its importance in the renin-angiotensin-aldosterone pathway, some sources identify it as a hormone. Specialized cells in the kidneys found in the juxtaglomerular apparatus respond to decreased blood flow by secreting renin into the blood. Renin converts the plasma protein angiotensinogen, which is produced by the liver, into its active form—angiotensin I. Angiotensin I circulates in the blood and is then converted into angiotensin II in the lungs. This reaction is catalyzed by the enzyme angiotensin-converting enzyme (ACE).

Angiotensin II is a powerful vasoconstrictor, greatly increasing blood pressure. It also stimulates the release of ADH and aldosterone, a hormone produced by the adrenal cortex. Aldosterone increases the reabsorption of sodium into the blood by the kidneys. Since water follows sodium, this increases the reabsorption of water. This in turn increases blood volume, raising blood pressure. Angiotensin II also stimulates the thirst center in the hypothalamus, so an individual will likely consume more fluids, again increasing blood volume and pressure.
[caption id="" align="aligncenter" width="560"]This flow chart shows the action of decreased blood pressure in the short and long term. Figure 3. Hormones Involved in Renal Control of Blood Pressure. In the renin-angiotensin-aldosterone mechanism, increasing angiotensin II will stimulate the production of antidiuretic hormone and aldosterone. In addition to renin, the kidneys produce erythropoietin, which stimulates the production of red blood cells, further increasing blood volume.[/caption]

Erythropoietin

Erythropoietin (EPO) is released by the kidneys when blood flow and/or oxygen levels decrease. EPO stimulates the production of erythrocytes within the bone marrow. Erythrocytes are the major formed element of the blood and may contribute 40 percent or more to blood volume, a significant factor of viscosity, resistance, pressure, and flow. In addition, EPO is a vasoconstrictor. Overproduction of EPO or excessive intake of synthetic EPO, often to enhance athletic performance, will increase viscosity, resistance, and pressure, and decrease flow in addition to its contribution as a vasoconstrictor.

Atrial Natriuretic Hormone

Secreted by cells in the atria of the heart, atrial natriuretic hormone (ANH) (also known as atrial natriuretic peptide) is secreted when blood volume is high enough to cause extreme stretching of the cardiac cells. Cells in the ventricle produce a hormone with similar effects, called B-type natriuretic hormone. Natriuretic hormones are antagonists to angiotensin II. They promote loss of sodium and water from the kidneys, and suppress renin, aldosterone, and ADH production and release. All of these actions promote loss of fluid from the body, so blood volume and blood pressure drop.

Autoregulation of Perfusion

As the name would suggest, autoregulation mechanisms require neither specialized nervous stimulation nor endocrine control. Rather, these are local, self-regulatory mechanisms that allow each region of tissue to adjust its blood flow—and thus its perfusion. These local mechanisms include chemical signals and myogenic controls.

Chemical Signals Involved in Autoregulation

Chemical signals work at the level of the precapillary sphincters to trigger either constriction or relaxation. As you know, opening a precapillary sphincter allows blood to flow into that particular capillary, whereas constricting a precapillary sphincter temporarily shuts off blood flow to that region. The factors involved in regulating the precapillary sphincters include the following:

  • Opening of the sphincter is triggered in response to decreased oxygen concentrations; increased carbon dioxide concentrations; increasing levels of lactic acid or other byproducts of cellular metabolism; increasing concentrations of potassium ions or hydrogen ions (falling pH); inflammatory chemicals such as histamines; and increased body temperature. These conditions in turn stimulate the release of NO, a powerful vasodilator, from endothelial cells (see Figure 1).
  • Contraction of the precapillary sphincter is triggered by the opposite levels of the regulators, which prompt the release of endothelins, powerful vasoconstricting peptides secreted by endothelial cells. Platelet secretions and certain prostaglandins may also trigger constriction.

Again, these factors alter tissue perfusion via their effects on the precapillary sphincter mechanism, which regulates blood flow to capillaries. Since the amount of blood is limited, not all capillaries can fill at once, so blood flow is allocated based upon the needs and metabolic state of the tissues as reflected in these parameters. Bear in mind, however, that dilation and constriction of the arterioles feeding the capillary beds is the primary control mechanism.

The Myogenic Response

The myogenic response is a reaction to the stretching of the smooth muscle in the walls of arterioles as changes in blood flow occur through the vessel. This may be viewed as a largely protective function against dramatic fluctuations in blood pressure and blood flow to maintain homeostasis. If perfusion of an organ is too low (ischemia), the tissue will experience low levels of oxygen (hypoxia). In contrast, excessive perfusion could damage the organ’s smaller and more fragile vessels. The myogenic response is a localized process that serves to stabilize blood flow in the capillary network that follows that arteriole.

When blood flow is low, the vessel’s smooth muscle will be only minimally stretched. In response, it relaxes, allowing the vessel to dilate and thereby increase the movement of blood into the tissue. When blood flow is too high, the smooth muscle will contract in response to the increased stretch, prompting vasoconstriction that reduces blood flow.

Figure 4 summarizes the effects of nervous, endocrine, and local controls on arterioles.

[caption id="" align="aligncenter" width="520"]This table summarizes mechanisms that regulate arteriole smooth muscle and veins. Neural controls are regulated by sympathetic stimulation and parasympathetic. Endocrine controls are regulated by epinephrine, norepinephrine, angiotensin II, ANH (peptide), and ADH. Other factors include decreasing levels of oxygen, decreasing pH, increasing levels of carbon dioxide, increasing levels of potassium ion, increasing levels of prostaglandins, increasing levels of andenosine, increasing levels of NO, increasing levels of lactic acid and other metabolites, increasing levels of endothelins, increasing levels of platelet secretions, increasing hyperhtermia, stretching of vascular wall (myogenic), and increasing levels of histamines from basophils and mast cells. Figure 4. Summary of Mechanisms Regulating Arteriole Smooth Muscle and Veins.[/caption]

Effect of Exercise on Vascular Homeostasis

The heart is a muscle and, like any muscle, it responds dramatically to exercise. For a healthy young adult, cardiac output (heart rate × stroke volume) increases in the nonathlete from approximately 5.0 liters (5.25 quarts) per minute to a maximum of about 20 liters (21 quarts) per minute. Accompanying this will be an increase in blood pressure from about 120/80 to 185/75. However, well-trained aerobic athletes can increase these values substantially. For these individuals, cardiac output soars from approximately 5.3 liters (5.57 quarts) per minute resting to more than 30 liters (31.5 quarts) per minute during maximal exercise. Along with this increase in cardiac output, blood pressure increases from 120/80 at rest to 200/90 at maximum values. In addition to improved cardiac function, exercise increases the size and mass of the heart. The average weight of the heart for the nonathlete is about 300 g, whereas in an athlete it will increase to 500 g. This increase in size generally makes the heart stronger and more efficient at pumping blood, increasing both stroke volume and cardiac output.

Tissue perfusion also increases as the body transitions from a resting state to light exercise and eventually to heavy exercise (see Figure 4). These changes result in selective vasodilation in the skeletal muscles, heart, lungs, liver, and integument. Simultaneously, vasoconstriction occurs in the vessels leading to the kidneys and most of the digestive and reproductive organs. The flow of blood to the brain remains largely unchanged whether at rest or exercising, since the vessels in the brain largely do not respond to regulatory stimuli, in most cases, because they lack the appropriate receptors.

As vasodilation occurs in selected vessels, resistance drops and more blood rushes into the organs they supply. This blood eventually returns to the venous system. Venous return is further enhanced by both the skeletal muscle and respiratory pumps. As blood returns to the heart more quickly, preload rises and the Frank-Starling principle tells us that contraction of the cardiac muscle in the atria and ventricles will be more forceful. Eventually, even the best-trained athletes will fatigue and must undergo a period of rest following exercise. Cardiac output and distribution of blood then return to normal.

Regular exercise promotes cardiovascular health in a variety of ways. Because an athlete’s heart is larger than a nonathlete’s, stroke volume increases, so the athletic heart can deliver the same amount of blood as the nonathletic heart but with a lower heart rate. This increased efficiency allows the athlete to exercise for longer periods of time before muscles fatigue and places less stress on the heart. Exercise also lowers overall cholesterol levels by removing from the circulation a complex form of cholesterol, triglycerides, and proteins known as low-density lipoproteins (LDLs), which are widely associated with increased risk of cardiovascular disease. Although there is no way to remove deposits of plaque from the walls of arteries other than specialized surgery, exercise does promote the health of vessels by decreasing the rate of plaque formation and reducing blood pressure, so the heart does not have to generate as much force to overcome resistance.

Generally as little as 30 minutes of noncontinuous exercise over the course of each day has beneficial effects and has been shown to lower the rate of heart attack by nearly 50 percent. While it is always advisable to follow a healthy diet, stop smoking, and lose weight, studies have clearly shown that fit, overweight people may actually be healthier overall than sedentary slender people. Thus, the benefits of moderate exercise are undeniable.

Clinical Considerations in Vascular Homeostasis

Any disorder that affects blood volume, vascular tone, or any other aspect of vascular functioning is likely to affect vascular homeostasis as well. That includes hypertension, hemorrhage, and shock.

Hypertension and Hypotension

Chronically elevated blood pressure is known clinically as hypertension. It is defined as chronic and persistent blood pressure measurements of 140/90 mm Hg or above. Pressures between 120/80 and 140/90 mm Hg are defined as prehypertension. About 68 million Americans currently suffer from hypertension. Unfortunately, hypertension is typically a silent disorder; therefore, hypertensive patients may fail to recognize the seriousness of their condition and fail to follow their treatment plan. The result is often a heart attack or stroke. Hypertension may also lead to an aneurism (ballooning of a blood vessel caused by a weakening of the wall), peripheral arterial disease (obstruction of vessels in peripheral regions of the body), chronic kidney disease, or heart failure. [caption id="" align="aligncenter" width="120"]QR Code representing a URL Listen to this CDC podcast to learn about hypertension, often described as a “silent killer.”[/caption]

Hemorrhage

Minor blood loss is managed by hemostasis and repair. Hemorrhage is a loss of blood that cannot be controlled by hemostatic mechanisms. Initially, the body responds to hemorrhage by initiating mechanisms aimed at increasing blood pressure and maintaining blood flow. Ultimately, however, blood volume will need to be restored, either through physiological processes or through medical intervention.

In response to blood loss, stimuli from the baroreceptors trigger the cardiovascular centers to stimulate sympathetic responses to increase cardiac output and vasoconstriction. This typically prompts the heart rate to increase to about 180–200 contractions per minute, restoring cardiac output to normal levels. Vasoconstriction of the arterioles increases vascular resistance, whereas constriction of the veins increases venous return to the heart. Both of these steps will help increase blood pressure. Sympathetic stimulation also triggers the release of epinephrine and norepinephrine, which enhance both cardiac output and vasoconstriction. If blood loss were less than 20 percent of total blood volume, these responses together would usually return blood pressure to normal and redirect the remaining blood to the tissues.

Additional endocrine involvement is necessary, however, to restore the lost blood volume. The angiotensin-renin-aldosterone mechanism stimulates the thirst center in the hypothalamus, which increases fluid consumption to help restore the lost blood. More importantly, it increases renal reabsorption of sodium and water, reducing water loss in urine output. The kidneys also increase the production of EPO, stimulating the formation of erythrocytes that not only deliver oxygen to the tissues but also increase overall blood volume. Figure 5 summarizes the responses to loss of blood volume.

[caption id="" align="aligncenter" width="490"]This flowchart shows the action of decreased blood pressure and volume in the neural and endocrine mechanisms. Figure 5. Homeostatic Responses to Loss of Blood Volume[/caption]
]]> 2286 2261 5 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2312 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2312 [caption id="" align="aligncenter" width="600"]The top panel shows a color-coded world map. The bottom panel shows many viruses on a cell. Figure 1. The Worldwide AIDS Epidemic. (a) As of 2008, more than 15 percent of adults were infected with HIV in certain African countries. This grim picture had changed little by 2012. (b) In this scanning electron micrograph, HIV virions (green particles) are budding off the surface of a macrophage (pink structure). (credit b: C. Goldsmith)[/caption]

Chapter Objectives

After studying this chapter, you will be able to:
  • Identify the components and anatomy of the lymphatic system
  • Discuss the role of the innate immune response against pathogens
  • Describe the power of the adaptive immune response to cure disease
  • Explain immunological deficiencies and over-reactions of the immune system
  • Discuss the role of the immune response in transplantation and cancer
  • Describe the interaction of the immune and lymphatic systems with other body systems
In June 1981, the Centers for Disease Control and Prevention (CDC), in Atlanta, Georgia, published a report of an unusual cluster of five patients in Los Angeles, California. All five were diagnosed with a rare pneumonia caused by a fungus called Pneumocystis jirovecii (formerly known as Pneumocystis carinii).

Why was this unusual? Although commonly found in the lungs of healthy individuals, this fungus is an opportunistic pathogen that causes disease in individuals with suppressed or underdeveloped immune systems. The very young, whose immune systems have yet to mature, and the elderly, whose immune systems have declined with age, are particularly susceptible. The five patients from LA, though, were between 29 and 36 years of age and should have been in the prime of their lives, immunologically speaking. What could be going on?

A few days later, a cluster of eight cases was reported in New York City, also involving young patients, this time exhibiting a rare form of skin cancer known as Kaposi’s sarcoma. This cancer of the cells that line the blood and lymphatic vessels was previously observed as a relatively innocuous disease of the elderly. The disease that doctors saw in 1981 was frighteningly more severe, with multiple, fast-growing lesions that spread to all parts of the body, including the trunk and face. Could the immune systems of these young patients have been compromised in some way? Indeed, when they were tested, they exhibited extremely low numbers of a specific type of white blood cell in their bloodstreams, indicating that they had somehow lost a major part of the immune system. Acquired immune deficiency syndrome, or AIDS, turned out to be a new disease caused by the previously unknown human immunodeficiency virus (HIV). Although nearly 100 percent fatal in those with active HIV infections in the early years, the development of anti-HIV drugs has transformed HIV infection into a chronic, manageable disease and not the certain death sentence it once was. One positive outcome resulting from the emergence of HIV disease was that the public’s attention became focused as never before on the importance of having a functional and healthy immune system.]]> 2312 2310 1 0 21.3 The Adaptive Immune Response: T lymphocytes and Their Functional Types https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2336 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2336

Learning Objectives

By the end of this section, you will be able to:
  • Explain the advantages of the adaptive immune response over the innate immune response
  • List the various characteristics of an antigen
  • Describe the types of T cell antigen receptors
  • Outline the steps of T cell development
  • Describe the major T cell types and their functions

Innate immune responses (and early induced responses) are in many cases ineffective at completely controlling pathogen growth. However, they slow pathogen growth and allow time for the adaptive immune response to strengthen and either control or eliminate the pathogen. The innate immune system also sends signals to the cells of the adaptive immune system, guiding them in how to attack the pathogen. Thus, these are the two important arms of the immune response.

The Benefits of the Adaptive Immune Response

The specificity of the adaptive immune response—its ability to specifically recognize and make a response against a wide variety of pathogens—is its great strength. Antigens, the small chemical groups often associated with pathogens, are recognized by receptors on the surface of B and T lymphocytes. The adaptive immune response to these antigens is so versatile that it can respond to nearly any pathogen. This increase in specificity comes because the adaptive immune response has a unique way to develop as many as 1011, or 100 trillion, different receptors to recognize nearly every conceivable pathogen. How could so many different types of antibodies be encoded? And what about the many specificities of T cells? There is not nearly enough DNA in a cell to have a separate gene for each specificity. The mechanism was finally worked out in the 1970s and 1980s using the new tools of molecular genetics

Primary Disease and Immunological Memory

The immune system’s first exposure to a pathogen is called a primary adaptive response. Symptoms of a first infection, called primary disease, are always relatively severe because it takes time for an initial adaptive immune response to a pathogen to become effective.

Upon re-exposure to the same pathogen, a secondary adaptive immune response is generated, which is stronger and faster that the primary response. The secondary adaptive response often eliminates a pathogen before it can cause significant tissue damage or any symptoms. Without symptoms, there is no disease, and the individual is not even aware of the infection. This secondary response is the basis of immunological memory, which protects us from getting diseases repeatedly from the same pathogen. By this mechanism, an individual’s exposure to pathogens early in life spares the person from these diseases later in life.

Self Recognition

A third important feature of the adaptive immune response is its ability to distinguish between self-antigens, those that are normally present in the body, and foreign antigens, those that might be on a potential pathogen. As T and B cells mature, there are mechanisms in place that prevent them from recognizing self-antigen, preventing a damaging immune response against the body. These mechanisms are not 100 percent effective, however, and their breakdown leads to autoimmune diseases, which will be discussed later in this chapter.

T Cell-Mediated Immune Responses

The primary cells that control the adaptive immune response are the lymphocytes, the T and B cells. T cells are particularly important, as they not only control a multitude of immune responses directly, but also control B cell immune responses in many cases as well. Thus, many of the decisions about how to attack a pathogen are made at the T cell level, and knowledge of their functional types is crucial to understanding the functioning and regulation of adaptive immune responses as a whole.

T lymphocytes recognize antigens based on a two-chain protein receptor. The most common and important of these are the alpha-beta T cell receptors (Figure 1).

[caption id="" align="aligncenter" width="300"]This figure shows the alpha beta T cell receptor in the plasma membrane. Figure 1. Alpha-beta T Cell Receptor. Notice the constant and variable regions of each chain, anchored by the transmembrane region.[/caption]

There are two chains in the T cell receptor, and each chain consists of two domains. The variable region domain is furthest away from the T cell membrane and is so named because its amino acid sequence varies between receptors. In contrast, the constant region domain has less variation. The differences in the amino acid sequences of the variable domains are the molecular basis of the diversity of antigens the receptor can recognize. Thus, the antigen-binding site of the receptor consists of the terminal ends of both receptor chains, and the amino acid sequences of those two areas combine to determine its antigenic specificity. Each T cell produces only one type of receptor and thus is specific for a single particular antigen.

Antigens

Antigens on pathogens are usually large and complex, and consist of many antigenic determinants. An antigenic determinant (epitope) is one of the small regions within an antigen to which a receptor can bind, and antigenic determinants are limited by the size of the receptor itself. They usually consist of six or fewer amino acid residues in a protein, or one or two sugar moieties in a carbohydrate antigen. Antigenic determinants on a carbohydrate antigen are usually less diverse than on a protein antigen. Carbohydrate antigens are found on bacterial cell walls and on red blood cells (the ABO blood group antigens). Protein antigens are complex because of the variety of three-dimensional shapes that proteins can assume, and are especially important for the immune responses to viruses and worm parasites. It is the interaction of the shape of the antigen and the complementary shape of the amino acids of the antigen-binding site that accounts for the chemical basis of specificity (Figure 2).

[caption id="" align="aligncenter" width="480"]This figure shows three T cells and the antigenic determinants in the center. Figure 2. Antigenic Determinants. A typical protein antigen has multiple antigenic determinants, shown by the ability of T cells with three different specificities to bind to different parts of the same antigen.[/caption]

Antigen Processing and Presentation

Although Figure 2 shows T cell receptors interacting with antigenic determinants directly, the mechanism that T cells use to recognize antigens is, in reality, much more complex. T cells do not recognize free-floating or cell-bound antigens as they appear on the surface of the pathogen. They only recognize antigen on the surface of specialized cells called antigen-presenting cells. Antigens are internalized by these cells. Antigen processing is a mechanism that enzymatically cleaves the antigen into smaller pieces. The antigen fragments are then brought to the cell’s surface and associated with a specialized type of antigen-presenting protein known as a major histocompatibility complex (MHC) molecule. The MHC is the cluster of genes that encode these antigen-presenting molecules. The association of the antigen fragments with an MHC molecule on the surface of a cell is known as antigen presentation and results in the recognition of antigen by a T cell. This association of antigen and MHC occurs inside the cell, and it is the complex of the two that is brought to the surface. The peptide-binding cleft is a small indentation at the end of the MHC molecule that is furthest away from the cell membrane; it is here that the processed fragment of antigen sits. MHC molecules are capable of presenting a variety of antigens, depending on the amino acid sequence, in their peptide-binding clefts. It is the combination of the MHC molecule and the fragment of the original peptide or carbohydrate that is actually physically recognized by the T cell receptor (Figure 3).

[caption id="" align="aligncenter" width="560"]This figure shows how an antigen-presenting cell deals with a pathogen or extracellular antigen. The different steps are shown with text callouts. Figure 3. Antigen Processing and Presentation.[/caption]

Two distinct types of MHC molecules, MHC class I and MHC class II, play roles in antigen presentation. Although produced from different genes, they both have similar functions. They bring processed antigen to the surface of the cell via a transport vesicle and present the antigen to the T cell and its receptor. Antigens from different classes of pathogens, however, use different MHC classes and take different routes through the cell to get to the surface for presentation. The basic mechanism, though, is the same. Antigens are processed by digestion, are brought into the endomembrane system of the cell, and then are expressed on the surface of the antigen-presenting cell for antigen recognition by a T cell. Intracellular antigens are typical of viruses, which replicate inside the cell, and certain other intracellular parasites and bacteria. These antigens are processed in the cytosol by an enzyme complex known as the proteasome and are then brought into the endoplasmic reticulum by the transporter associated with antigen processing (TAP) system, where they interact with class I MHC molecules and are eventually transported to the cell surface by a transport vesicle.

Extracellular antigens, characteristic of many bacteria, parasites, and fungi that do not replicate inside the cell’s cytoplasm, are brought into the endomembrane system of the cell by receptor-mediated endocytosis. The resulting vesicle fuses with vesicles from the Golgi complex, which contain pre-formed MHC class II molecules. After fusion of these two vesicles and the association of antigen and MHC, the new vesicle makes its way to the cell surface.

Professional Antigen-presenting Cells

Many cell types express class I molecules for the presentation of intracellular antigens. These MHC molecules may then stimulate a cytotoxic T cell immune response, eventually destroying the cell and the pathogen within. This is especially important when it comes to the most common class of intracellular pathogens, the virus. Viruses infect nearly every tissue of the body, so all these tissues must necessarily be able to express class I MHC or no T cell response can be made.

On the other hand, class II MHC molecules are expressed only on the cells of the immune system, specifically cells that affect other arms of the immune response. Thus, these cells are called “professional” antigen-presenting cells to distinguish them from those that bear class I MHC. The three types of professional antigen presenters are macrophages, dendritic cells, and B cells (Table 4).

Macrophages stimulate T cells to release cytokines that enhance phagocytosis. Dendritic cells also kill pathogens by phagocytosis (see Figure 3), but their major function is to bring antigens to regional draining lymph nodes. The lymph nodes are the locations in which most T cell responses against pathogens of the interstitial tissues are mounted. Macrophages are found in the skin and in the lining of mucosal surfaces, such as the nasopharynx, stomach, lungs, and intestines. B cells may also present antigens to T cells, which are necessary for certain types of antibody responses, to be covered later in this chapter.

Classes of Antigen-presenting Cells (Table 4)
MHC Cell type Phagocytic? Function
Class I Many No Stimulates cytotoxic T cell immune response
Class II Macrophage Yes Stimulates phagocytosis and presentation at primary infection site
Class II Dendritic Yes, in tissues Brings antigens to regional lymph nodes
Class II B cell Yes, internalizes surface Ig and antigen Stimulates antibody secretion by B cells

T Cell Development and Differentiation

The process of eliminating T cells that might attack the cells of one’s own body is referred to as T cell tolerance. While thymocytes are in the cortex of the thymus, they are referred to as “double negatives,” meaning that they do not bear the CD4 or CD8 molecules that you can use to follow their pathways of differentiation (Figure 4). In the cortex of the thymus, they are exposed to cortical epithelial cells. In a process known as positive selection, double-negative thymocytes bind to the MHC molecules they observe on the thymic epithelia, and the MHC molecules of “self” are selected. This mechanism kills many thymocytes during T cell differentiation. In fact, only two percent of the thymocytes that enter the thymus leave it as mature, functional T cells.

[caption id="" align="aligncenter" width="500"]This multipart figure shows the different steps in the differentiation of a thymocyte into T cells. For each step of the process, accompanying text details the steps in the process. The right panel of this image shows the location of the different steps in the process. Figure 4. Differentiation of T Cells within the Thymus. Thymocytes enter the thymus and go through a series of developmental stages that ensures both function and tolerance before they leave and become functional components of the adaptive immune response.[/caption]

Later, the cells become double positives that express both CD4 and CD8 markers and move from the cortex to the junction between the cortex and medulla. It is here that negative selection takes place. In negative selection, self-antigens are brought into the thymus from other parts of the body by professional antigen-presenting cells. The T cells that bind to these self-antigens are selected for negatively and are killed by apoptosis. In summary, the only T cells left are those that can bind to MHC molecules of the body with foreign antigens presented on their binding clefts, preventing an attack on one’s own body tissues, at least under normal circumstances. Tolerance can be broken, however, by the development of an autoimmune response, to be discussed later in this chapter.

The cells that leave the thymus become single positives, expressing either CD4 or CD8, but not both (see Figure 4). The CD4+ T cells will bind to class II MHC and the CD8+ cells will bind to class I MHC. The discussion that follows explains the functions of these molecules and how they can be used to differentiate between the different T cell functional types.

Mechanisms of T Cell-mediated Immune Responses

Mature T cells become activated by recognizing processed foreign antigen in association with a self-MHC molecule and begin dividing rapidly by mitosis. This proliferation of T cells is called clonal expansion and is necessary to make the immune response strong enough to effectively control a pathogen. How does the body select only those T cells that are needed against a specific pathogen? Again, the specificity of a T cell is based on the amino acid sequence and the three-dimensional shape of the antigen-binding site formed by the variable regions of the two chains of the T cell receptor (Figure 5). Clonal selection is the process of antigen binding only to those T cells that have receptors specific to that antigen. Each T cell that is activated has a specific receptor “hard-wired” into its DNA, and all of its progeny will have identical DNA and T cell receptors, forming clones of the original T cell.

[caption id="" align="aligncenter" width="450"]This flowchart shows the process in which a naïve T cell become activated T cells in the left part of the pathway and memory cells in the right part of the pathway. Figure 5. Clonal Selection and Expansion of T Lymphocytes. Stem cells differentiate into T cells with specific receptors, called clones. The clones with receptors specific for antigens on the pathogen are selected for and expanded.[/caption]

Clonal Selection and Expansion

The clonal selection theory was proposed by Frank Burnet in the 1950s. However, the term clonal selection is not a complete description of the theory, as clonal expansion goes hand in glove with the selection process. The main tenet of the theory is that a typical individual has a multitude (1011) of different types of T cell clones based on their receptors. In this use, a clone is a group of lymphocytes that share the same antigen receptor. Each clone is necessarily present in the body in low numbers. Otherwise, the body would not have room for lymphocytes with so many specificities.

Only those clones of lymphocytes whose receptors are activated by the antigen are stimulated to proliferate. Keep in mind that most antigens have multiple antigenic determinants, so a T cell response to a typical antigen involves a polyclonal response. A polyclonal response is the stimulation of multiple T cell clones. Once activated, the selected clones increase in number and make many copies of each cell type, each clone with its unique receptor. By the time this process is complete, the body will have large numbers of specific lymphocytes available to fight the infection (see Figure 5).

The Cellular Basis of Immunological Memory

As already discussed, one of the major features of an adaptive immune response is the development of immunological memory.

During a primary adaptive immune response, both memory T cells and effector T cells are generated. Memory T cells are long-lived and can even persist for a lifetime. Memory cells are primed to act rapidly. Thus, any subsequent exposure to the pathogen will elicit a very rapid T cell response. This rapid, secondary adaptive response generates large numbers of effector T cells so fast that the pathogen is often overwhelmed before it can cause any symptoms of disease. This is what is meant by immunity to a disease. The same pattern of primary and secondary immune responses occurs in B cells and the antibody response, as will be discussed later in the chapter.

T Cell Types and their Functions

In the discussion of T cell development, you saw that mature T cells express either the CD4 marker or the CD8 marker, but not both. These markers are cell adhesion molecules that keep the T cell in close contact with the antigen-presenting cell by directly binding to the MHC molecule (to a different part of the molecule than does the antigen). Thus, T cells and antigen-presenting cells are held together in two ways: by CD4 or CD8 attaching to MHC and by the T cell receptor binding to antigen (Figure 6).

[caption id="" align="aligncenter" width="380"]This figure shows the different steps in processing an extracellular pathogen. Figure 6. Pathogen Presentation. (a) CD4 is associated with helper and regulatory T cells. An extracellular pathogen is processed and presented in the binding cleft of a class II MHC molecule, and this interaction is strengthened by the CD4 molecule. (b) CD8 is associated with cytotoxic T cells. An intracellular pathogen is presented by a class I MHC molecule, and CD8 interacts with it.[/caption]

Although the correlation is not 100 percent, CD4-bearing T cells are associated with helper functions and CD8-bearing T cells are associated with cytotoxicity. These functional distinctions based on CD4 and CD8 markers are useful in defining the function of each type.

Helper T Cells and their Cytokines

Helper T cells (Th), bearing the CD4 molecule, function by secreting cytokines that act to enhance other immune responses. There are two classes of Th cells, and they act on different components of the immune response. These cells are not distinguished by their surface molecules but by the characteristic set of cytokines they secrete (Table 5).

Th1 cells are a type of helper T cell that secretes cytokines that regulate the immunological activity and development of a variety of cells, including macrophages and other types of T cells.

Th2 cells, on the other hand, are cytokine-secreting cells that act on B cells to drive their differentiation into plasma cells that make antibody. In fact, T cell help is required for antibody responses to most protein antigens, and these are called T cell-dependent antigens.

Cytotoxic T cells

Cytotoxic T cells (Tc) are T cells that kill target cells by inducing apoptosis using the same mechanism as NK cells. They either express Fas ligand, which binds to the fas molecule on the target cell, or act by using perforins and granzymes contained in their cytoplasmic granules. As was discussed earlier with NK cells, killing a virally infected cell before the virus can complete its replication cycle results in the production of no infectious particles. As more Tc cells are developed during an immune response, they overwhelm the ability of the virus to cause disease. In addition, each Tc cell can kill more than one target cell, making them especially effective. Tc cells are so important in the antiviral immune response that some speculate that this was the main reason the adaptive immune response evolved in the first place.

Regulatory T Cells

Regulatory T cells (Treg), or suppressor T cells, are the most recently discovered of the types listed here, so less is understood about them. In addition to CD4, they bear the molecules CD25 and FOXP3. Exactly how they function is still under investigation, but it is known that they suppress other T cell immune responses. This is an important feature of the immune response, because if clonal expansion during immune responses were allowed to continue uncontrolled, these responses could lead to autoimmune diseases and other medical issues.

Not only do T cells directly destroy pathogens, but they regulate nearly all other types of the adaptive immune response as well, as evidenced by the functions of the T cell types, their surface markers, the cells they work on, and the types of pathogens they work against (see Table 5).
Functions of T Cell Types and Their Cytokines (Table 5)
T cell Main target Function Pathogen Surface marker MHC Cytokines or mediators
Tc Infected cells Cytotoxicity Intracellular CD8 Class I Perforins, granzymes, and fas ligand
Th1 Macrophage Helper inducer Extracellular CD4 Class II Interferon-γ and TGF-β
Th2 B cell Helper inducer Extracellular CD4 Class II IL-4, IL-6, IL-10, and others
Treg Th cell Suppressor None CD4, CD25 ? TGF-β and IL-10
]]> 2336 2310 4 0 21.5 The Immune Response against Pathogens https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2345 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2345

Learning Objectives

By the end of this section, you will be able to:
  • Explain the development of immunological competence
  • Describe the mucosal immune response
  • Discuss immune responses against bacterial, viral, fungal, and animal pathogens
  • Describe different ways pathogens evade immune responses

Now that you understand the development of mature, naïve B cells and T cells, and some of their major functions, how do all of these various cells, proteins, and cytokines come together to actually resolve an infection? Ideally, the immune response will rid the body of a pathogen entirely. The adaptive immune response, with its rapid clonal expansion, is well suited to this purpose. Think of a primary infection as a race between the pathogen and the immune system. The pathogen bypasses barrier defenses and starts multiplying in the host’s body. During the first 4 to 5 days, the innate immune response will partially control, but not stop, pathogen growth. As the adaptive immune response gears up, however, it will begin to clear the pathogen from the body, while at the same time becoming stronger and stronger. When following antibody responses in patients with a particular disease such as a virus, this clearance is referred to as seroconversion (sero- = “serum”). Seroconversion is the reciprocal relationship between virus levels in the blood and antibody levels. As the antibody levels rise, the virus levels decline, and this is a sign that the immune response is being at least partially effective (partially, because in many diseases, seroconversion does not necessarily mean a patient is getting well).

An excellent example of this is seroconversion during HIV disease (Figure 1). Notice that antibodies are made early in this disease, and the increase in anti-HIV antibodies correlates with a decrease in detectable virus in the blood. Although these antibodies are an important marker for diagnosing the disease, they are not sufficient to completely clear the virus. Several years later, the vast majority of these individuals, if untreated, will lose their entire adaptive immune response, including the ability to make antibodies, during the final stages of AIDS.

[caption id="" align="aligncenter" width="480"]This graph shows the concentration of HIV viral particles in blood over time in years. Figure 1. HIV Disease Progression. Seroconversion, the rise of anti-HIV antibody levels and the concomitant decline in measurable virus levels, happens during the first several months of HIV disease. Unfortunately, this antibody response is ineffective at controlling the disease, as seen by the progression of the disease towards AIDS, in which all adaptive immune responses are compromised.[/caption]

The Mucosal Immune Response

Mucosal tissues are major barriers to the entry of pathogens into the body. The IgA (and sometimes IgM) antibodies in mucus and other secretions can bind to the pathogen, and in the cases of many viruses and bacteria, neutralize them. Neutralization is the process of coating a pathogen with antibodies, making it physically impossible for the pathogen to bind to receptors. Neutralization, which occurs in the blood, lymph, and other body fluids and secretions, protects the body constantly. Neutralizing antibodies are the basis for the disease protection offered by vaccines. Vaccinations for diseases that commonly enter the body via mucous membranes, such as influenza, are usually formulated to enhance IgA production.

Immune responses in some mucosal tissues such as the Peyer’s patches (see Chapter 21.1 Figure 10) in the small intestine take up particulate antigens by specialized cells known as microfold or M cells (Figure 2). These cells allow the body to sample potential pathogens from the intestinal lumen. Dendritic cells then take the antigen to the regional lymph nodes, where an immune response is mounted.

[caption id="" align="aligncenter" width="480"]This diagram shows the process in which cells of the small intestine generate IgA immunity. Figure 2. IgA Immunity. The nasal-associated lymphoid tissue and Peyer’s patches of the small intestine generate IgA immunity. Both use M cells to transport antigen inside the body so that immune responses can be mounted.[/caption]

Defenses against Bacteria and Fungi

The body fights bacterial pathogens with a wide variety of immunological mechanisms, essentially trying to find one that is effective. Bacteria such as Mycobacterium leprae, the cause of leprosy, are resistant to lysosomal enzymes and can persist in macrophage organelles or escape into the cytosol. In such situations, infected macrophages receiving cytokine signals from Th1 cells turn on special metabolic pathways. Macrophage oxidative metabolism is hostile to intracellular bacteria, often relying on the production of nitric oxide to kill the bacteria inside the macrophage.

Fungal infections, such as those from Aspergillus, Candida, and Pneumocystis, are largely opportunistic infections that take advantage of suppressed immune responses. Most of the same immune mechanisms effective against bacteria have similar effects on fungi, both of which have characteristic cell wall structures that protect their cells.

Defenses against Parasites

Worm parasites such as helminths are seen as the primary reason why the mucosal immune response, IgE-mediated allergy and asthma, and eosinophils evolved. These parasites were at one time very common in human society. When infecting a human, often via contaminated food, some worms take up residence in the gastrointestinal tract. Eosinophils are attracted to the site by T cell cytokines, which release their granule contents upon their arrival. Mast cell degranulation also occurs, and the fluid leakage caused by the increase in local vascular permeability is thought to have a flushing action on the parasite, expelling its larvae from the body. Furthermore, if IgE labels the parasite, the eosinophils can bind to it by its Fc receptor.

Defenses against Viruses

The primary mechanisms against viruses are NK cells, interferons, and cytotoxic T cells. Antibodies are effective against viruses mostly during protection, where an immune individual can neutralize them based on a previous exposure. Antibodies have no effect on viruses or other intracellular pathogens once they enter the cell, since antibodies are not able to penetrate the plasma membrane of the cell. Many cells respond to viral infections by downregulating their expression of MHC class I molecules. This is to the advantage of the virus, because without class I expression, cytotoxic T cells have no activity. NK cells, however, can recognize virally infected class I-negative cells and destroy them. Thus, NK and cytotoxic T cells have complementary activities against virally infected cells.

Interferons have activity in slowing viral replication and are used in the treatment of certain viral diseases, such as hepatitis B and C, but their ability to eliminate the virus completely is limited. The cytotoxic T cell response, though, is key, as it eventually overwhelms the virus and kills infected cells before the virus can complete its replicative cycle. Clonal expansion and the ability of cytotoxic T cells to kill more than one target cell make these cells especially effective against viruses. In fact, without cytotoxic T cells, it is likely that humans would all die at some point from a viral infection (if no vaccine were available).

Evasion of the Immune System by Pathogens

It is important to keep in mind that although the immune system has evolved to be able to control many pathogens, pathogens themselves have evolved ways to evade the immune response. An example already mentioned is in Mycobactrium tuberculosis, which has evolved a complex cell wall that is resistant to the digestive enzymes of the macrophages that ingest them, and thus persists in the host, causing the chronic disease tuberculosis. This section briefly summarizes other ways in which pathogens can “outwit” immune responses. But keep in mind, although it seems as if pathogens have a will of their own, they do not. All of these evasive “strategies” arose strictly by evolution, driven by selection.

Bacteria sometimes evade immune responses because they exist in multiple strains, such as different groups of Staphylococcus aureus. S. aureus is commonly found in minor skin infections, such as boils, and some healthy people harbor it in their nose. One small group of strains of this bacterium, however, called methicillin-resistant Staphylococcus aureus, has become resistant to multiple antibiotics and is essentially untreatable. Different bacterial strains differ in the antigens on their surfaces. The immune response against one strain (antigen) does not affect the other; thus, the species survives.

Another method of immune evasion is mutation. Because viruses’ surface molecules mutate continuously, viruses like influenza change enough each year that the flu vaccine for one year may not protect against the flu common to the next. New vaccine formulations must be derived for each flu season.

Genetic recombination—the combining of gene segments from two different pathogens—is an efficient form of immune evasion. For example, the influenza virus contains gene segments that can recombine when two different viruses infect the same cell. Recombination between human and pig influenza viruses led to the 2010 H1N1 swine flu outbreak.

Pathogens can produce immunosuppressive molecules that impair immune function, and there are several different types. Viruses are especially good at evading the immune response in this way, and many types of viruses have been shown to suppress the host immune response in ways much more subtle than the wholesale destruction caused by HIV.

]]> 2345 2310 6 0 21.6 Diseases Associated with Depressed or Overactive Immune Responses https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2348 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2348

Learning Objectives

By the end of this section, you will be able to:
  • Discuss inherited and acquired immunodeficiencies
  • Explain the four types of hypersensitivity and how they differ
  • Give an example of how autoimmune disease breaks tolerance

This section is about how the immune system goes wrong. When it goes haywire, and becomes too weak or too strong, it leads to a state of disease. The factors that maintain immunological homeostasis are complex and incompletely understood.

Immunodeficiencies

As you have seen, the immune system is quite complex. It has many pathways using many cell types and signals. Because it is so complex, there are many ways for it to go wrong. Inherited immunodeficiencies arise from gene mutations that affect specific components of the immune response. There are also acquired immunodeficiencies with potentially devastating effects on the immune system, such as HIV.

Inherited Immunodeficiencies

A list of all inherited immunodeficiencies is well beyond the scope of this book. The list is almost as long as the list of cells, proteins, and signaling molecules of the immune system itself. Some deficiencies, such as those for complement, cause only a higher susceptibility to some Gram-negative bacteria. Others are more severe in their consequences. Certainly, the most serious of the inherited immunodeficiencies is severe combined immunodeficiency disease (SCID). This disease is complex because it is caused by many different genetic defects. What groups them together is the fact that both the B cell and T cell arms of the adaptive immune response are affected.

Children with this disease usually die of opportunistic infections within their first year of life unless they receive a bone marrow transplant. Such a procedure had not yet been perfected for David Vetter, the “boy in the bubble,” who was treated for SCID by having to live almost his entire life in a sterile plastic cocoon for the 12 years before his death from infection in 1984. One of the features that make bone marrow transplants work as well as they do is the proliferative capability of hematopoietic stem cells of the bone marrow. Only a small amount of bone marrow from a healthy donor is given intravenously to the recipient. It finds its own way to the bone where it populates it, eventually reconstituting the patient’s immune system, which is usually destroyed beforehand by treatment with radiation or chemotherapeutic drugs.

New treatments for SCID using gene therapy, inserting nondefective genes into cells taken from the patient and giving them back, have the advantage of not needing the tissue match required for standard transplants. Although not a standard treatment, this approach holds promise, especially for those in whom standard bone marrow transplantation has failed.

Human Immunodeficiency Virus/AIDS

Although many viruses cause suppression of the immune system, only one wipes it out completely, and that is the previously mentioned HIV. It is worth discussing the biology of this virus, which can lead to the well-known AIDS, so that its full effects on the immune system can be understood. The virus is transmitted through semen, vaginal fluids, and blood, and can be caught by risky sexual behaviors and the sharing of needles by intravenous drug users. There are sometimes, but not always, flu-like symptoms in the first 1 to 2 weeks after infection. This is later followed by seroconversion. The anti-HIV antibodies formed during seroconversion are the basis for most initial HIV screening done in the United States. Because seroconversion takes different lengths of time in different individuals, multiple AIDS tests are given months apart to confirm or eliminate the possibility of infection.

After seroconversion, the amount of virus circulating in the blood drops and stays at a low level for several years. During this time, the levels of CD4+ cells, especially helper T cells, decline steadily, until at some point, the immune response is so weak that opportunistic disease and eventually death result. CD4 is the receptor that HIV uses to get inside T cells and reproduce. Given that CD4+ helper T cells play an important role in other in T cell immune responses and antibody responses, it should be no surprise that both types of immune responses are eventually seriously compromised.

Treatment for the disease consists of drugs that target virally encoded proteins that are necessary for viral replication but are absent from normal human cells. By targeting the virus itself and sparing the cells, this approach has been successful in significantly prolonging the lives of HIV-positive individuals. On the other hand, an HIV vaccine has been 30 years in development and is still years away. Because the virus mutates rapidly to evade the immune system, scientists have been looking for parts of the virus that do not change and thus would be good targets for a vaccine candidate.

Hypersensitivities

The word “hypersensitivity” simply means sensitive beyond normal levels of activation. Allergies and inflammatory responses to nonpathogenic environmental substances have been observed since the dawn of history. Hypersensitivity is a medical term describing symptoms that are now known to be caused by unrelated mechanisms of immunity. Still, it is useful for this discussion to use the four types of hypersensitivities as a guide to understand these mechanisms (Figure 1).

[caption id="" align="aligncenter" width="525"]This table describes different types of hypersensitivity. In Type I (IgE-Mediated Hypersensitivity), IgE is bound to mast cells via its Fc portion. When an allergen binds to these antibodies, crosslinking of IgE induces degranulation. Type I causes localized and systemic anaphylaxis, seasonal allergies including hay fever, food allergies such as those to shellfish and peanuts, hives, and eczema. In Type II (IgG-Mediated Hypersensitivity), cells are destroyed by bound antibody, either by activation of complement or by a cytotoxic T cell with an Fc receptor for the antibody (ADCC). Examples are when red blood cells are destroyed by complement and antibody during a transfusion of mismatched blood types or during erythroblastosis fetalis. In Type III (Immune Complex-Mediated Hypersensitivity), antigen-antibody complexes are deposited in tissues, causing activation of complement, which attracts neutrophils to the site. Most common forms of immune complex disease are seen in glomerulonephritis, rheumatoid arthritis, and systemic lupus erythematosus. In Type IV (Cell-Mediated Hypersensitivity), Th1 cells secrete cytokines, which activate macrophages and cytotoxic T cells and can cause macrophage accumulation at the site. Most common forms are contact dermatitis, tuberculin reaction, and autoimmune diseases such as diabetes mellitus type I, multiple sclerosis, and rheumatoid arthritis. Figure 1. Immune Hypersensitivity. Components of the immune system cause four types of hypersensitivity. Notice that types I–III are B cell mediated, whereas type IV hypersensitivity is exclusively a T cell phenomenon.[/caption]

Immediate (Type I) Hypersensitivity

Antigens that cause allergic responses are often referred to as allergens. The specificity of the immediate hypersensitivity response is predicated on the binding of allergen-specific IgE to the mast cell surface. The process of producing allergen-specific IgE is called sensitization, and is a necessary prerequisite for the symptoms of immediate hypersensitivity to occur. Allergies and allergic asthma are mediated by mast cell degranulation that is caused by the crosslinking of the antigen-specific IgE molecules on the mast cell surface. The mediators released have various vasoactive effects already discussed, but the major symptoms of inhaled allergens are the nasal edema and runny nose caused by the increased vascular permeability and increased blood flow of nasal blood vessels. As these mediators are released with mast cell degranulation, type I hypersensitivity reactions are usually rapid and occur within just a few minutes, hence the term immediate hypersensitivity.

Most allergens are in themselves nonpathogenic and therefore innocuous. Some individuals develop mild allergies, which are usually treated with antihistamines. Others develop severe allergies that may cause anaphylactic shock, which can potentially be fatal within 20 to 30 minutes if untreated. This drop in blood pressure (shock) with accompanying contractions of bronchial smooth muscle is caused by systemic mast cell degranulation when an allergen is eaten (for example, shellfish and peanuts), injected (by a bee sting or being administered penicillin), or inhaled (asthma). Because epinephrine raises blood pressure and relaxes bronchial smooth muscle, it is routinely used to counteract the effects of anaphylaxis and can be lifesaving. Patients with known severe allergies are encouraged to keep automatic epinephrine injectors with them at all times, especially when away from easy access to hospitals.

Allergists use skin testing to identify allergens in type I hypersensitivity. In skin testing, allergen extracts are injected into the epidermis, and a positive result of a soft, pale swelling at the site surrounded by a red zone (called the wheal and flare response), caused by the release of histamine and the granule mediators, usually occurs within 30 minutes. The soft center is due to fluid leaking from the blood vessels and the redness is caused by the increased blood flow to the area that results from the dilation of local blood vessels at the site.

Type II and Type III Hypersensitivities

Type II hypersensitivity, which involves IgG-mediated lysis of cells by complement proteins, occurs during mismatched blood transfusions and blood compatibility diseases such as erythroblastosis fetalis (see section on transplantation). Type III hypersensitivity occurs with diseases such as systemic lupus erythematosus, where soluble antigens, mostly DNA and other material from the nucleus, and antibodies accumulate in the blood to the point that the antigen and antibody precipitate along blood vessel linings. These immune complexes often lodge in the kidneys, joints, and other organs where they can activate complement proteins and cause inflammation.

Delayed (Type IV) Hypersensitivity

Delayed hypersensitivity, or type IV hypersensitivity, is basically a standard cellular immune response. In delayed hypersensitivity, the first exposure to an antigen is called sensitization, such that on re-exposure, a secondary cellular response results, secreting cytokines that recruit macrophages and other phagocytes to the site. These sensitized T cells, of the Th1 class, will also activate cytotoxic T cells. The time it takes for this reaction to occur accounts for the 24- to 72-hour delay in development.

The classical test for delayed hypersensitivity is the tuberculin test for tuberculosis, where bacterial proteins from M. tuberculosis are injected into the skin. A couple of days later, a positive test is indicated by a raised red area that is hard to the touch, called an induration, which is a consequence of the cellular infiltrate, an accumulation of activated macrophages. A positive tuberculin test means that the patient has been exposed to the bacteria and exhibits a cellular immune response to it.

Another type of delayed hypersensitivity is contact sensitivity, where substances such as the metal nickel cause a red and swollen area upon contact with the skin. The individual must have been previously sensitized to the metal. A much more severe case of contact sensitivity is poison ivy, but many of the harshest symptoms of the reaction are associated with the toxicity of its oils and are not T cell mediated.

Autoimmune Responses

The worst cases of the immune system over-reacting are autoimmune diseases. Somehow, tolerance breaks down and the immune systems in individuals with these diseases begin to attack their own bodies, causing significant damage. The trigger for these diseases is, more often than not, unknown, and the treatments are usually based on resolving the symptoms using immunosuppressive and anti-inflammatory drugs such as steroids. These diseases can be localized and crippling, as in rheumatoid arthritis, or diffuse in the body with multiple symptoms that differ in different individuals, as is the case with systemic lupus erythematosus (Figure 2).

[caption id="" align="aligncenter" width="500"]The left panel of this figure shows an x-ray image of a person’s hand with rheumatoid arthritis, and the right panel of this figure shows a woman’s body with labels showing the different responses in the body when the patient suffers from lupus. Figure 2. Autoimmune Disorders: Rheumatoid Arthritis and Lupus. (a) Extensive damage to the right hand of a rheumatoid arthritis sufferer is shown in the x-ray. (b) The diagram shows a variety of possible symptoms of systemic lupus erythematosus.[/caption]

Environmental triggers seem to play large roles in autoimmune responses. One explanation for the breakdown of tolerance is that, after certain bacterial infections, an immune response to a component of the bacterium cross-reacts with a self-antigen. This mechanism is seen in rheumatic fever, a result of infection with Streptococcus bacteria, which causes strep throat. The antibodies to this pathogen’s M protein cross-react with an antigenic component of heart myosin, a major contractile protein of the heart that is critical to its normal function. The antibody binds to these molecules and activates complement proteins, causing damage to the heart, especially to the heart valves. On the other hand, some theories propose that having multiple common infectious diseases actually prevents autoimmune responses. The fact that autoimmune diseases are rare in countries that have a high incidence of infectious diseases supports this idea, another example of the hygiene hypothesis discussed earlier in this chapter.

There are genetic factors in autoimmune diseases as well. Some diseases are associated with the MHC genes that an individual expresses. The reason for this association is likely because if one’s MHC molecules are not able to present a certain self-antigen, then that particular autoimmune disease cannot occur. Overall, there are more than 80 different autoimmune diseases, which are a significant health problem in the elderly. Table 7 lists several of the most common autoimmune diseases, the antigens that are targeted, and the segment of the adaptive immune response that causes the damage.

Autoimmune Diseases (Table 7)
Disease Autoantigen Symptoms
Celiac disease Tissue transglutaminase Damage to small intestine
Diabetes mellitus type I Beta cells of pancreas Low insulin production; inability to regulate serum glucose
Graves’ disease Thyroid-stimulating hormone receptor (antibody blocks receptor) Hyperthyroidism
Hashimoto’s thyroiditis Thyroid-stimulating hormone receptor (antibody mimics hormone and stimulates receptor) Hypothyroidism
Lupus erythematosus Nuclear DNA and proteins Damage of many body systems
Myasthenia gravis Acetylcholine receptor in neuromuscular junctions Debilitating muscle weakness
Rheumatoid arthritis Joint capsule antigens Chronic inflammation of joints
]]> 2348 2310 7 0 21.7 Transplantation and Cancer Immunology https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2351 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2351

Learning Objectives

By the end of this section, you will be able to:
  • Explain why blood typing is important and what happens when mismatched blood is used in a transfusion
  • Describe how tissue typing is done during organ transplantation and the role of transplant anti-rejection drugs
  • Show how the immune response is able to control some cancers and how this immune response might be enhanced by cancer vaccines

The immune responses to transplanted organs and to cancer cells are both important medical issues. With the use of tissue typing and anti-rejection drugs, transplantation of organs and the control of the anti-transplant immune response have made huge strides in the past 50 years. Today, these procedures are commonplace. Tissue typing is the determination of MHC molecules in the tissue to be transplanted to better match the donor to the recipient. The immune response to cancer, on the other hand, has been more difficult to understand and control. Although it is clear that the immune system can recognize some cancers and control them, others seem to be resistant to immune mechanisms.

The Rh Factor

Red blood cells can be typed based on their surface antigens. ABO blood type, in which individuals are type A, B, AB, or O according to their genetics, is one example. A separate antigen system seen on red blood cells is the Rh antigen. When someone is “A positive” for example, the positive refers to the presence of the Rh antigen, whereas someone who is “A negative” would lack this molecule.

An interesting consequence of Rh factor expression is seen in erythroblastosis fetalis, a hemolytic disease of the newborn (Figure 1). This disease occurs when mothers negative for Rh antigen have multiple Rh-positive children. During the birth of a first Rh-positive child, the mother makes a primary anti-Rh antibody response to the fetal blood cells that enter the maternal bloodstream. If the mother has a second Rh-positive child, IgG antibodies against Rh-positive blood mounted during this secondary response cross the placenta and attack the fetal blood, causing anemia. This is a consequence of the fact that the fetus is not genetically identical to the mother, and thus the mother is capable of mounting an immune response against it. This disease is treated with antibodies specific for Rh factor. These are given to the mother during the subsequent births, destroying any fetal blood that might enter her system and preventing the immune response.

[caption id="" align="aligncenter" width="500"]This figure shows the progression of thedisease called erythroblastosis fetalis. The top panel shows the umbilical artery and vein and the placenta. The center panel shows the response in the immune system of a first Rh+ infant. The bottom panel shows the response in the case of a second exposure for a Rh+ infant. Figure 1. Erythroblastosis Fetalis. Erythroblastosis fetalis (hemolytic disease of the newborn) is the result of an immune response in an Rh-negative mother who has multiple children with an Rh-positive father. During the first birth, fetal blood enters the mother’s circulatory system, and anti-Rh antibodies are made. During the gestation of the second child, these antibodies cross the placenta and attack the blood of the fetus. The treatment for this disease is to give the mother anti-Rh antibodies (RhoGAM) during the first pregnancy to destroy Rh-positive fetal red blood cells from entering her system and causing the anti-Rh antibody response in the first place.[/caption]

Tissue Transplantation

Tissue transplantation is more complicated than blood transfusions because of two characteristics of MHC molecules. These molecules are the major cause of transplant rejection (hence the name “histocompatibility”). MHC polygeny refers to the multiple MHC proteins on cells, and MHC polymorphism refers to the multiple alleles for each individual MHC locus. Thus, there are many alleles in the human population that can be expressed (Table 8 and Table 9). When a donor organ expresses MHC molecules that are different from the recipient, the latter will often mount a cytotoxic T cell response to the organ and reject it. Histologically, if a biopsy of a transplanted organ exhibits massive infiltration of T lymphocytes within the first weeks after transplant, it is a sign that the transplant is likely to fail. The response is a classical, and very specific, primary T cell immune response. As far as medicine is concerned, the immune response in this scenario does the patient no good at all and causes significant harm.

Partial Table of Alleles of the Human MHC (Class I) (Table 8)
Gene # of alleles # of possible MHC I protein components
A 2132 1527
B 2798 2110
C 1672 1200
E 11 3
F 22 4
G 50 16
Partial Table of Alleles of the Human MHC (Class II) (Table 9)
Gene # of alleles # of possible MHC II protein components
DRA 7 2
DRB 1297 958
DQA1 49 31
DQB1 179 128
DPA1 36 18
DPB1 158 136
DMA 7 4
DMB 13 7
DOA 12 3
DOB 13 5

Immunosuppressive drugs such as cyclosporine A have made transplants more successful, but matching the MHC molecules is still key. In humans, there are six MHC molecules that show the most polymorphisms, three class I molecules (A, B, and C) and three class II molecules called DP, DQ, and DR. A successful transplant usually requires a match between at least 3–4 of these molecules, with more matches associated with greater success. Family members, since they share a similar genetic background, are much more likely to share MHC molecules than unrelated individuals do. In fact, due to the extensive polymorphisms in these MHC molecules, unrelated donors are found only through a worldwide database. The system is not foolproof however, as there are not enough individuals in the system to provide the organs necessary to treat all patients needing them.

One disease of transplantation occurs with bone marrow transplants, which are used to treat various diseases, including SCID and leukemia. Because the bone marrow cells being transplanted contain lymphocytes capable of mounting an immune response, and because the recipient’s immune response has been destroyed before receiving the transplant, the donor cells may attack the recipient tissues, causing graft-versus-host disease. Symptoms of this disease, which usually include a rash and damage to the liver and mucosa, are variable, and attempts have been made to moderate the disease by first removing mature T cells from the donor bone marrow before transplanting it.

Immune Responses Against Cancer

It is clear that with some cancers, for example Kaposi’s sarcoma, a healthy immune system does a good job at controlling them (Figure 2). This disease, which is caused by the human herpesvirus, is almost never observed in individuals with strong immune systems, such as the young and immunocompetent. Other examples of cancers caused by viruses include liver cancer caused by the hepatitis B virus and cervical cancer caused by the human papilloma virus. As these last two viruses have vaccines available for them, getting vaccinated can help prevent these two types of cancer by stimulating the immune response.

[caption id="" align="aligncenter" width="380"]This photograph shows lesions on the surface of skin. Figure 2. Karposi’s Sarcoma Lesions. (credit: National Cancer Institute)[/caption]

On the other hand, as cancer cells are often able to divide and mutate rapidly, they may escape the immune response, just as certain pathogens such as HIV do. There are three stages in the immune response to many cancers: elimination, equilibrium, and escape. Elimination occurs when the immune response first develops toward tumor-specific antigens specific to the cancer and actively kills most cancer cells, followed by a period of controlled equilibrium during which the remaining cancer cells are held in check. Unfortunately, many cancers mutate, so they no longer express any specific antigens for the immune system to respond to, and a subpopulation of cancer cells escapes the immune response, continuing the disease process.

This fact has led to extensive research in trying to develop ways to enhance the early immune response to completely eliminate the early cancer and thus prevent a later escape. One method that has shown some success is the use of cancer vaccines, which differ from viral and bacterial vaccines in that they are directed against the cells of one’s own body. Treated cancer cells are injected into cancer patients to enhance their anti-cancer immune response and thereby prolong survival. The immune system has the capability to detect these cancer cells and proliferate faster than the cancer cells do, overwhelming the cancer in a similar way as they do for viruses. Cancer vaccines have been developed for malignant melanoma, a highly fatal skin cancer, and renal (kidney) cell carcinoma. These vaccines are still in the development stages, but some positive and encouraging results have been obtained clinically.

It is tempting to focus on the complexity of the immune system and the problems it causes as a negative. The upside to immunity, however, is so much greater: The benefit of staying alive far outweighs the negatives caused when the system does sometimes go awry. Working on “autopilot,” the immune system helps to maintain your health and kill pathogens. The only time you really miss the immune response is when it is not being effective and illness results, or, as in the extreme case of HIV disease, the immune system is gone completely.

]]> 2351 2310 8 0 24.6 Energy and Heat Balance https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2407 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2407

Learning Objectives

By the end of this section, you will be able to:
  • Describe how the body regulates temperature
  • Explain the significance of the metabolic rate

The body tightly regulates the body temperature through a process called thermoregulation, in which the body can maintain its temperature within certain boundaries, even when the surrounding temperature is very different. The core temperature of the body remains steady at around 36.5–37.5 °C (or 97.7–99.5 °F). In the process of ATP production by cells throughout the body, approximately 60 percent of the energy produced is in the form of heat used to maintain body temperature. Thermoregulation is an example of negative feedback.

The hypothalamus in the brain is the master switch that works as a thermostat to regulate the body’s core temperature (Figure 1). If the temperature is too high, the hypothalamus can initiate several processes to lower it. These include increasing the circulation of the blood to the surface of the body to allow for the dissipation of heat through the skin and initiation of sweating to allow evaporation of water on the skin to cool its surface. Conversely, if the temperature falls below the set core temperature, the hypothalamus can initiate shivering to generate heat. The body uses more energy and generates more heat. In addition, thyroid hormone will stimulate more energy use and heat production by cells throughout the body. An environment is said to be thermoneutral when the body does not expend or release energy to maintain its core temperature. For a naked human, this is an ambient air temperature of around 84 °F. If the temperature is higher, for example, when wearing clothes, the body compensates with cooling mechanisms. The body loses heat through the mechanisms of heat exchange.

[caption id="attachment_1826" align="aligncenter" width="500"]This figure shows the pathways in which body temperature is controlled by the hypothalamus. Figure 1. Hypothalamus Controls Thermoregulation. The hypothalamus controls thermoregulation.[/caption]

Mechanisms of Heat Exchange

When the environment is not thermoneutral, the body uses four mechanisms of heat exchange to maintain homeostasis: conduction, convection, radiation, and evaporation. Each of these mechanisms relies on the property of heat to flow from a higher concentration to a lower concentration; therefore, each of the mechanisms of heat exchange varies in rate according to the temperature and conditions of the environment.

Conduction is the transfer of heat by two objects that are in direct contact with one another. It occurs when the skin comes in contact with a cold or warm object. For example, when holding a glass of ice water, the heat from your skin will warm the glass and in turn melt the ice. Alternatively, on a cold day, you might warm up by wrapping your cold hands around a hot mug of coffee. Only about 3 percent of the body’s heat is lost through conduction.

Convection is the transfer of heat to the air surrounding the skin. The warmed air rises away from the body and is replaced by cooler air that is subsequently heated. Convection can also occur in water. When the water temperature is lower than the body’s temperature, the body loses heat by warming the water closest to the skin, which moves away to be replaced by cooler water. The convection currents created by the temperature changes continue to draw heat away from the body more quickly than the body can replace it, resulting in hyperthermia. About 15 percent of the body’s heat is lost through convection.

Radiation is the transfer of heat via infrared waves. This occurs between any two objects when their temperatures differ. A radiator can warm a room via radiant heat. On a sunny day, the radiation from the sun warms the skin. The same principle works from the body to the environment. About 60 percent of the heat lost by the body is lost through radiation.

Evaporation is the transfer of heat by the evaporation of water. Because it takes a great deal of energy for a water molecule to change from a liquid to a gas, evaporating water (in the form of sweat) takes with it a great deal of energy from the skin. However, the rate at which evaporation occurs depends on relative humidity—more sweat evaporates in lower humidity environments. Sweating is the primary means of cooling the body during exercise, whereas at rest, about 20 percent of the heat lost by the body occurs through evaporation.

]]> 2407 2405 1 0 Muscle Anatomy Review https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2419 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2419 All images by Henry Gray (1918), Anatomy of the Human Body, Bartleby.com imageGray410.pngimageimageimageimageimageimageimageimage http://www.bartleby.com/107/indexillus.html

]]> 2419 2408 1 0 12.2 Nervous Tissue https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2491 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2491

Learning Objectives

By the end of this section, you will be able to:
  • Describe the basic structure of a neuron
  • Identify the different types of neurons on the basis of polarity
  • List the glial cells of the CNS and describe their function
  • List the glial cells of the PNS and describe their function

Nervous tissue is composed of two types of cells, neurons and glial cells. Neurons are the primary type of cell that most anyone associates with the nervous system. They are responsible for the computation and communication that the nervous system provides. They are electrically active and release chemical signals to target cells. Glial cells, or glia, are known to play a supporting role for nervous tissue. Ongoing research pursues an expanded role that glial cells might play in signaling, but neurons are still considered the basis of this function. Neurons are important, but without glial support they would not be able to perform their function.

Neurons

Neurons are the cells considered to be the basis of nervous tissue. They are responsible for the electrical signals that communicate information about sensations, and that produce movements in response to those stimuli, along with inducing thought processes within the brain. An important part of the function of neurons is in their structure, or shape. The three-dimensional shape of these cells makes the immense numbers of connections within the nervous system possible.

Parts of a Neuron

As you learned in the first section, the main part of a neuron is the cell body, which is also known as the soma (soma = “body”). The cell body contains the nucleus and most of the major organelles. But what makes neurons special is that they have many extensions of their cell membranes, which are generally referred to as processes. Neurons are usually described as having one, and only one, axon—a fiber that emerges from the cell body and projects to target cells. That single axon can branch repeatedly to communicate with many target cells. It is the axon that propagates the nerve impulse, which is communicated to one or more cells. The other processes of the neuron are dendrites, which receive information from other neurons at specialized areas of contact called synapses. The dendrites are usually highly branched processes, providing locations for other neurons to communicate with the cell body. Information flows through a neuron from the dendrites, across the cell body, and down the axon. This gives the neuron a polarity—meaning that information flows in this one direction. Figure 1 shows the relationship of these parts to one another.

[caption id="" align="aligncenter" width="380"]This illustration shows the anatomy of a neuron. The neuron has a very irregular cell body (soma) containing a purple nucleus. There are six projections protruding from the top, bottom and left side of the cell body. Each of the projections branches many times, forming small, tree-shaped structures protruding from the cell body. The right side of the cell body tapers into a long cord called the axon. The axon is insulated by segments of myelin sheath, which resemble a semitransparent toilet paper roll wound around the axon. The myelin sheath is not continuous, but is separated into equally spaced segments. The bare axon segments between the sheath segments are called nodes of Ranvier. An oligodendrocyte is reaching its two arm like projections onto two myelin sheath segments. The axon branches many times at its end, where it connects to the dendrites of another neuron. Each connection between an axon branch and a dendrite is called a synapse. The cell membrane completely surrounds the cell body, dendrites, and its axon. The axon of another nerve is seen in the upper left of the diagram connecting with the dendrites of the central neuron. Figure 1. Parts of a Neuron. The major parts of the neuron are labeled on a multipolar neuron from the CNS.[/caption]

Where the axon emerges from the cell body, there is a special region referred to as the axon hillock. This is a tapering of the cell body toward the axon fiber. Within the axon hillock, the cytoplasm changes to a solution of limited components called axoplasm. Because the axon hillock represents the beginning of the axon, it is also referred to as the initial segment.

Many axons are wrapped by an insulating substance called myelin, which is actually made from glial cells. Myelin acts as insulation much like the plastic or rubber that is used to insulate electrical wires. A key difference between myelin and the insulation on a wire is that there are gaps in the myelin covering of an axon. Each gap is called a node of Ranvier and is important to the way that electrical signals travel down the axon. The length of the axon between each gap, which is wrapped in myelin, is referred to as an axon segment. At the end of the axon is the axon terminal, where there are usually several branches extending toward the target cell, each of which ends in an enlargement called a synaptic end bulb. These bulbs are what make the connection with the target cell at the synapse.

Types of Neurons

There are many neurons in the nervous system—a number in the trillions. And there are many different types of neurons. They can be classified by many different criteria. The first way to classify them is by the number of processes attached to the cell body. Using the standard model of neurons, one of these processes is the axon, and the rest are dendrites. Because information flows through the neuron from dendrites or cell bodies toward the axon, these names are based on the neuron's polarity (Figure 2).

[caption id="" align="aligncenter" width="420"]Three illustrations show some of the possible shapes that neurons can take. In the unipolar neuron, the dendrite enters from the left and merges with the axon into a common pathway, which is connected to the cell body. The axon leaves the cell body through the common pathway, the branches off to the right, in the opposite direction as the dendrite. Therefore, this neuron is T shaped. In the bipolar neuron, the dendrite enters into the left side of the cell body while the axon emerges from the opposite (right) side. In a multipolar neuron, multiple dendrites enter into the cell body. The only part of the cell body that does not have dendrites is the part that elongates into the axon. Figure 2. Neuron Classification by Shape. Unipolar cells have one process that includes both the axon and dendrite. Bipolar cells have two processes, the axon and a dendrite. Multipolar cells have more than two processes, the axon and two or more dendrites.[/caption]

Unipolar cells have only one process emerging from the cell. True unipolar cells are only found in invertebrate animals, so the unipolar cells in humans are more appropriately called “pseudo-unipolar” cells. Invertebrate unipolar cells do not have dendrites. Human unipolar cells have an axon that emerges from the cell body, but it splits so that the axon can extend along a very long distance. At one end of the axon are dendrites, and at the other end, the axon forms synaptic connections with a target. Unipolar cells are exclusively sensory neurons and have two unique characteristics. First, their dendrites are receiving sensory information, sometimes directly from the stimulus itself. Secondly, the cell bodies of unipolar neurons are always found in ganglia. Sensory reception is a peripheral function (those dendrites are in the periphery, perhaps in the skin) so the cell body is in the periphery, though closer to the CNS in a ganglion. The axon projects from the dendrite endings, past the cell body in a ganglion, and into the central nervous system.

Bipolar cells have two processes, which extend from each end of the cell body, opposite to each other. One is the axon and one the dendrite. Bipolar cells are not very common. They are found mainly in the olfactory epithelium (where smell stimuli are sensed), and as part of the retina.

Multipolar neurons are all of the neurons that are not unipolar or bipolar. They have one axon and two or more dendrites (usually many more). With the exception of the unipolar sensory ganglion cells, and the two specific bipolar cells mentioned above, all other neurons are multipolar. Some cutting edge research suggests that certain neurons in the CNS do not conform to the standard model of “one, and only one” axon. Some sources describe a fourth type of neuron, called an anaxonic neuron. The name suggests that it has no axon (an- = “without”), but this is not accurate. Anaxonic neurons are very small, and if you look through a microscope at the standard resolution used in histology (approximately 400X to 1000X total magnification), you will not be able to distinguish any process specifically as an axon or a dendrite. Any of those processes can function as an axon depending on the conditions at any given time. Nevertheless, even if they cannot be easily seen, and one specific process is definitively the axon, these neurons have multiple processes and are therefore multipolar.

Neurons can also be classified on the basis of where they are found, who found them, what they do, or even what chemicals they use to communicate with each other. Some neurons referred to in this section on the nervous system are named on the basis of those sorts of classifications (Figure 3). For example, a multipolar neuron that has a very important role to play in a part of the brain called the cerebellum is known as a Purkinje (commonly pronounced per-KIN-gee) cell. It is named after the anatomist who discovered it (Jan Evangilista Purkinje, 1787–1869).

[caption id="" align="aligncenter" width="450"]This diagram contains three black and white drawings of more specialized nerve cells. Part A shows a pyramidal cell of the cerebral cortex, which has two, long, nerve tracts attached to the top and bottom of the cell body. However, the cell body also has many shorter dendrites projecting out a short distance from the cell body. Part B shows a Purkinje cell of the cerebellar cortex. This cell has a single, long, nerve tract entering the bottom of the cell body. Two large nerve tracts leave the top of the cell body but immediately branch many times to form a large web of nerve fibers. Therefore, the purkinje cell somewhat resembles a shrub or coral in shape. Part C shows the olfactory cells in the olfactory epithelium and olfactory bulbs. It contains several cell groups linked together. At the bottom, there is a row of olfactory epithelial cells that are tightly packed, side-by-side, somewhat resembling the slats on a fence. There are six neurons embedded in this epithelium. Each neuron connects to the epithelium through branching nerve fibers projecting from the bottom of their cell bodies. A single nerve fiber projects from the top of each neuron and synapses with nerve fibers from the neurons above. These upper neurons are cross shaped, with one nerve fiber projecting from the bottom, top, right and left sides. The upper cells synapse with the epithelial nerve cells using the nerve tract projecting from the bottom of their cell body. The nerve tract projecting from the top continues the pathway, making a ninety degree turn to the right and continuing to the right border of the image. Figure 3. Other Neuron Classifications. Three examples of neurons that are classified on the basis of other criteria. (a) The pyramidal cell is a multipolar cell with a cell body that is shaped something like a pyramid. (b) The Purkinje cell in the cerebellum was named after the scientist who originally described it. (c) Olfactory neurons are named for the functional group with which they belong.[/caption]

Glial Cells

Glial cells, or neuroglia or simply glia, are the other type of cell found in nervous tissue. They are considered to be supporting cells, and many functions are directed at helping neurons complete their function for communication. The name glia comes from the Greek word that means “glue,” and was coined by the German pathologist Rudolph Virchow, who wrote in 1856: “This connective substance, which is in the brain, the spinal cord, and the special sense nerves, is a kind of glue (neuroglia) in which the nervous elements are planted.” Today, research into nervous tissue has shown that there are many deeper roles that these cells play. And research may find much more about them in the future.

There are six types of glial cells. Four of them are found in the CNS and two are found in the PNS. Table 2 outlines some common characteristics and functions.

Glial Cell Types by Location and Basic Function (Table 2)
CNS glia PNS glia Basic function
Astrocyte Satellite cell Support
Oligodendrocyte Schwann cell Insulation, myelination
Microglia - Immune surveillance and phagocytosis
Ependymal cell - Creating CSF

Glial Cells of the CNS

One cell providing support to neurons of the CNS is the astrocyte, so named because it appears to be star-shaped under the microscope (astro- = “star”). Astrocytes have many processes extending from their main cell body (not axons or dendrites like neurons, just cell extensions). Those processes extend to interact with neurons, blood vessels, or the connective tissue covering the CNS that is called the pia mater (Figure 4). Generally, they are supporting cells for the neurons in the central nervous system. Some ways in which they support neurons in the central nervous system are by maintaining the concentration of chemicals in the extracellular space, removing excess signaling molecules, reacting to tissue damage, and contributing to the blood-brain barrier (BBB). The blood-brain barrier is a physiological barrier that keeps many substances that circulate in the rest of the body from getting into the central nervous system, restricting what can cross from circulating blood into the CNS. Nutrient molecules, such as glucose or amino acids, can pass through the BBB, but other molecules cannot. This actually causes problems with drug delivery to the CNS. Pharmaceutical companies are challenged to design drugs that can cross the BBB as well as have an effect on the nervous system.

[caption id="" align="aligncenter" width="420"]This diagram shows several types of nervous system cells associated with two multipolar neurons. Astrocytes are star shaped-cells with many dendrite like projections but no axon. They are connected with the multipolar neurons and other cells in the diagram through their dendrite like projections. Ependymal cells have a teardrop shaped cell body and a long tail that branches several times before connecting with astrocytes and the multipolar neuron. Microglial cells are small cells with rectangular bodies and many dendrite like projections stemming from their shorter sides. The projections are so extensive that they give the microglial cell a fuzzy appearance. The oligodendrocytes have circular cell bodies with four dendrite like projections. Each projection is connected to a segment of myelin sheath on the axons of the multipolar neurons. The oligodendrocytes are the same color as the myelin sheath segment and are adding layers to the sheath using their projections. Figure 4. Glial Cells of the CNS. The CNS has astrocytes, oligodendrocytes, microglia, and ependymal cells that support the neurons of the CNS in several ways.[/caption]

Like a few other parts of the body, the brain has a privileged blood supply. Very little can pass through by diffusion. Most substances that cross the wall of a blood vessel into the CNS must do so through an active transport process. Because of this, only specific types of molecules can enter the CNS. Glucose—the primary energy source—is allowed, as are amino acids. Water and some other small particles, like gases and ions, can enter. But most everything else cannot, including white blood cells, which are one of the body’s main lines of defense. While this barrier protects the CNS from exposure to toxic or pathogenic substances, it also keeps out the cells that could protect the brain and spinal cord from disease and damage. The BBB also makes it harder for pharmaceuticals to be developed that can affect the nervous system. Aside from finding efficacious substances, the means of delivery is also crucial.

Also found in CNS tissue is the oligodendrocyte, sometimes called just “oligo,” which is the glial cell type that insulates axons in the CNS. The name means “cell of a few branches” (oligo- = “few”; dendro- = “branches”; -cyte = “cell”). There are a few processes that extend from the cell body. Each one reaches out and surrounds an axon to insulate it in myelin. One oligodendrocyte will provide the myelin for multiple axon segments, either for the same axon or for separate axons. The function of myelin will be discussed below.

Microglia are, as the name implies, smaller than most of the other glial cells. Ongoing research into these cells, although not entirely conclusive, suggests that they may originate as white blood cells, called macrophages, that become part of the CNS during early development. While their origin is not conclusively determined, their function is related to what macrophages do in the rest of the body. When macrophages encounter diseased or damaged cells in the rest of the body, they ingest and digest those cells or the pathogens that cause disease. Microglia are the cells in the CNS that can do this in normal, healthy tissue, and they are therefore also referred to as CNS-resident macrophages.

The ependymal cell is a glial cell that filters blood to make cerebrospinal fluid (CSF), the fluid that circulates through the CNS. Because of the privileged blood supply inherent in the BBB, the extracellular space in nervous tissue does not easily exchange components with the blood. Ependymal cells line each ventricle, one of four central cavities that are remnants of the hollow center of the neural tube formed during the embryonic development of the brain. The choroid plexus is a specialized structure in the ventricles where ependymal cells come in contact with blood vessels and filter and absorb components of the blood to produce cerebrospinal fluid. Because of this, ependymal cells can be considered a component of the BBB, or a place where the BBB breaks down. These glial cells appear similar to epithelial cells, making a single layer of cells with little intracellular space and tight connections between adjacent cells. They also have cilia on their apical surface to help move the CSF through the ventricular space. The relationship of these glial cells to the structure of the CNS is seen in Figure 4.

Glial Cells of the PNS

One of the two types of glial cells found in the PNS is the satellite cell. Satellite cells are found in sensory and autonomic ganglia, where they surround the cell bodies of neurons. This accounts for the name, based on their appearance under the microscope. They provide support, performing similar functions in the periphery as astrocytes do in the CNS—except, of course, for establishing the BBB.

The second type of glial cell is the Schwann cell, which insulate axons with myelin in the periphery. Schwann cells are different than oligodendrocytes, in that a Schwann cell wraps around a portion of only one axon segment and no others. Oligodendrocytes have processes that reach out to multiple axon segments, whereas the entire Schwann cell surrounds just one axon segment. The nucleus and cytoplasm of the Schwann cell are on the edge of the myelin sheath. The relationship of these two types of glial cells to ganglia and nerves in the PNS is seen in Figure 5.

[caption id="" align="aligncenter" width="420"]This diagram shows a collection of PNS glial cells. The largest cell is a unipolar peripheral ganglionic neuron which has a common nerve tract projecting from the bottom of its cell body. The common nerve tract then splits into the axon, going off to the left, and the dendrite, going off to the right. The cell body of the neuron is covered with several satellite cells that are irregular, flattened, and take on the appearance of fried eggs. Schwann cells wrap around each myelin sheath segment on the axon, with their nucleus creating a small bump on each segment. Figure 5. Glial Cells of the PNS. The PNS has satellite cells and Schwann cells.[/caption]

Myelin

The insulation for axons in the nervous system is provided by glial cells, oligodendrocytes in the CNS, and Schwann cells in the PNS. Whereas the manner in which either cell is associated with the axon segment, or segments, that it insulates is different, the means of myelinating an axon segment is mostly the same in the two situations. Myelin is a lipid-rich sheath that surrounds the axon and by doing so creates a myelin sheath that facilitates the transmission of electrical signals along the axon. The lipids are essentially the phospholipids of the glial cell membrane. Myelin, however, is more than just the membrane of the glial cell. It also includes important proteins that are integral to that membrane. Some of the proteins help to hold the layers of the glial cell membrane closely together.

The appearance of the myelin sheath can be thought of as similar to the pastry wrapped around a hot dog for “pigs in a blanket” or a similar food. The glial cell is wrapped around the axon several times with little to no cytoplasm between the glial cell layers. For oligodendrocytes, the rest of the cell is separate from the myelin sheath as a cell process extends back toward the cell body. A few other processes provide the same insulation for other axon segments in the area. For Schwann cells, the outermost layer of the cell membrane contains cytoplasm and the nucleus of the cell as a bulge on one side of the myelin sheath. During development, the glial cell is loosely or incompletely wrapped around the axon (Figure 6a). The edges of this loose enclosure extend toward each other, and one end tucks under the other. The inner edge wraps around the axon, creating several layers, and the other edge closes around the outside so that the axon is completely enclosed.

Myelin sheaths can extend for one or two millimeters, depending on the diameter of the axon. Axon diameters can be as small as 1 to 20 micrometers. Because a micrometer is 1/1000 of a millimeter, this means that the length of a myelin sheath can be 100–1000 times the diameter of the axon. Figure 1, Figure 4, and Figure 5 show the myelin sheath surrounding an axon segment, but are not to scale. If the myelin sheath were drawn to scale, the neuron would have to be immense—possibly covering an entire wall of the room in which you are sitting.

[caption id="" align="aligncenter" width="550"]This three-part diagram shows the process of myelination. In step A, the cell membrane of a cylindrical Schwann cell, which has a blue nucleus, has indented around an axon. An upper and lower lip of the cell membrane is visible where the membrane indents around the axon. In part B, the lower lip of the cell membrane dives under the upper lip and wraps around the axon. In part C, the process in part B has continued, forming many layers of myelin that wrap around the axon. The nucleus of the Schwann cell is still visible in the outermost layer, just to the left of the upper lip. The area of the axon next to the Schwann cell, which has no myelin, is labeled as a node of Ranvier. Figure 6. The Process of Myelination. Myelinating glia wrap several layers of cell membrane around the cell membrane of an axon segment. A single Schwann cell insulates a segment of a peripheral nerve, whereas in the CNS, an oligodendrocyte may provide insulation for a few separate axon segments. EM × 1,460,000. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]
Disorders of the…

Nervous Tissue Several diseases can result from the demyelination of axons. The causes of these diseases are not the same; some have genetic causes, some are caused by pathogens, and others are the result of autoimmune disorders. Though the causes are varied, the results are largely similar. The myelin insulation of axons is compromised, making electrical signaling slower.

Multiple sclerosis (MS) is one such disease. It is an example of an autoimmune disease. The antibodies produced by lymphocytes (a type of white blood cell) mark myelin as something that should not be in the body. This causes inflammation and the destruction of the myelin in the central nervous system. As the insulation around the axons is destroyed by the disease, scarring becomes obvious. This is where the name of the disease comes from; sclerosis means hardening of tissue, which is what a scar is. Multiple scars are found in the white matter of the brain and spinal cord. The symptoms of MS include both somatic and autonomic deficits. Control of the musculature is compromised, as is control of organs such as the bladder.

Guillain-Barré (pronounced gee-YAN bah-RAY) syndrome is an example of a demyelinating disease of the peripheral nervous system. It is also the result of an autoimmune reaction, but the inflammation is in peripheral nerves. Sensory symptoms or motor deficits are common, and autonomic failures can lead to changes in the heart rhythm or a drop in blood pressure, especially when standing, which causes dizziness.

]]> 2491 2477 3 0 12.5 Communication Between Neurons https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2505 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2505

Learning Objectives

By the end of this section, you will be able to:
  • Explain the differences between the types of graded potentials
  • Categorize the major neurotransmitters by chemical type and effect

The electrical changes taking place within a neuron, as described in the previous section, are similar to a light switch being turned on. A stimulus starts the depolarization, but the action potential runs on its own once a threshold has been reached. The question is now, “What flips the light switch on?” Temporary changes to the cell membrane voltage can result from neurons receiving information from the environment, or from the action of one neuron on another. These special types of potentials influence a neuron and determine whether an action potential will occur or not. Many of these transient signals originate at the synapse.

Graded Potentials

Local changes in the membrane potential are called graded potentials and are usually associated with the dendrites of a neuron. The amount of change in the membrane potential is determined by the size of the stimulus that causes it. In the example of testing the temperature of the shower, slightly warm water would only initiate a small change in a thermoreceptor, whereas hot water would cause a large amount of change in the membrane potential.

Graded potentials can be of two sorts, either they are depolarizing or hyperpolarizing (Figure 1). For a membrane at the resting potential, a graded potential represents a change in that voltage either above -70 mV or below -70 mV. Depolarizing graded potentials are often the result of Na+ or Ca2+ entering the cell. Both of these ions have higher concentrations outside the cell than inside; because they have a positive charge, they will move into the cell causing it to become less negative relative to the outside. Hyperpolarizing graded potentials can be caused by K+ leaving the cell or Cl- entering the cell. If a positive charge moves out of a cell, the cell becomes more negative; if a negative charge enters the cell, the same thing happens.

[caption id="" align="aligncenter" width="500"]The graph has membrane potential, in millivolts, on the X axis, ranging from negative 90 to positive 30. Time is on the X axis. The left half of the plot line is labeled the depolarizing graded potential. The plot has four progressively larger peaks, with each starting at the resting membrane potential of negative 70. The lowest peak reaches to about negative 65 and is narrow in width, as this represents a small stimulus that causes a small depolarization of the cell membrane. The second peak reaches to about negative 60 but is still narrow. This represents a larger stimulus causing more depolarization. The third peak also reaches to negative 60, but is about twice as wide as the other two peaks. This represents a stimulus of longer duration, which causes a longer lasting depolarization. However, this stimulus is not greater in strength than the previous stimulus. The rightmost peak among the depolarizing graded potentials reaches above the threshold line to about negative 51. This represents a stimulus of sufficient strength to trigger an action potential. The right half of the plot is labeled the hyperpolarizing graded potential. The plot line in this half begins at the resting potential of negative 70, but then drops to more negative membrane potentials. The first peak drops to negative 75 EV, the second peak drops to negative 80 EV and the third peak drops to negative 88 EV. These peaks represent a stimulus that results in hyperpolarization, which is triggered by the activation of specific ion channels in the cell membrane. Figure 1. Graded Potentials. Graded potentials are temporary changes in the membrane voltage, the characteristics of which depend on the size of the stimulus. Some types of stimuli cause depolarization of the membrane, whereas others cause hyperpolarization. It depends on the specific ion channels that are activated in the cell membrane.[/caption]

Types of Graded Potentials

For the unipolar cells of sensory neurons—both those with free nerve endings and those within encapsulations—graded potentials develop in the dendrites that influence the generation of an action potential in the axon of the same cell. This is called a generator potential. For other sensory receptor cells, such as taste cells or photoreceptors of the retina, graded potentials in their membranes result in the release of neurotransmitters at synapses with sensory neurons. This is called a receptor potential.

A postsynaptic potential (PSP) is the graded potential in the dendrites of a neuron that is receiving synapses from other cells. Postsynaptic potentials can be depolarizing or hyperpolarizing. Depolarization in a postsynaptic potential is called an excitatory postsynaptic potential (EPSP) because it causes the membrane potential to move toward threshold. Hyperpolarization in a postsynaptic potential is an inhibitory postsynaptic potential (IPSP) because it causes the membrane potential to move away from threshold.

Summation

All types of graded potentials will result in small changes of either depolarization or hyperpolarization in the voltage of a membrane. These changes can lead to the neuron reaching threshold if the changes add together, or summate. The combined effects of different types of graded potentials are illustrated in Figure 2. If the total change in voltage in the membrane is a positive 15 mV, meaning that the membrane depolarizes from -70 mV to -55 mV, then the graded potentials will result in the membrane reaching threshold.

For receptor potentials, threshold is not a factor because the change in membrane potential for receptor cells directly causes neurotransmitter release. However, generator potentials can initiate action potentials in the sensory neuron axon, and postsynaptic potentials can initiate an action potential in the axon of other neurons. Graded potentials summate at a specific location at the beginning of the axon to initiate the action potential, namely the initial segment. For sensory neurons, which do not have a cell body between the dendrites and the axon, the initial segment is directly adjacent to the dendritic endings. For all other neurons, the axon hillock is essentially the initial segment of the axon, and it is where summation takes place. These locations have a high density of voltage-gated Na+ channels that initiate the depolarizing phase of the action potential.

Summation can be spatial or temporal, meaning it can be the result of multiple graded potentials at different locations on the neuron, or all at the same place but separated in time. Spatial summation is related to associating the activity of multiple inputs to a neuron with each other. Temporal summation is the relationship of multiple action potentials from a single cell resulting in a significant change in the membrane potential. Spatial and temporal summation can act together, as well.

[caption id="" align="aligncenter" width="380"]This graph has membrane potential, in millivolts, on the X axis, ranging from negative 90 to negative 40. Time is on the X axis. The plot line is moving up and down between the resting membrane potential of minus 70 EV and the threshold potential of minus 55 EV. An EPSP causes the plot line to move higher, closer to the threshold potential. An IPSP causes the plot line to move lower, further away from the threshold potential. Toward the right side of the graph, the neuron receives an EPSP that pushes the membrane potential above the threshold, triggering an action potential that causes the plot line to quickly rise above positive 30 EV. The plot line then quickly drops back below minus 70 EV but then gradually increases back to minus 70. A picture of a neuron indicates that excitatory post synaptic potentials are commonly provided by synapses on the neuron’s dendrites. Inhibitory post synaptic potentials are commonly provided by synapses near the neuron’s axon hillock. Figure 2. Postsynaptic Potential Summation. The result of summation of postsynaptic potentials is the overall change in the membrane potential. At point A, several different excitatory postsynaptic potentials add up to a large depolarization. At point B, a mix of excitatory and inhibitory postsynaptic potentials result in a different end result for the membrane potential.[/caption]

Synapses

There are two types of connections between electrically active cells, chemical synapses and electrical synapses. In a chemical synapse, a chemical signal—namely, a neurotransmitter—is released from one cell and it affects the other cell. In an electrical synapse, there is a direct connection between the two cells so that ions can pass directly from one cell to the next. If one cell is depolarized in an electrical synapse, the joined cell also depolarizes because the ions pass between the cells. Chemical synapses involve the transmission of chemical information from one cell to the next. This section will concentrate on the chemical type of synapse.

An example of a chemical synapse is the neuromuscular junction (NMJ) described in the chapter on muscle tissue. In the nervous system, there are many more synapses that are essentially the same as the NMJ. All synapses have common characteristics, which can be summarized in this list:

  • presynaptic element
  • neurotransmitter (packaged in vesicles)
  • synaptic cleft
  • receptor proteins
  • postsynaptic element
  • neurotransmitter elimination or re-uptake

For the NMJ, these characteristics are as follows: the presynaptic element is the motor neuron's axon terminals, the neurotransmitter is acetylcholine, the synaptic cleft is the space between the cells where the neurotransmitter diffuses, the receptor protein is the nicotinic acetylcholine receptor, the postsynaptic element is the sarcolemma of the muscle cell, and the neurotransmitter is eliminated by acetylcholinesterase. Other synapses are similar to this, and the specifics are different, but they all contain the same characteristics.

Neurotransmitter Release

When an action potential reaches the axon terminals, voltage-gated Ca2+ channels in the membrane of the synaptic end bulb open. The concentration of Ca2+ increases inside the end bulb, and the Ca2+ ion associates with proteins in the outer surface of neurotransmitter vesicles. The Ca2+ facilitates the merging of the vesicle with the presynaptic membrane so that the neurotransmitter is released through exocytosis into the small gap between the cells, known as the synaptic cleft.

Once in the synaptic cleft, the neurotransmitter diffuses the short distance to the postsynaptic membrane and can interact with neurotransmitter receptors. Receptors are specific for the neurotransmitter, and the two fit together like a key and lock. One neurotransmitter binds to its receptor and will not bind to receptors for other neurotransmitters, making the binding a specific chemical event (Figure 3).

[caption id="" align="aligncenter" width="480"]This diagram shows a postsynaptic neuron. An axon from a presynaptic neuron is synapsing with the dendrites on the post synaptic neuron. The axon of the presynaptic neuron branches into several club shaped axon terminals. A magnified view of one of the synapses reveals that the axon terminal does not contact the dendrite of the postsynaptic neuron. Instead, there is a small space between the two structures, called the synaptic cleft. The axon terminal of the presynaptic neuron contains several synaptic vesicles, each holding about a dozen neurotransmitter particles. The synaptic vesicles travel to the edge of the axon terminal and release their neurotransmitters into the synaptic clefts The neurotransmitters travel through the synaptic cleft and bind to carrier proteins on the postsynaptic neuron that contain receptors foe neurotransmitters. Figure 3. The Synapse. The synapse is a connection between a neuron and its target cell (which is not necessarily a neuron). The presynaptic element is the synaptic end bulb of the axon where Ca2+ enters the bulb to cause vesicle fusion and neurotransmitter release. The neurotransmitter diffuses across the synaptic cleft to bind to its receptor. The neurotransmitter is cleared from the synapse either by enzymatic degradation, neuronal reuptake, or glial reuptake.[/caption]

Neurotransmitter Systems

There are several systems of neurotransmitters that are found at various synapses in the nervous system. These groups refer to the chemicals that are the neurotransmitters, and within the groups are specific systems.

The first group, which is a neurotransmitter system of its own, is the cholinergic system. It is the system based on acetylcholine. This includes the NMJ as an example of a cholinergic synapse, but cholinergic synapses are found in other parts of the nervous system. They are in the autonomic nervous system, as well as distributed throughout the brain.

The cholinergic system has two types of receptors, the nicotinic receptor is found in the NMJ as well as other synapses. There is also an acetylcholine receptor known as the muscarinic receptor. Both of these receptors are named for drugs that interact with the receptor in addition to acetylcholine. Nicotine will bind to the nicotinic receptor and activate it similar to acetylcholine. Muscarine, a product of certain mushrooms, will bind to the muscarinic receptor. However, nicotine will not bind to the muscarinic receptor and muscarine will not bind to the nicotinic receptor.

Another group of neurotransmitters are amino acids. This includes glutamate (Glu), GABA (gamma-aminobutyric acid, a derivative of glutamate), and glycine (Gly). These amino acids have an amino group and a carboxyl group in their chemical structures. Glutamate is one of the 20 amino acids that are used to make proteins. Each amino acid neurotransmitter would be part of its own system, namely the glutamatergic, GABAergic, and glycinergic systems. They each have their own receptors and do not interact with each other. Amino acid neurotransmitters are eliminated from the synapse by reuptake. A pump in the cell membrane of the presynaptic element, or sometimes a neighboring glial cell, will clear the amino acid from the synaptic cleft so that it can be recycled, repackaged in vesicles, and released again.

Another class of neurotransmitter is the biogenic amine, a group of neurotransmitters that are enzymatically made from amino acids. They have amino groups in them, but no longer have carboxyl groups and are therefore no longer classified as amino acids. Serotonin is made from tryptophan. It is the basis of the serotonergic system, which has its own specific receptors. Serotonin is transported back into the presynaptic cell for repackaging.

Other biogenic amines are made from tyrosine, and include dopamine, norepinephrine, and epinephrine. Dopamine is part of its own system, the dopaminergic system, which has dopamine receptors. Dopamine is removed from the synapse by transport proteins in the presynaptic cell membrane. Norepinephrine and epinephrine belong to the adrenergic neurotransmitter system. The two molecules are very similar and bind to the same receptors, which are referred to as alpha and beta receptors. Norepinephrine and epinephrine are also transported back into the presynaptic cell. The chemical epinephrine (epi- = “on”; “-nephrine” = kidney) is also known as adrenaline (renal = “kidney”), and norepinephrine is sometimes referred to as noradrenaline. The adrenal gland produces epinephrine and norepinephrine to be released into the blood stream as hormones.

A neuropeptide is a neurotransmitter molecule made up of chains of amino acids connected by peptide bonds. This is what a protein is, but the term protein implies a certain length to the molecule. Some neuropeptides are quite short, such as met-enkephalin, which is five amino acids long. Others are long, such as beta-endorphin, which is 31 amino acids long. Neuropeptides are often released at synapses in combination with another neurotransmitter, and they often act as hormones in other systems of the body, such as vasoactive intestinal peptide (VIP) or substance P.

The effect of a neurotransmitter on the postsynaptic element is entirely dependent on the receptor protein. First, if there is no receptor protein in the membrane of the postsynaptic element, then the neurotransmitter has no effect. The depolarizing or hyperpolarizing effect is also dependent on the receptor. When acetylcholine binds to the nicotinic receptor, the postsynaptic cell is depolarized. This is because the receptor is a cation channel and positively charged Na+ will rush into the cell. However, when acetylcholine binds to the muscarinic receptor, of which there are several variants, it might cause depolarization or hyperpolarization of the target cell.

The amino acid neurotransmitters, glutamate, glycine, and GABA, are almost exclusively associated with just one effect. Glutamate is considered an excitatory amino acid, but only because Glu receptors in the adult cause depolarization of the postsynaptic cell. Glycine and GABA are considered inhibitory amino acids, again because their receptors cause hyperpolarization.

The biogenic amines have mixed effects. For example, the dopamine receptors that are classified as D1 receptors are excitatory whereas D2-type receptors are inhibitory. Biogenic amine receptors and neuropeptide receptors can have even more complex effects because some may not directly affect the membrane potential, but rather have an effect on gene transcription or other metabolic processes in the neuron. The characteristics of the various neurotransmitter systems presented in this section are organized in Table 3.

The important thing to remember about neurotransmitters, and signaling chemicals in general, is that the effect is entirely dependent on the receptor. Neurotransmitters bind to one of two classes of receptors at the cell surface, ionotropic or metabotropic (Figure 4). Ionotropic receptors are ligand-gated ion channels, such as the nicotinic receptor for acetylcholine or the glycine receptor. A metabotropic receptor involves a complex of proteins that result in metabolic changes within the cell. The receptor complex includes the transmembrane receptor protein, a G protein, and an effector protein. The neurotransmitter, referred to as the first messenger, binds to the receptor protein on the extracellular surface of the cell, and the intracellular side of the protein initiates activity of the G protein. The G protein is a guanosine triphosphate (GTP) hydrolase that physically moves from the receptor protein to the effector protein to activate the latter. An effector protein is an enzyme that catalyzes the generation of a new molecule, which acts as the intracellular mediator of the signal that binds to the receptor. This intracellular mediator is called the second messenger.

Different receptors use different second messengers. Two common examples of second messengers are cyclic adenosine monophosphate (cAMP) and inositol triphosphate (IP3). The enzyme adenylate cyclase (an example of an effector protein) makes cAMP, and phospholipase C is the enzyme that makes IP3. Second messengers, after they are produced by the effector protein, cause metabolic changes within the cell. These changes are most likely the activation of other enzymes in the cell. In neurons, they often modify ion channels, either opening or closing them. These enzymes can also cause changes in the cell, such as the activation of genes in the nucleus, and therefore the increased synthesis of proteins. In neurons, these kinds of changes are often the basis of stronger connections between cells at the synapse and may be the basis of learning and memory.

[caption id="" align="aligncenter" width="450"]This diagram contains two images, labeled A and B. Both images show a cross section of a postsynaptic membrane. There are two proteins embedded in each of the two membrane cross sections. In diagram A, direct activation brings about an immediate response. Here, both of the membrane proteins are ion channels. Several hexagonal neurotransmitters bind to ionotropic receptors on the extracellular fluid side of the channels. The binding of neurotransmitters causes the channels to open, allowing ions to flow from the extracellular fluid into the cytosol. Image B shows indirect activation, which involves a prolonged response, amplified over time. Here, one of the cell membrane proteins is solid while the other is a channel. Neurotransmitters bind to metabotropic receptors on the extracellular side of the solid protein. This triggers the solid protein to activate a G protein in the cytoplasm. The G protein binds to an effector protein in the cytoplasm, which results in the production of several second messenger particles. The second messenger activates enzymes that open the channel protein, allowing ions to enter the cytoplasm. Figure 4. Receptor Types. (a) An ionotropic receptor is a channel that opens when the neurotransmitter binds to it. (b) A metabotropic receptor is a complex that causes metabolic changes in the cell when the neurotransmitter binds to it (1). After binding, the G protein hydrolyzes GTP and moves to the effector protein (2). When the G protein contacts the effector protein, a second messenger is generated, such as cAMP (3). The second messenger can then go on to cause changes in the neuron, such as opening or closing ion channels, metabolic changes, and changes in gene transcription.[/caption]
Characteristics of Neurotransmitter Systems (Table 3)
System Cholinergic Amino acids Biogenic amines Neuropeptides
Neurotransmitters Acetylcholine Glutamate, glycine, GABA Serotonin (5-HT), dopamine, norepinephrine, (epinephrine) Met-enkephalin, beta-endorphin, VIP, Substance P, etc.
Receptors Nicotinic and muscarinic receptors Glu receptors, gly receptors, GABA receptors 5-HT receptors, D1 and D2 receptors, α-adrenergic and β-adrenergic receptors Receptors are too numerous to list, but are specific to the peptides.
Elimination Degradation by acetylcholinesterase Reuptake by neurons or glia Reuptake by neurons Degradation by enzymes called peptidases
Postsynaptic effect Nicotinic receptor causes depolarization. Muscarinic receptors can cause both depolarization or hyperpolarization depending on the subtype. Glu receptors cause depolarization. Gly and GABA receptors cause hyperpolarization. Depolarization or hyperpolarization depends on the specific receptor. For example, D1 receptors cause depolarization and D2 receptors cause hyperpolarization. Depolarization or hyperpolarization depends on the specific receptor.
Disorders of the…
Nervous SystemThe underlying cause of some neurodegenerative diseases, such as Alzheimer’s and Parkinson’s, appears to be related to proteins—specifically, to proteins behaving badly. One of the strongest theories of what causes Alzheimer’s disease is based on the accumulation of beta-amyloid plaques, dense conglomerations of a protein that is not functioning correctly. Parkinson’s disease is linked to an increase in a protein known as alpha-synuclein that is toxic to the cells of the substantia nigra nucleus in the midbrain. For proteins to function correctly, they are dependent on their three-dimensional shape. The linear sequence of amino acids folds into a three-dimensional shape that is based on the interactions between and among those amino acids. When the folding is disturbed, and proteins take on a different shape, they stop functioning correctly. But the disease is not necessarily the result of functional loss of these proteins; rather, these altered proteins start to accumulate and may become toxic. For example, in Alzheimer’s, the hallmark of the disease is the accumulation of these amyloid plaques in the cerebral cortex. The term coined to describe this sort of disease is “proteopathy” and it includes other diseases. Creutzfeld-Jacob disease, the human variant of the prion disease known as mad cow disease in the bovine, also involves the accumulation of amyloid plaques, similar to Alzheimer’s. Diseases of other organ systems can fall into this group as well, such as cystic fibrosis or type 2 diabetes. Recognizing the relationship between these diseases has suggested new therapeutic possibilities. Interfering with the accumulation of the proteins, and possibly as early as their original production within the cell, may unlock new ways to alleviate these devastating diseases.
]]> 2505 2477 6 0 12.3 The Function of Nervous Tissue https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2509 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2509

Learning Objectives

By the end of this section, you will be able to:
  • Distinguish the major functions of the nervous system: sensation, integration, and response
  • List the sequence of events in a simple sensory receptor–motor response pathway

Having looked at the components of nervous tissue, and the basic anatomy of the nervous system, next comes an understanding of how nervous tissue is capable of communicating within the nervous system. Before getting to the nuts and bolts of how this works, an illustration of how the components come together will be helpful. An example is summarized in Figure 1.

[caption id="" align="aligncenter" width="450"]This diagram shows the complete pathway a nerve impulse takes when a person tests the temperature of shower water with their hand. First, a sensory nerve ending in the index finger sends a nerve impulse to the spinal cord. A cross section of one segment of the vertebrae is shown from a superior view. The sensory nerve connected to the nerve ending is located in the dorsal root ganglion. The nerve ending is a dendrite of the sensory neuron, as it also has an axon that synapses with an interneuron. The interneuron then synapses with a second interneuron in the thalamus. This second interneuron synapses with brain tissue in the cerebral cortex, allowing conscious perception of the water temperature. The brain then initiates a motor command by stimulating an upper motor neuron in the cerebral cortex. The axon of the upper motor neuron extends all the way to the spinal cord, where it synapses with a lower motor neuron in the gray matter of the spinal cord. The impulse then travels down the lower motor neuron back to the hand where it synapses with the skeletal muscles of the hand. This triggers the muscle contractions that turn the dials of the shower to adjust the water temperature. Figure 1. Testing the Water. (1) The sensory neuron has endings in the skin that sense a stimulus such as water temperature. The strength of the signal that starts here is dependent on the strength of the stimulus. (2) The graded potential from the sensory endings, if strong enough, will initiate an action potential at the initial segment of the axon (which is immediately adjacent to the sensory endings in the skin). (3) The axon of the peripheral sensory neuron enters the spinal cord and contacts another neuron in the gray matter. The contact is a synapse where another graded potential is caused by the release of a chemical signal from the axon terminals. (4) An action potential is initiated at the initial segment of this neuron and travels up the sensory pathway to a region of the brain called the thalamus. Another synapse passes the information along to the next neuron. (5) The sensory pathway ends when the signal reaches the cerebral cortex. (6) After integration with neurons in other parts of the cerebral cortex, a motor command is sent from the precentral gyrus of the frontal cortex. (7) The upper motor neuron sends an action potential down to the spinal cord. The target of the upper motor neuron is the dendrites of the lower motor neuron in the gray matter of the spinal cord. (8) The axon of the lower motor neuron emerges from the spinal cord in a nerve and connects to a muscle through a neuromuscular junction to cause contraction of the target muscle.[/caption]

Imagine you are about to take a shower in the morning before going to school. You have turned on the faucet to start the water as you prepare to get in the shower. After a few minutes, you expect the water to be a temperature that will be comfortable to enter. So you put your hand out into the spray of water. What happens next depends on how your nervous system interacts with the stimulus of the water temperature and what you do in response to that stimulus.

Found in the skin of your fingers or toes is a type of sensory receptor that is sensitive to temperature, called a thermoreceptor. When you place your hand under the shower (Figure 2), the cell membrane of the thermoreceptors changes its electrical state (voltage). The amount of change is dependent on the strength of the stimulus (how hot the water is). This is called a graded potential. If the stimulus is strong, the voltage of the cell membrane will change enough to generate an electrical signal that will travel down the axon. You have learned about this type of signaling before, with respect to the interaction of nerves and muscles at the neuromuscular junction. The voltage at which such a signal is generated is called the threshold, and the resulting electrical signal is called an action potential. In this example, the action potential travels—a process known as propagation—along the axon from the axon hillock to the axon terminals and into the synaptic end bulbs. When this signal reaches the end bulbs, it causes the release of a signaling molecule called a neurotransmitter.
[caption id="" align="aligncenter" width="380"]This diagram shows the first step of the previous figure. A hand is placed under flowing water, causing a sensory receptor in the index finger to send a nerve impulse down the arm, to the spinal cord. Figure 2. The Sensory Input. Receptors in the skin sense the temperature of the water.[/caption]

The neurotransmitter diffuses across the short distance of the synapse and binds to a receptor protein of the target neuron. When the molecular signal binds to the receptor, the cell membrane of the target neuron changes its electrical state and a new graded potential begins. If that graded potential is strong enough to reach threshold, the second neuron generates an action potential at its axon hillock. The target of this neuron is another neuron in the thalamus of the brain, the part of the CNS that acts as a relay for sensory information. At another synapse, neurotransmitter is released and binds to its receptor. The thalamus then sends the sensory information to the cerebral cortex, the outermost layer of gray matter in the brain, where conscious perception of that water temperature begins.

Within the cerebral cortex, information is processed among many neurons, integrating the stimulus of the water temperature with other sensory stimuli, with your emotional state (you just aren't ready to wake up; the bed is calling to you), memories (perhaps of the lab notes you have to study before a quiz). Finally, a plan is developed about what to do, whether that is to turn the temperature up, turn the whole shower off and go back to bed, or step into the shower. To do any of these things, the cerebral cortex has to send a command out to your body to move muscles (Figure 3).
[caption id="" align="aligncenter" width="380"]This diagram shows the later steps of Figure 12.13. A hand is placed under flowing water. The axon of a motor neuron travels down the forearm and then branches as it reaches the hand. Each branch synapses with a different skeletal muscle in the hand. The synapse between the axon branches and the muscle is a neuromuscular junction. An impulse travelling down the motor neuron will cause the skeletal muscles to contract, resulting in muscle movement. In this case, the movement results in the person adjusting the faucet dials to change the temperature of the water. Figure 3. The Motor Response. On the basis of the sensory input and the integration in the CNS, a motor response is formulated and executed.[/caption]
A region of the cortex is specialized for sending signals down to the spinal cord for movement. The upper motor neuron is in this region, called the precentral gyrus of the frontal cortex, which has an axon that extends all the way down the spinal cord. At the level of the spinal cord at which this axon makes a synapse, a graded potential occurs in the cell membrane of a lower motor neuron. This second motor neuron is responsible for causing muscle fibers to contract. In the manner described in the chapter on muscle tissue, an action potential travels along the motor neuron axon into the periphery. The axon terminates on muscle fibers at the neuromuscular junction. Acetylcholine is released at this specialized synapse, which causes the muscle action potential to begin, following a large potential known as an end plate potential. When the lower motor neuron excites the muscle fiber, it contracts. All of this occurs in a fraction of a second, but this story is the basis of how the nervous system functions.
Career Connections

Neurophysiologist Understanding how the nervous system works could be a driving force in your career. Studying neurophysiology is a very rewarding path to follow. It means that there is a lot of work to do, but the rewards are worth the effort.

The career path of a research scientist can be straightforward: college, graduate school, postdoctoral research, academic research position at a university. A Bachelor’s degree in science will get you started, and for neurophysiology that might be in biology, psychology, computer science, engineering, or neuroscience. But the real specialization comes in graduate school. There are many different programs out there to study the nervous system, not just neuroscience itself. Most graduate programs are doctoral, meaning that a Master’s degree is not part of the work. These are usually considered five-year programs, with the first two years dedicated to course work and finding a research mentor, and the last three years dedicated to finding a research topic and pursuing that with a near single-mindedness. The research will usually result in a few publications in scientific journals, which will make up the bulk of a doctoral dissertation. After graduating with a Ph.D., researchers will go on to find specialized work called a postdoctoral fellowship within established labs. In this position, a researcher starts to establish their own research career with the hopes of finding an academic position at a research university.

Other options are available if you are interested in how the nervous system works. Especially for neurophysiology, a medical degree might be more suitable so you can learn about the clinical applications of neurophysiology and possibly work with human subjects. An academic career is not a necessity. Biotechnology firms are eager to find motivated scientists ready to tackle the tough questions about how the nervous system works so that therapeutic chemicals can be tested on some of the most challenging disorders such as Alzheimer’s disease or Parkinson’s disease, or spinal cord injury.

Others with a medical degree and a specialization in neuroscience go on to work directly with patients, diagnosing and treating mental disorders. You can do this as a psychiatrist, a neuropsychologist, a neuroscience nurse, or a neurodiagnostic technician, among other possible career paths.

]]> 2509 2477 4 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2511 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2511 [caption id="" align="aligncenter" width="500"]This photo shows a person playing foosball. The person has had both of their lower arms amputated. The left arm was replaced with a replica of a human hand and the right arm was replaced with a manipulator that resembles a pair of tongs. Figure 1. Robotic Arms Playing Foosball. As the neural circuitry of the nervous system has become more fully understood and robotics more sophisticated, it is now possible to integrate technology with the body and restore abilities following traumatic events. At some point in the future, will this type of technology lead to the ability to augment our nervous systems? (credit: U.S. Army/Wikimedia Commons)[/caption]

Chapter Objectives

After studying this chapter, you will be able to:
  • Name the major divisions of the nervous system, both anatomical and functional
  • Describe the functional and structural differences between gray matter and white matter structures
  • Name the parts of the multipolar neuron in order of polarity
  • List the types of glial cells and assign each to the proper division of the nervous system, along with their function(s)
  • Distinguish the major functions of the nervous system: sensation, integration, and response
  • Describe the components of the membrane that establish the resting membrane potential
  • Describe the changes that occur to the membrane that result in the action potential
  • Explain the differences between types of graded potentials
  • Categorize the major neurotransmitters by chemical type and effect
The nervous system is a very complex organ system. In Peter D. Kramer’s book Listening to Prozac, a pharmaceutical researcher is quoted as saying, “If the human brain were simple enough for us to understand, we would be too simple to understand it” (1994). That quote is from the early 1990s; in the two decades since, progress has continued at an amazing rate within the scientific disciplines of neuroscience. It is an interesting conundrum to consider that the complexity of the nervous system may be too complex for it (that is, for us) to completely unravel. But our current level of understanding is probably nowhere close to that limit. One easy way to begin to understand the structure of the nervous system is to start with the large divisions and work through to a more in-depth understanding. In other chapters, the finer details of the nervous system will be explained, but first looking at an overview of the system will allow you to begin to understand how its parts work together. The focus of this chapter is on nervous (neural) tissue, both its structure and its function. But before you learn about that, you will see a big picture of the system—actually, a few big pictures.]]> 2511 2477 1 0 13.3 Circulation and the Central Nervous System https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2527 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2527

Learning Objectives

By the end of this section, you will be able to:
  • Describe the vessels that supply the CNS with blood
  • Name the components of the ventricular system and the regions of the brain in which each is located
  • Explain the production of cerebrospinal fluid and its flow through the ventricles
  • Explain how a disruption in circulation would result in a stroke

The CNS is crucial to the operation of the body, and any compromise in the brain and spinal cord can lead to severe difficulties. The CNS has a privileged blood supply, as suggested by the blood-brain barrier. The function of the tissue in the CNS is crucial to the survival of the organism, so the contents of the blood cannot simply pass into the central nervous tissue. To protect this region from the toxins and pathogens that may be traveling through the blood stream, there is strict control over what can move out of the general systems and into the brain and spinal cord. Because of this privilege, the CNS needs specialized structures for the maintenance of circulation. This begins with a unique arrangement of blood vessels carrying fresh blood into the CNS. Beyond the supply of blood, the CNS filters that blood into cerebrospinal fluid (CSF), which is then circulated through the cavities of the brain and spinal cord called ventricles.

Blood Supply to the Brain

A lack of oxygen to the CNS can be devastating, and the cardiovascular system has specific regulatory reflexes to ensure that the blood supply is not interrupted. There are multiple routes for blood to get into the CNS, with specializations to protect that blood supply and to maximize the ability of the brain to get an uninterrupted perfusion.

Arterial Supply

The major artery carrying recently oxygenated blood away from the heart is the aorta. The very first branches off the aorta supply the heart with nutrients and oxygen. The next branches give rise to the common carotid arteries, which further branch into the internal carotid arteries. The external carotid arteries supply blood to the tissues on the surface of the cranium. The bases of the common carotids contain stretch receptors that immediately respond to the drop in blood pressure upon standing. The orthostatic reflex is a reaction to this change in body position, so that blood pressure is maintained against the increasing effect of gravity (orthostatic means “standing up”). Heart rate increases—a reflex of the sympathetic division of the autonomic nervous system—and this raises blood pressure.

The internal carotid artery enters the cranium through the carotid canal in the temporal bone. A second set of vessels that supply the CNS are the vertebral arteries, which are protected as they pass through the neck region by the transverse foramina of the cervical vertebrae. The vertebral arteries enter the cranium through the foramen magnum of the occipital bone. Branches off the left and right vertebral arteries merge into the anterior spinal artery supplying the anterior aspect of the spinal cord, found along the anterior median fissure. The two vertebral arteries then merge into the basilar artery, which gives rise to branches to the brain stem and cerebellum. The left and right internal carotid arteries and branches of the basilar artery all become the circle of Willis, a confluence of arteries that can maintain perfusion of the brain even if narrowing or a blockage limits flow through one part (Figure 1).

[caption id="" align="aligncenter" width="320"]This diagram shows a series of interconnected blood vessels and capillaries. Figure 1. Circle of Willis. The blood supply to the brain enters through the internal carotid arteries and the vertebral arteries, eventually giving rise to the circle of Willis.[/caption]

Venous Return

After passing through the CNS, blood returns to the circulation through a series of dural sinuses and veins (Figure 2). The superior sagittal sinus runs in the groove of the longitudinal fissure, where it absorbs CSF from the meninges. The superior sagittal sinus drains to the confluence of sinuses, along with the occipital sinuses and straight sinus, to then drain into the transverse sinuses. The transverse sinuses connect to the sigmoid sinuses, which then connect to the jugular veins. From there, the blood continues toward the heart to be pumped to the lungs for reoxygenation.
[caption id="" align="aligncenter" width="550"]This diagram shows a lateral view of the brain and labels the location of the different sinuses. Figure 2. Dural Sinuses and Veins. Blood drains from the brain through a series of sinuses that connect to the jugular veins.[/caption]

Protective Coverings of the Brain and Spinal Cord

The outer surface of the CNS is covered by a series of membranes composed of connective tissue called the meninges, which protect the brain. The dura mater is a thick fibrous layer and a strong protective sheath over the entire brain and spinal cord. It is anchored to the inner surface of the cranium and vertebral cavity. The arachnoid mater is a membrane of thin fibrous tissue that forms a loose sac around the CNS. Beneath the arachnoid is a thin, filamentous mesh called the arachnoid trabeculae, which looks like a spider web, giving this layer its name. Directly adjacent to the surface of the CNS is the pia mater, a thin fibrous membrane that follows the convolutions of gyri and sulci in the cerebral cortex and fits into other grooves and indentations (Figure 3).
[caption id="" align="aligncenter" width="500"]This image shows a cross-section through the brain. The different meningeal layers are labeled. Figure 3. Meningeal Layers of Superior Sagittal Sinus. The layers of the meninges in the longitudinal fissure of the superior sagittal sinus are shown, with the dura mater adjacent to the inner surface of the cranium, the pia mater adjacent to the surface of the brain, and the arachnoid and subarachnoid space between them. An arachnoid villus is shown emerging into the dural sinus to allow CSF to filter back into the blood for drainage.[/caption]

Dura Mater

Like a thick cap covering the brain, the dura mater is a tough outer covering. The name comes from the Latin for “tough mother” to represent its physically protective role. It encloses the entire CNS and the major blood vessels that enter the cranium and vertebral cavity. It is directly attached to the inner surface of the bones of the cranium and to the very end of the vertebral cavity.

There are infoldings of the dura that fit into large crevasses of the brain. Two infoldings go through the midline separations of the cerebrum and cerebellum; one forms a shelf-like tent between the occipital lobes of the cerebrum and the cerebellum, and the other surrounds the pituitary gland. The dura also surrounds and supports the venous sinuses.

Arachnoid Mater

The middle layer of the meninges is the arachnoid, named for the spider-web–like trabeculae between it and the pia mater. The arachnoid defines a sac-like enclosure around the CNS. The trabeculae are found in the subarachnoid space, which is filled with circulating CSF. The arachnoid emerges into the dural sinuses as the arachnoid granulations, where the CSF is filtered back into the blood for drainage from the nervous system.

The subarachnoid space is filled with circulating CSF, which also provides a liquid cushion to the brain and spinal cord. Similar to clinical blood work, a sample of CSF can be withdrawn to find chemical evidence of neuropathology or metabolic traces of the biochemical functions of nervous tissue.

Pia Mater

The outer surface of the CNS is covered in the thin fibrous membrane of the pia mater. It is thought to have a continuous layer of cells providing a fluid-impermeable membrane. The name pia mater comes from the Latin for “tender mother,” suggesting the thin membrane is a gentle covering for the brain. The pia extends into every convolution of the CNS, lining the inside of the sulci in the cerebral and cerebellar cortices. At the end of the spinal cord, a thin filament extends from the inferior end of CNS at the upper lumbar region of the vertebral column to the sacral end of the vertebral column. Because the spinal cord does not extend through the lower lumbar region of the vertebral column, a needle can be inserted through the dura and arachnoid layers to withdraw CSF. This procedure is called a lumbar puncture and avoids the risk of damaging the central tissue of the spinal cord. Blood vessels that are nourishing the central nervous tissue are between the pia mater and the nervous tissue.

Disorders of the…
Meninges Meningitis is an inflammation of the meninges, the three layers of fibrous membrane that surround the CNS. Meningitis can be caused by infection by bacteria or viruses. The particular pathogens are not special to meningitis; it is just an inflammation of that specific set of tissues from what might be a broader infection. Bacterial meningitis can be caused by Streptococcus, Staphylococcus, or the tuberculosis pathogen, among many others. Viral meningitis is usually the result of common enteroviruses (such as those that cause intestinal disorders), but may be the result of the herpes virus or West Nile virus. Bacterial meningitis tends to be more severe. The symptoms associated with meningitis can be fever, chills, nausea, vomiting, light sensitivity, soreness of the neck, or severe headache. More important are the neurological symptoms, such as changes in mental state (confusion, memory deficits, and other dementia-type symptoms). A serious risk of meningitis can be damage to peripheral structures because of the nerves that pass through the meninges. Hearing loss is a common result of meningitis. The primary test for meningitis is a lumbar puncture. A needle inserted into the lumbar region of the spinal column through the dura mater and arachnoid membrane into the subarachnoid space can be used to withdraw the fluid for chemical testing. Fatality occurs in 5 to 40 percent of children and 20 to 50 percent of adults with bacterial meningitis. Treatment of bacterial meningitis is through antibiotics, but viral meningitis cannot be treated with antibiotics because viruses do not respond to that type of drug. Fortunately, the viral forms are milder.

The Ventricular System

Cerebrospinal fluid (CSF) circulates throughout and around the CNS. In other tissues, water and small molecules are filtered through capillaries as the major contributor to the interstitial fluid. In the brain, CSF is produced in special structures to perfuse through the nervous tissue of the CNS and is continuous with the interstitial fluid. Specifically, CSF circulates to remove metabolic wastes from the interstitial fluids of nervous tissues and return them to the blood stream. The ventricles are the open spaces within the brain where CSF circulates. In some of these spaces, CSF is produced by filtering of the blood that is performed by a specialized membrane known as a choroid plexus. The CSF circulates through all of the ventricles to eventually emerge into the subarachnoid space where it will be reabsorbed into the blood.

The Ventricles

There are four ventricles within the brain, all of which developed from the original hollow space within the neural tube, the central canal. The first two are named the lateral ventricles and are deep within the cerebrum. These ventricles are connected to the third ventricle by two openings called the interventricular foramina. The third ventricle is the space between the left and right sides of the diencephalon, which opens into the cerebral aqueduct that passes through the midbrain. The aqueduct opens into the fourth ventricle, which is the space between the cerebellum and the pons and upper medulla (Figure 4).
[caption id="" align="aligncenter" width="550"]This diagram shows the cross section of the brain and the major parts are labeled. Arrows on the figure show the direction of circulation of the cerebro-spinal fluid. Figure 4. Cerebrospinal Fluid Circulation. The choroid plexus in the four ventricles produce CSF, which is circulated through the ventricular system and then enters the subarachnoid space through the median and lateral apertures. The CSF is then reabsorbed into the blood at the arachnoid granulations, where the arachnoid membrane emerges into the dural sinuses.[/caption]

As the telencephalon enlarges and grows into the cranial cavity, it is limited by the space within the skull. The telencephalon is the most anterior region of what was the neural tube, but cannot grow past the limit of the frontal bone of the skull. Because the cerebrum fits into this space, it takes on a C-shaped formation, through the frontal, parietal, occipital, and finally temporal regions. The space within the telencephalon is stretched into this same C-shape. The two ventricles are in the left and right sides, and were at one time referred to as the first and second ventricles. The interventricular foramina connect the frontal region of the lateral ventricles with the third ventricle.

The third ventricle is the space bounded by the medial walls of the hypothalamus and thalamus. The two thalami touch in the center in most brains as the massa intermedia, which is surrounded by the third ventricle. The cerebral aqueduct opens just inferior to the epithalamus and passes through the midbrain. The tectum and tegmentum of the midbrain are the roof and floor of the cerebral aqueduct, respectively. The aqueduct opens up into the fourth ventricle. The floor of the fourth ventricle is the dorsal surface of the pons and upper medulla (that gray matter making a continuation of the tegmentum of the midbrain). The fourth ventricle then narrows into the central canal of the spinal cord.

The ventricular system opens up to the subarachnoid space from the fourth ventricle. The single median aperture and the pair of lateral apertures connect to the subarachnoid space so that CSF can flow through the ventricles and around the outside of the CNS. Cerebrospinal fluid is produced within the ventricles by a type of specialized membrane called a choroid plexus. Ependymal cells (one of the types of glial cells described in the introduction to the nervous system) surround blood capillaries and filter the blood to make CSF. The fluid is a clear solution with a limited amount of the constituents of blood. It is essentially water, small molecules, and electrolytes. Oxygen and carbon dioxide are dissolved into the CSF, as they are in blood, and can diffuse between the fluid and the nervous tissue.

Cerebrospinal Fluid Circulation

The choroid plexuses are found in all four ventricles. Observed in dissection, they appear as soft, fuzzy structures that may still be pink, depending on how well the circulatory system is cleared in preparation of the tissue. The CSF is produced from components extracted from the blood, so its flow out of the ventricles is tied to the pulse of cardiovascular circulation.

From the lateral ventricles, the CSF flows into the third ventricle, where more CSF is produced, and then through the cerebral aqueduct into the fourth ventricle where even more CSF is produced. A very small amount of CSF is filtered at any one of the plexuses, for a total of about 500 milliliters daily, but it is continuously made and pulses through the ventricular system, keeping the fluid moving. From the fourth ventricle, CSF can continue down the central canal of the spinal cord, but this is essentially a cul-de-sac, so more of the fluid leaves the ventricular system and moves into the subarachnoid space through the median and lateral apertures.

Within the subarachnoid space, the CSF flows around all of the CNS, providing two important functions. As with elsewhere in its circulation, the CSF picks up metabolic wastes from the nervous tissue and moves it out of the CNS. It also acts as a liquid cushion for the brain and spinal cord. By surrounding the entire system in the subarachnoid space, it provides a thin buffer around the organs within the strong, protective dura mater. The arachnoid granulations are outpocketings of the arachnoid membrane into the dural sinuses so that CSF can be reabsorbed into the blood, along with the metabolic wastes. From the dural sinuses, blood drains out of the head and neck through the jugular veins, along with the rest of the circulation for blood, to be reoxygenated by the lungs and wastes to be filtered out by the kidneys (Table 2).

Components of CSF Circulation (Table 2)
Lateral ventricles Third ventricle Cerebral aqueduct Fourth ventricle Central canal Subarachnoid space
Location in CNS Cerebrum Diencephalon Midbrain Between pons/upper medulla and cerebellum Spinal cord External to entire CNS
Blood vessel structure Choroid plexus Choroid plexus None Choroid plexus None Arachnoid granulations
]]> 2527 2512 2 0 17.2 Hormones https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2593 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2593

Learning Objectives

By the end of this section, you will be able to:
  • Identify the three major classes of hormones on the basis of chemical structure
  • Compare and contrast intracellular and cell membrane hormone receptors
  • Describe signaling pathways that involve cAMP and IP3
  • Identify several factors that influence a target cell’s response
  • Discuss the role of feedback loops and humoral, hormonal, and neural stimuli in hormone control

Although a given hormone may travel throughout the body in the bloodstream, it will affect the activity only of its target cells; that is, cells with receptors for that particular hormone. Once the hormone binds to the receptor, a chain of events is initiated that leads to the target cell’s response. Hormones play a critical role in the regulation of physiological processes because of the target cell responses they regulate. These responses contribute to human reproduction, growth and development of body tissues, metabolism, fluid, and electrolyte balance, sleep, and many other body functions. The major hormones of the human body and their effects are identified in Table 2.

Endocrine Glands and Their Major Hormones (Table 2)
Endocrine gland Associated hormones Chemical class Effect
Pituitary (anterior) Growth hormone (GH) Protein Promotes growth of body tissues
Pituitary (anterior) Prolactin (PRL) Peptide Promotes milk production
Pituitary (anterior) Thyroid-stimulating hormone (TSH) Glycoprotein Stimulates thyroid hormone release
Pituitary (anterior) Adrenocorticotropic hormone (ACTH) Peptide Stimulates hormone release by adrenal cortex
Pituitary (anterior) Follicle-stimulating hormone (FSH) Glycoprotein Stimulates gamete production
Pituitary (anterior) Luteinizing hormone (LH) Glycoprotein Stimulates androgen production by gonads
Pituitary (posterior) Antidiuretic hormone (ADH) Peptide Stimulates water reabsorption by kidneys
Pituitary (posterior) Oxytocin Peptide Stimulates uterine contractions during childbirth
Thyroid Thyroxine (T4), triiodothyronine (T3) Amine Stimulate basal metabolic rate
Thyroid Calcitonin Peptide Reduces blood Ca2+ levels
Parathyroid Parathyroid hormone (PTH) Peptide Increases blood Ca2+ levels
Adrenal (cortex) Aldosterone Steroid Increases blood Na+ levels
Adrenal (cortex) Cortisol, corticosterone, cortisone Steroid Increase blood glucose levels
Adrenal (medulla) Epinephrine, norepinephrine Amine Stimulate fight-or-flight response
Pineal Melatonin Amine Regulates sleep cycles
Pancreas Insulin Protein Reduces blood glucose levels
Pancreas Glucagon Protein Increases blood glucose levels
Testes Testosterone Steroid Stimulates development of male secondary sex characteristics and sperm production
Ovaries Estrogens and progesterone Steroid Stimulate development of female secondary sex characteristics and prepare the body for childbirth

Types of Hormones

The hormones of the human body can be divided into two major groups on the basis of their chemical structure. Hormones derived from amino acids include amines, peptides, and proteins. Those derived from lipids include steroids (Figure 1). These chemical groups affect a hormone’s distribution, the type of receptors it binds to, and other aspects of its function.

[caption id="" align="aligncenter" width="550"]This table shows the chemical structure of amine hormones, peptide hormones, protein hormones, and steroid hormones. Amine hormones are amino acids with modified side groups. The example given is norepinephrine, which contains the NH two group typical of an amino acid, along with a hydroxyl (OH) group. The carboxyl group typical of most amino acids is replaced with a benzene ring, depicted as a hexagon of carbons that are connected by alternating single and double bonds. Peptide hormones are composed of short chains of amino acids. The example given is oxytocin, which has a chain of the following amino acids: GLY, LEU, PRO. The PRO is the bottom of the chain, which connects to a ring of the following amino acids: CYS, CYS, TYR, ILE, GLU, and ASP. Protein hormones are composed of long chains of linked amino acids. The example given is human growth hormone, which is composed of a bundle of amino acid strands, some thread-like, some coiled, and some in flat, folded sheets. Finally, steroid hormones are derived from the lipid cholesterol. Testosterone and progesterone are given as examples, which each contain several hexagonal and pentagonal carbon rings linked together. Figure 1. Amine, Peptide, Protein, and Steroid Hormone Structure[/caption]

Amine Hormones

Hormones derived from the modification of amino acids are referred to as amine hormones. Typically, the original structure of the amino acid is modified such that a –COOH, or carboxyl, group is removed, whereas the −NH3+−NH3+, or amine, group remains.

Amine hormones are synthesized from the amino acids tryptophan or tyrosine. An example of a hormone derived from tryptophan is melatonin, which is secreted by the pineal gland and helps regulate circadian rhythm. Tyrosine derivatives include the metabolism-regulating thyroid hormones, as well as the catecholamines, such as epinephrine, norepinephrine, and dopamine. Epinephrine and norepinephrine are secreted by the adrenal medulla and play a role in the fight-or-flight response, whereas dopamine is secreted by the hypothalamus and inhibits the release of certain anterior pituitary hormones.

Peptide and Protein Hormones

Whereas the amine hormones are derived from a single amino acid, peptide and protein hormones consist of multiple amino acids that link to form an amino acid chain. Peptide hormones consist of short chains of amino acids, whereas protein hormones are longer polypeptides. Both types are synthesized like other body proteins: DNA is transcribed into mRNA, which is translated into an amino acid chain.

Examples of peptide hormones include antidiuretic hormone (ADH), a pituitary hormone important in fluid balance, and atrial-natriuretic peptide, which is produced by the heart and helps to decrease blood pressure. Some examples of protein hormones include growth hormone, which is produced by the pituitary gland, and follicle-stimulating hormone (FSH), which has an attached carbohydrate group and is thus classified as a glycoprotein. FSH helps stimulate the maturation of eggs in the ovaries and sperm in the testes.

Steroid Hormones

The primary hormones derived from lipids are steroids. Steroid hormones are derived from the lipid cholesterol. For example, the reproductive hormones testosterone and the estrogens—which are produced by the gonads (testes and ovaries)—are steroid hormones. The adrenal glands produce the steroid hormone aldosterone, which is involved in osmoregulation, and cortisol, which plays a role in metabolism.

Like cholesterol, steroid hormones are not soluble in water (they are hydrophobic). Because blood is water-based, lipid-derived hormones must travel to their target cell bound to a transport protein. This more complex structure extends the half-life of steroid hormones much longer than that of hormones derived from amino acids. A hormone’s half-life is the time required for half the concentration of the hormone to be degraded. For example, the lipid-derived hormone cortisol has a half-life of approximately 60 to 90 minutes. In contrast, the amino acid–derived hormone epinephrine has a half-life of approximately one minute.

Pathways of Hormone Action

The message a hormone sends is received by a hormone receptor, a protein located either inside the cell or within the cell membrane. The receptor will process the message by initiating other signaling events or cellular mechanisms that result in the target cell’s response. Hormone receptors recognize molecules with specific shapes and side groups, and respond only to those hormones that are recognized. The same type of receptor may be located on cells in different body tissues, and trigger somewhat different responses. Thus, the response triggered by a hormone depends not only on the hormone, but also on the target cell.

Once the target cell receives the hormone signal, it can respond in a variety of ways. The response may include the stimulation of protein synthesis, activation or deactivation of enzymes, alteration in the permeability of the cell membrane, altered rates of mitosis and cell growth, and stimulation of the secretion of products. Moreover, a single hormone may be capable of inducing different responses in a given cell.

Pathways Involving Intracellular Hormone Receptors

Intracellular hormone receptors are located inside the cell. Hormones that bind to this type of receptor must be able to cross the cell membrane. Steroid hormones are derived from cholesterol and therefore can readily diffuse through the lipid bilayer of the cell membrane to reach the intracellular receptor (Figure 2). Thyroid hormones, which contain benzene rings studded with iodine, are also lipid-soluble and can enter the cell.

The location of steroid and thyroid hormone binding differs slightly: a steroid hormone may bind to its receptor within the cytosol or within the nucleus. In either case, this binding generates a hormone-receptor complex that moves toward the chromatin in the cell nucleus and binds to a particular segment of the cell’s DNA. In contrast, thyroid hormones bind to receptors already bound to DNA. For both steroid and thyroid hormones, binding of the hormone-receptor complex with DNA triggers transcription of a target gene to mRNA, which moves to the cytosol and directs protein synthesis by ribosomes.

[caption id="" align="aligncenter" width="480"]This illustration shows the steps involved with the binding of lipid-soluble hormones. Lipid-soluble hormones, such as steroid hormones, easily diffuse through the cell membrane. The hormone binds to its receptor in the cytosol, forming a receptor-hormone complex. The receptor-hormone complex then enters the nucleus and binds to the target gene on the cell’s DNA. Transcription of the gene creates a messenger RNA that is translated into the desired protein within the cytoplasm. It is these proteins that alter the cell’s activity. Figure 2. Binding of Lipid-Soluble Hormones. A steroid hormone directly initiates the production of proteins within a target cell. Steroid hormones easily diffuse through the cell membrane. The hormone binds to its receptor in the cytosol, forming a receptor–hormone complex. The receptor–hormone complex then enters the nucleus and binds to the target gene on the DNA. Transcription of the gene creates a messenger RNA that is translated into the desired protein within the cytoplasm.[/caption]

Pathways Involving Cell Membrane Hormone Receptors

Hydrophilic, or water-soluble, hormones are unable to diffuse through the lipid bilayer of the cell membrane and must therefore pass on their message to a receptor located at the surface of the cell. Except for thyroid hormones, which are lipid-soluble, all amino acid–derived hormones bind to cell membrane receptors that are located, at least in part, on the extracellular surface of the cell membrane. Therefore, they do not directly affect the transcription of target genes, but instead initiate a signaling cascade that is carried out by a molecule called a second messenger. In this case, the hormone is called a first messenger.

The second messenger used by most hormones is cyclic adenosine monophosphate (cAMP). In the cAMP second messenger system, a water-soluble hormone binds to its receptor in the cell membrane (Step 1 in Figure 3). This receptor is associated with an intracellular component called a G protein, and binding of the hormone activates the G-protein component (Step 2). The activated G protein in turn activates an enzyme called adenylyl cyclase, also known as adenylate cyclase (Step 3), which converts adenosine triphosphate (ATP) to cAMP (Step 4). As the second messenger, cAMP activates a type of enzyme called a protein kinase that is present in the cytosol (Step 5). Activated protein kinases initiate a phosphorylation cascade, in which multiple protein kinases phosphorylate (add a phosphate group to) numerous and various cellular proteins, including other enzymes (Step 6).
[caption id="" align="aligncenter" width="480"]This illustration shows the binding of water-soluble hormones. Water-soluble hormones cannot diffuse through the cell membrane. These hormones must bind to a receptor on the outer surface of the cell membrane. The receptor then activates a G protein in the cytoplasm, which travels to and activates adenylyl cyclase. Adenylyl cyclase catalyzes the conversion of ATP to CAMP, the secondary messenger in this pathway. CAMP, in turn, activates protein kinases, which phosphorylate proteins in the cytoplasm. This phosphorylation, shown as a P being added to a polypeptide chain, activates the proteins, allowing them to alter cell activity. Figure 3. Binding of Water-Soluble Hormones. Water-soluble hormones cannot diffuse through the cell membrane. These hormones must bind to a surface cell-membrane receptor. The receptor then initiates a cell-signaling pathway within the cell involving G proteins, adenylyl cyclase, the secondary messenger cyclic AMP (cAMP), and protein kinases. In the final step, these protein kinases phosphorylate proteins in the cytoplasm. This activates proteins in the cell that carry out the changes specified by the hormone.[/caption]
The phosphorylation of cellular proteins can trigger a wide variety of effects, from nutrient metabolism to the synthesis of different hormones and other products. The effects vary according to the type of target cell, the G proteins and kinases involved, and the phosphorylation of proteins. Examples of hormones that use cAMP as a second messenger include calcitonin, which is important for bone construction and regulating blood calcium levels; glucagon, which plays a role in blood glucose levels; and thyroid-stimulating hormone, which causes the release of T3 and T4 from the thyroid gland. Overall, the phosphorylation cascade significantly increases the efficiency, speed, and specificity of the hormonal response, as thousands of signaling events can be initiated simultaneously in response to a very low concentration of hormone in the bloodstream. However, the duration of the hormone signal is short, as cAMP is quickly deactivated by the enzyme phosphodiesterase (PDE), which is located in the cytosol. The action of PDE helps to ensure that a target cell’s response ceases quickly unless new hormones arrive at the cell membrane.

Importantly, there are also G proteins that decrease the levels of cAMP in the cell in response to hormone binding. For example, when growth hormone–inhibiting hormone (GHIH), also known as somatostatin, binds to its receptors in the pituitary gland, the level of cAMP decreases, thereby inhibiting the secretion of human growth hormone.

Not all water-soluble hormones initiate the cAMP second messenger system. One common alternative system uses calcium ions as a second messenger. In this system, G proteins activate the enzyme phospholipase C (PLC), which functions similarly to adenylyl cyclase. Once activated, PLC cleaves a membrane-bound phospholipid into two molecules: diacylglycerol (DAG) and inositol triphosphate (IP3). Like cAMP, DAG activates protein kinases that initiate a phosphorylation cascade. At the same time, IP3 causes calcium ions to be released from storage sites within the cytosol, such as from within the smooth endoplasmic reticulum. The calcium ions then act as second messengers in two ways: they can influence enzymatic and other cellular activities directly, or they can bind to calcium-binding proteins, the most common of which is calmodulin. Upon binding calcium, calmodulin is able to modulate protein kinase within the cell. Examples of hormones that use calcium ions as a second messenger system include angiotensin II, which helps regulate blood pressure through vasoconstriction, and growth hormone–releasing hormone (GHRH), which causes the pituitary gland to release growth hormones.

Factors Affecting Target Cell Response

You will recall that target cells must have receptors specific to a given hormone if that hormone is to trigger a response. But several other factors influence the target cell response. For example, the presence of a significant level of a hormone circulating in the bloodstream can cause its target cells to decrease their number of receptors for that hormone. This process is called downregulation, and it allows cells to become less reactive to the excessive hormone levels. When the level of a hormone is chronically reduced, target cells engage in upregulation to increase their number of receptors. This process allows cells to be more sensitive to the hormone that is present. Cells can also alter the sensitivity of the receptors themselves to various hormones. Two or more hormones can interact to affect the response of cells in a variety of ways. The three most common types of interaction are as follows:
  • The permissive effect, in which the presence of one hormone enables another hormone to act. For example, thyroid hormones have complex permissive relationships with certain reproductive hormones. A dietary deficiency of iodine, a component of thyroid hormones, can therefore affect reproductive system development and functioning.
  • The synergistic effect, in which two hormones with similar effects produce an amplified response. In some cases, two hormones are required for an adequate response. For example, two different reproductive hormones—FSH from the pituitary gland and estrogens from the ovaries—are required for the maturation of female ova (egg cells).
  • The antagonistic effect, in which two hormones have opposing effects. A familiar example is the effect of two pancreatic hormones, insulin and glucagon. Insulin increases the liver’s storage of glucose as glycogen, decreasing blood glucose, whereas glucagon stimulates the breakdown of glycogen stores, increasing blood glucose.

Regulation of Hormone Secretion

To prevent abnormal hormone levels and a potential disease state, hormone levels must be tightly controlled. The body maintains this control by balancing hormone production and degradation. Feedback loops govern the initiation and maintenance of most hormone secretion in response to various stimuli.

Role of Feedback Loops

The contribution of feedback loops to homeostasis will only be briefly reviewed here. Positive feedback loops are characterized by the release of additional hormone in response to an original hormone release. The release of oxytocin during childbirth is a positive feedback loop. The initial release of oxytocin begins to signal the uterine muscles to contract, which pushes the fetus toward the cervix, causing it to stretch. This, in turn, signals the pituitary gland to release more oxytocin, causing labor contractions to intensify. The release of oxytocin decreases after the birth of the child.

The more common method of hormone regulation is the negative feedback loop. Negative feedback is characterized by the inhibition of further secretion of a hormone in response to adequate levels of that hormone. This allows blood levels of the hormone to be regulated within a narrow range. An example of a negative feedback loop is the release of glucocorticoid hormones from the adrenal glands, as directed by the hypothalamus and pituitary gland. As glucocorticoid concentrations in the blood rise, the hypothalamus and pituitary gland reduce their signaling to the adrenal glands to prevent additional glucocorticoid secretion (Figure 4).

[caption id="" align="aligncenter" width="520"]This diagram shows a negative feedback loop using the example of glucocorticoid regulation in the blood. Step 1 in the cycle is when an imbalance occurs. The hypothalamus perceives low blood concentrations of glucocorticoids in the blood. This is illustrated by there being only 5 glucocorticoids floating in a cross section of an artery. Step 2 in the cycle is hormone release, where the hypothalamus releases corticotropin-releasing hormone (CRH). Step 3 is labeled correction. Here, the CRH release starts a hormone cascade that triggers the adrenal gland to release glucocorticoid into the blood. This allows the blood concentration of glucocorticoid to increase, as illustrated by 8 glucocorticoid molecules now being present in the cross section of the artery. Step 4 is labeled negative feedback. Here, the hypothalamus perceives normal concentrations of glucocorticoids in the blood and stops releasing CRH. This brings blood glucocorticoid levels back to homeostasis. Figure 4. Negative Feedback Loop. The release of adrenal glucocorticoids is stimulated by the release of hormones from the hypothalamus and pituitary gland. This signaling is inhibited when glucocorticoid levels become elevated by causing negative signals to the pituitary gland and hypothalamus.[/caption]

Role of Endocrine Gland Stimuli

Reflexes triggered by both chemical and neural stimuli control endocrine activity. These reflexes may be simple, involving only one hormone response, or they may be more complex and involve many hormones, as is the case with the hypothalamic control of various anterior pituitary–controlled hormones.

Humoral stimuli are changes in blood levels of non-hormone chemicals, such as nutrients or ions, which cause the release or inhibition of a hormone to, in turn, maintain homeostasis. For example, osmoreceptors in the hypothalamus detect changes in blood osmolarity (the concentration of solutes in the blood plasma). If blood osmolarity is too high, meaning that the blood is not dilute enough, osmoreceptors signal the hypothalamus to release ADH. The hormone causes the kidneys to reabsorb more water and reduce the volume of urine produced. This reabsorption causes a reduction of the osmolarity of the blood, diluting the blood to the appropriate level. The regulation of blood glucose is another example. High blood glucose levels cause the release of insulin from the pancreas, which increases glucose uptake by cells and liver storage of glucose as glycogen.

An endocrine gland may also secrete a hormone in response to the presence of another hormone produced by a different endocrine gland. Such hormonal stimuli often involve the hypothalamus, which produces releasing and inhibiting hormones that control the secretion of a variety of pituitary hormones.

In addition to these chemical signals, hormones can also be released in response to neural stimuli. A common example of neural stimuli is the activation of the fight-or-flight response by the sympathetic nervous system. When an individual perceives danger, sympathetic neurons signal the adrenal glands to secrete norepinephrine and epinephrine. The two hormones dilate blood vessels, increase the heart and respiratory rate, and suppress the digestive and immune systems. These responses boost the body’s transport of oxygen to the brain and muscles, thereby improving the body’s ability to fight or flee.

]]> 2593 2585 2 0 17.3 The Pituitary Gland and Hypothalamus https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2599 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2599

Learning Objectives

By the end of this section, you will be able to:
  • Explain the interrelationships of the anatomy and functions of the hypothalamus and the posterior and anterior lobes of the pituitary gland
  • Identify the two hormones released from the posterior pituitary, their target cells, and their principal actions
  • Identify the six hormones produced by the anterior lobe of the pituitary gland, their target cells, their principal actions, and their regulation by the hypothalamus

The hypothalamus–pituitary complex can be thought of as the “command center” of the endocrine system. This complex secretes several hormones that directly produce responses in target tissues, as well as hormones that regulate the synthesis and secretion of hormones of other glands. In addition, the hypothalamus–pituitary complex coordinates the messages of the endocrine and nervous systems. In many cases, a stimulus received by the nervous system must pass through the hypothalamus–pituitary complex to be translated into hormones that can initiate a response.

The hypothalamus is a structure of the diencephalon of the brain located anterior and inferior to the thalamus (Figure 1). It has both neural and endocrine functions, producing and secreting many hormones. In addition, the hypothalamus is anatomically and functionally related to the pituitary gland (or hypophysis), a bean-sized organ suspended from it by a stem called the infundibulum (or pituitary stalk). The pituitary gland is cradled within the sellaturcica of the sphenoid bone of the skull. It consists of two lobes that arise from distinct parts of embryonic tissue: the posterior pituitary (neurohypophysis) is neural tissue, whereas the anterior pituitary (also known as the adenohypophysis) is glandular tissue that develops from the primitive digestive tract. The hormones secreted by the posterior and anterior pituitary, and the intermediate zone between the lobes are summarized in Table 3.

[caption id="" align="aligncenter" width="520"]This illustration shows the hypothalamus-pituitary complex, which is located at the base of the brain and shown here from a lateral view. The hypothalamus lies inferior and anterior to the thalamus, which is sits atop the brainstem. The hypothalamus connects to the pituitary gland by the stalk-like infundibulum. The pituitary gland looks like a sac containing two balls hanging from the infundibulum. The “balls” are the anterior and posterior lobes of the pituitary. Each lobe secretes different hormones in response to signals from the hypothalamus. Figure 1. Hypothalamus–Pituitary Complex. The hypothalamus region lies inferior and anterior to the thalamus. It connects to the pituitary gland by the stalk-like infundibulum. The pituitary gland consists of an anterior and posterior lobe, with each lobe secreting different hormones in response to signals from the hypothalamus.[/caption]
Pituitary Hormones (Table 3)
Pituitary lobe Associated hormones Chemical class Effect
Anterior Growth hormone (GH) Protein Promotes growth of body tissues
Anterior Prolactin (PRL) Peptide Promotes milk production from mammary glands
Anterior Thyroid-stimulating hormone (TSH) Glycoprotein Stimulates thyroid hormone release from thyroid
Anterior Adrenocorticotropic hormone (ACTH) Peptide Stimulates hormone release by adrenal cortex
Anterior Follicle-stimulating hormone (FSH) Glycoprotein Stimulates gamete production in gonads
Anterior Luteinizing hormone (LH) Glycoprotein Stimulates androgen production by gonads
Posterior Antidiuretic hormone (ADH) Peptide Stimulates water reabsorption by kidneys
Posterior Oxytocin Peptide Stimulates uterine contractions during childbirth
Intermediate zone Melanocyte-stimulating hormone Peptide Stimulates melanin formation in melanocytes

Posterior Pituitary

The posterior pituitary is actually an extension of the neurons of the paraventricular and supraoptic nuclei of the hypothalamus. The cell bodies of these regions rest in the hypothalamus, but their axons descend as the hypothalamic–hypophyseal tract within the infundibulum, and end in axon terminals that comprise the posterior pituitary (Figure 2).

[caption id="" align="aligncenter" width="520"]This illustration zooms in on the hypothalamus and the attached pituitary gland. The posterior pituitary is highlighted. Two nuclei in the hypothalamus contain neurosecretory cells that release different hormones. The neurosecretory cells of the paraventricular nucleus release oxytocin (OT) while the neurosecretory cells of the supraoptic nucleus release anti-diuretic hormone (ADH). The neurosecretory cells stretch down the infundibulum into the posterior pituitary. The tube-like extensions of the neurosecretory cells within the infundibulum are labeled the hypothalamophypophyseal tracts. These tracts connect with a web-like network of blood vessels in the posterior pituitary called the capillary plexus. From the capillary plexus, the posterior pituitary secretes the OT or ADH into a single vein that exits the pituitary. Figure 2. Posterior Pituitary. Neurosecretory cells in the hypothalamus release oxytocin (OT) or ADH into the posterior lobe of the pituitary gland. These hormones are stored or released into the blood via the capillary plexus.[/caption]

The posterior pituitary gland does not produce hormones, but rather stores and secretes hormones produced by the hypothalamus. The paraventricular nuclei produce the hormone oxytocin, whereas the supraoptic nuclei produce ADH. These hormones travel along the axons into storage sites in the axon terminals of the posterior pituitary. In response to signals from the same hypothalamic neurons, the hormones are released from the axon terminals into the bloodstream.

Oxytocin

When fetal development is complete, the peptide-derived hormone oxytocin (tocia- = “childbirth”) stimulates uterine contractions and dilation of the cervix. Throughout most of pregnancy, oxytocin hormone receptors are not expressed at high levels in the uterus. Toward the end of pregnancy, the synthesis of oxytocin receptors in the uterus increases, and the smooth muscle cells of the uterus become more sensitive to its effects. Oxytocin is continually released throughout childbirth through a positive feedback mechanism. As noted earlier, oxytocin prompts uterine contractions that push the fetal head toward the cervix. In response, cervical stretching stimulates additional oxytocin to be synthesized by the hypothalamus and released from the pituitary. This increases the intensity and effectiveness of uterine contractions and prompts additional dilation of the cervix. The feedback loop continues until birth.

Although the mother’s high blood levels of oxytocin begin to decrease immediately following birth, oxytocin continues to play a role in maternal and newborn health. First, oxytocin is necessary for the milk ejection reflex (commonly referred to as “let-down”) in breastfeeding women. As the newborn begins suckling, sensory receptors in the nipples transmit signals to the hypothalamus. In response, oxytocin is secreted and released into the bloodstream. Within seconds, cells in the mother’s milk ducts contract, ejecting milk into the infant’s mouth. Secondly, in both males and females, oxytocin is thought to contribute to parent–newborn bonding, known as attachment. Oxytocin is also thought to be involved in feelings of love and closeness, as well as in the sexual response.

Antidiuretic Hormone (ADH)

The solute concentration of the blood, or blood osmolarity, may change in response to the consumption of certain foods and fluids, as well as in response to disease, injury, medications, or other factors. Blood osmolarity is constantly monitored by osmoreceptors—specialized cells within the hypothalamus that are particularly sensitive to the concentration of sodium ions and other solutes.

In response to high blood osmolarity, which can occur during dehydration or following a very salty meal, the osmoreceptors signal the posterior pituitary to release antidiuretic hormone (ADH). The target cells of ADH are located in the tubular cells of the kidneys. Its effect is to increase epithelial permeability to water, allowing increased water reabsorption. The more water reabsorbed from the filtrate, the greater the amount of water that is returned to the blood and the less that is excreted in the urine. A greater concentration of water results in a reduced concentration of solutes. ADH is also known as vasopressin because, in very high concentrations, it causes constriction of blood vessels, which increases blood pressure by increasing peripheral resistance. The release of ADH is controlled by a negative feedback loop. As blood osmolarity decreases, the hypothalamic osmoreceptors sense the change and prompt a corresponding decrease in the secretion of ADH. As a result, less water is reabsorbed from the urine filtrate.

Interestingly, drugs can affect the secretion of ADH. For example, alcohol consumption inhibits the release of ADH, resulting in increased urine production that can eventually lead to dehydration and a hangover. A disease called diabetes insipidus is characterized by chronic underproduction of ADH that causes chronic dehydration. Because little ADH is produced and secreted, not enough water is reabsorbed by the kidneys. Although patients feel thirsty, and increase their fluid consumption, this doesn’t effectively decrease the solute concentration in their blood because ADH levels are not high enough to trigger water reabsorption in the kidneys. Electrolyte imbalances can occur in severe cases of diabetes insipidus.

Anterior Pituitary

The anterior pituitary originates from the digestive tract in the embryo and migrates toward the brain during fetal development. There are three regions: the pars distalis is the most anterior, the pars intermedia is adjacent to the posterior pituitary, and the pars tuberalis is a slender “tube” that wraps the infundibulum. Recall that the posterior pituitary does not synthesize hormones, but merely stores them. In contrast, the anterior pituitary does manufacture hormones. However, the secretion of hormones from the anterior pituitary is regulated by two classes of hormones. These hormones—secreted by the hypothalamus—are the releasing hormones that stimulate the secretion of hormones from the anterior pituitary and the inhibiting hormones that inhibit secretion. Hypothalamic hormones are secreted by neurons, but enter the anterior pituitary through blood vessels (Figure 3). Within the infundibulum is a bridge of capillaries that connects the hypothalamus to the anterior pituitary. This network, called the hypophyseal portal system, allows hypothalamic hormones to be transported to the anterior pituitary without first entering the systemic circulation. The system originates from the superior hypophyseal artery, which branches off the carotid arteries and transports blood to the hypothalamus. The branches of the superior hypophyseal artery form the hypophyseal portal system (see Figure 3). Hypothalamic releasing and inhibiting hormones travel through a primary capillary plexus to the portal veins, which carry them into the anterior pituitary. Hormones produced by the anterior pituitary (in response to releasing hormones) enter a secondary capillary plexus, and from there drain into the circulation.
[caption id="" align="aligncenter" width="520"]This illustration zooms in on the hypothalamus and the attached pituitary gland. The anterior pituitary is highlighted. Three neurosecretory cells are secreting hormones into a web-like network of arteries within the infundibulum. The artery net is labeled the primary capillary plexus of the hypophyseal portal system. The superior hypophysel artery enters the primary capillary plexus from outside of the infundibulum. The hypophyseal portal vein runs down from the primary capillary plexus, through the infundibulum, and connects to the secondary capillary plexus of the hypophyseal portal system. The secondary capillary plexus is located within the anterior pituitary. The hormones released from the neurosecretory cells of the hypothalamus travel through the primary capillary plexus, down the hypophyseal portal vein, and into the secondary capillary plexus. There, the hypothalamus hormones stimulate the anterior pituitary to release its hormones. The anterior pituitary hormones leave the primary capillary plexus from a single vein at the bottom of the anterior lobe. Figure 3. Anterior Pituitary. The anterior pituitary manufactures seven hormones. The hypothalamus produces separate hormones that stimulate or inhibit hormone production in the anterior pituitary. Hormones from the hypothalamus reach the anterior pituitary via the hypophyseal portal system.[/caption]

The anterior pituitary produces seven hormones. These are the growth hormone (GH), thyroid-stimulating hormone (TSH), adrenocorticotropic hormone (ACTH), follicle-stimulating hormone (FSH), luteinizing hormone (LH), beta endorphin, and prolactin. Of the hormones of the anterior pituitary, TSH, ACTH, FSH, and LH are collectively referred to as tropic hormones (trope- = “turning”) because they turn on or off the function of other endocrine glands.

Growth Hormone

The endocrine system regulates the growth of the human body, protein synthesis, and cellular replication. A major hormone involved in this process is growth hormone (GH), also called somatotropin—a protein hormone produced and secreted by the anterior pituitary gland. Its primary function is anabolic; it promotes protein synthesis and tissue building through direct and indirect mechanisms (Figure 4). GH levels are controlled by the release of GHRH and GHIH (also known as somatostatin) from the hypothalamus.

[caption id="" align="aligncenter" width="550"]This flow chart illustrates the hormone cascade that stimulates human growth. In step 1, the hypothalamus releases growth hormone-releasing hormone (GHRH). GHRH travels into the primary capillary plexus of the anterior pituitary, where it stimulates the anterior pituitary to release growth hormone (GH). The release of growth hormone causes three types of effects. In the glucose-sparing effect, GH stimulates adipose cells to break down stored fat, fueling the growth effects (discussed next). The target cells for the glucose-sparing effects are adipose cells. In the growth effects, GH increases the uptake of amino acids from the blood and enhances cellular proliferation while also reducing apoptosis. The target cells for the growth effects are bone cells, muscle cells, nervous system cells, and immune system cells. In the diabetogenic effect, GH stimulates the liver to break down glycogen into glucose, fueling the growth effects. The liver also releases IGF in response to GH. The IGF further stimulates the growth effects but also negatively feeds back to the hypothalamus. When high IGF one levels are perceived by the hypothalamus, it releases growth hormone inhibiting hormone (GHIH). GHIH inhibits GH release by the anterior pituitary. Figure 4. Hormonal Regulation of Growth. Growth hormone (GH) directly accelerates the rate of protein synthesis in skeletal muscle and bones. Insulin-like growth factor 1 (IGF-1) is activated by growth hormone and indirectly supports the formation of new proteins in muscle cells and bone.[/caption]
A glucose-sparing effect occurs when GH stimulates lipolysis, or the breakdown of adipose tissue, releasing fatty acids into the blood. As a result, many tissues switch from glucose to fatty acids as their main energy source, which means that less glucose is taken up from the bloodstream.

GH also initiates the diabetogenic effect in which GH stimulates the liver to break down glycogen to glucose, which is then deposited into the blood. The name “diabetogenic” is derived from the similarity in elevated blood glucose levels observed between individuals with untreated diabetes mellitus and individuals experiencing GH excess. Blood glucose levels rise as the result of a combination of glucose-sparing and diabetogenic effects.

GH indirectly mediates growth and protein synthesis by triggering the liver and other tissues to produce a group of proteins called insulin-like growth factors (IGFs). These proteins enhance cellular proliferation and inhibit apoptosis, or programmed cell death. IGFs stimulate cells to increase their uptake of amino acids from the blood for protein synthesis. Skeletal muscle and cartilage cells are particularly sensitive to stimulation from IGFs.

Dysfunction of the endocrine system’s control of growth can result in several disorders. For example, gigantism is a disorder in children that is caused by the secretion of abnormally large amounts of GH, resulting in excessive growth. A similar condition in adults is acromegaly, a disorder that results in the growth of bones in the face, hands, and feet in response to excessive levels of GH in individuals who have stopped growing. Abnormally low levels of GH in children can cause growth impairment—a disorder called pituitary dwarfism (also known as growth hormone deficiency).

Thyroid-Stimulating Hormone

The activity of the thyroid gland is regulated by thyroid-stimulating hormone (TSH), also called thyrotropin. TSH is released from the anterior pituitary in response to thyrotropin-releasing hormone (TRH) from the hypothalamus. As discussed shortly, it triggers the secretion of thyroid hormones by the thyroid gland. In a classic negative feedback loop, elevated levels of thyroid hormones in the bloodstream then trigger a drop in production of TRH and subsequently TSH.

Adrenocorticotropic Hormone

The adrenocorticotropic hormone (ACTH), also called corticotropin, stimulates the adrenal cortex (the more superficial “bark” of the adrenal glands) to secrete corticosteroid hormones such as cortisol. ACTH come from a precursor molecule known as pro-opiomelanotropin (POMC) which produces several biologically active molecules when cleaved, including ACTH, melanocyte-stimulating hormone, and the brain opioid peptides known as endorphins. The release of ACTH is regulated by the corticotropin-releasing hormone (CRH) from the hypothalamus in response to normal physiologic rhythms. A variety of stressors can also influence its release, and the role of ACTH in the stress response is discussed later in this chapter.

Follicle-Stimulating Hormone and Luteinizing Hormone

The endocrine glands secrete a variety of hormones that control the development and regulation of the reproductive system (these glands include the anterior pituitary, the adrenal cortex, and the gonads—the testes in males and the ovaries in females). Much of the development of the reproductive system occurs during puberty and is marked by the development of sex-specific characteristics in both male and female adolescents. Puberty is initiated by gonadotropin-releasing hormone (GnRH), a hormone produced and secreted by the hypothalamus. GnRH stimulates the anterior pituitary to secrete gonadotropins—hormones that regulate the function of the gonads. The levels of GnRH are regulated through a negative feedback loop; high levels of reproductive hormones inhibit the release of GnRH. Throughout life, gonadotropins regulate reproductive function and, in the case of women, the onset and cessation of reproductive capacity.

The gonadotropins include two glycoprotein hormones: follicle-stimulating hormone (FSH) stimulates the production and maturation of sex cells, or gametes, including ova in women and sperm in men. FSH also promotes follicular growth; these follicles then release estrogens in the female ovaries. Luteinizing hormone (LH) triggers ovulation in women, as well as the production of estrogens and progesterone by the ovaries. LH stimulates production of testosterone by the male testes.

Prolactin

As its name implies, prolactin (PRL) promotes lactation (milk production) in women. During pregnancy, it contributes to development of the mammary glands, and after birth, it stimulates the mammary glands to produce breast milk. However, the effects of prolactin depend heavily upon the permissive effects of estrogens, progesterone, and other hormones. And as noted earlier, the let-down of milk occurs in response to stimulation from oxytocin. In a non-pregnant woman, prolactin secretion is inhibited by prolactin-inhibiting hormone (PIH), which is actually the neurotransmitter dopamine, and is released from neurons in the hypothalamus. Only during pregnancy do prolactin levels rise in response to prolactin-releasing hormone (PRH) from the hypothalamus.

Intermediate Pituitary: Melanocyte-Stimulating Hormone

The cells in the zone between the pituitary lobes secrete a hormone known as melanocyte-stimulating hormone (MSH) that is formed by cleavage of the pro-opiomelanocortin (POMC) precursor protein. Local production of MSH in the skin is responsible for melanin production in response to UV light exposure. The role of MSH made by the pituitary is more complicated. For instance, people with lighter skin generally have the same amount of MSH as people with darker skin. Nevertheless, this hormone is capable of darkening of the skin by inducing melanin production in the skin’s melanocytes. Women also show increased MSH production during pregnancy; in combination with estrogens, it can lead to darker skin pigmentation, especially the skin of the areolas and labia minora. Figure 5 is a summary of the pituitary hormones and their principal effects.

[caption id="" align="aligncenter" width="550"]These two diagrammatic tables show the major pituitary hormones, their releasing hormone from the hypothalamus, their target organs, and their effects. The top part of the diagram shows the posterior pituitary hormones. ADH is produced by the hypothalamus and stored in the posterior pituitary. The targets of ADH are the kidneys, sweat glands and circulatory system, as this hormone affects water balance. OT is produced by the posterior pituitary and has no releasing hormone. Its target is the female reproductive system, as this hormone triggers uterine contractions during childbirth. The anterior pituitary hormones are listed in the lower diagram. The release of LH by the anterior pituitary is triggered by the release of GNRH from the hypothalamus. The target of LH is the reproductive system, as this hormone stimulates the production of sex hormones by the gonads. The release of FSH by the anterior pituitary is triggered by the release of GNRH from the hypothalamus. The target of FSH is the reproductive system, as this hormone stimulates the production of sperm and eggs. The release of TSH by the anterior pituitary is triggered by the release of TRH from the hypothalamus. The target of TSH is the thyroid gland, as this hormone stimulates the release of thyroid hormone (TH). TH regulates metabolism. The release of PRL by the anterior pituitary is triggered by the release of PRH and inhibited by the release of PIH from the hypothalamus. The target of PRL is the mammary glands, as this hormone promotes milk production. The release of GH by the anterior pituitary is triggered by the release of GHRH and inhibited by the release of GHIH from the hypothalamus. The targets of GH are the liver, bones and muscles, as it induces its targets to produce insulin-like growth factors (IGH), as this hormone stimulates body growth and a higher metabolic rate. The release of ACTH by the anterior pituitary is triggered by the release of CRH from the hypothalamus. The targets of ACTH are the adrenal glands, as this hormone induces its targets to produce glucocorticoids, which regulate metabolism and the stress response. Figure 5. Major Pituitary Hormones. Major pituitary hormones and their target organs.[/caption]
 

]]> 2599 2585 3 0 17.4 The Thyroid Gland https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2603 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2603

Learning Objectives

By the end of this section, you will be able to:
  • Describe the location and anatomy of the thyroid gland
  • Discuss the synthesis of triiodothyronine and thyroxine
  • Explain the role of thyroid hormones in the regulation of basal metabolism
  • Identify the hormone produced by the parafollicular cells of the thyroid

A butterfly-shaped organ, the thyroid gland is located anterior to the trachea, just inferior to the larynx (Figure 1). The medial region, called the isthmus, is flanked by wing-shaped left and right lobes. Each of the thyroid lobes are embedded with parathyroid glands, primarily on their posterior surfaces. The tissue of the thyroid gland is composed mostly of thyroid follicles. The follicles are made up of a central cavity filled with a sticky fluid called colloid. Surrounded by a wall of epithelial follicle cells, the colloid is the center of thyroid hormone production, and that production is dependent on the hormones’ essential and unique component: iodine.

[caption id="" align="aligncenter" width="380"]Part A of this figure is a diagram of the anterior view of the thyroid gland. The thyroid gland is a butterfly-shaped gland wrapping around the trachea. It narrows at its center, just under the thyroid cartilage of the larynx. This narrow area is called the isthmus of the thyroid. Two large arteries, the common carotid arteries, run parallel to the trachea on the outer border of the thyroid. A small artery enters the superior edge of the thyroid, near the isthmus, and branches throughout the two “wings” of the thyroid. Part B of this figure is a posterior view of the thyroid. The posterior view shows that the thyroid does not completely wrap around the posterior of the trachea. The posterior sides of the thyroid wings can be seen protruding from under the cricoid cartilage of the larynx. The posterior sides of the thyroid “wings” each contain two small, disc-shaped parathyroid glands embedded in the thyroid tissue. Within each wing, one disc is located superior to the other. These are labeled the left and right parathyroid glands. Just under the inferior parathyroid glands are two arteries that bring blood to the thyroid from the left and right subclavian arteries. Part C of this figure is a micrograph of thyroid tissue. The thyroid follicle cells are cuboidal epithelial cells. These cells form a ring around irregular-shaped cavities called follicles. The follicles contain light colored colloid. A larger parafollicular cell is embedded between two of the follicular cells near the edge of a follicle. Figure 1. Thyroid Gland. The thyroid gland is located in the neck where it wraps around the trachea. (a) Anterior view of the thyroid gland. (b) Posterior view of the thyroid gland. (c) The glandular tissue is composed primarily of thyroid follicles. The larger parafollicular cells often appear within the matrix of follicle cells. LM × 1332. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Synthesis and Release of Thyroid Hormones

Hormones are produced in the colloid when atoms of the mineral iodine attach to a glycoprotein, called thyroglobulin, that is secreted into the colloid by the follicle cells. The following steps outline the hormones’ assembly:

  1. Binding of TSH to its receptors in the follicle cells of the thyroid gland causes the cells to actively transport iodide ions (I–) across their cell membrane, from the bloodstream into the cytosol. As a result, the concentration of iodide ions “trapped” in the follicular cells is many times higher than the concentration in the bloodstream.
  2. Iodide ions then move to the lumen of the follicle cells that border the colloid. There, the ions undergo oxidation (their negatively charged electrons are removed). The oxidation of two iodide ions (2 I–) results in iodine (I2), which passes through the follicle cell membrane into the colloid.
  3. In the colloid, peroxidase enzymes link the iodine to the tyrosine amino acids in thyroglobulin to produce two intermediaries: a tyrosine attached to one iodine and a tyrosine attached to two iodines. When one of each of these intermediaries is linked by covalent bonds, the resulting compound is triiodothyronine (T3), a thyroid hormone with three iodines. Much more commonly, two copies of the second intermediary bond, forming tetraiodothyronine, also known as thyroxine (T4), a thyroid hormone with four iodines.

These hormones remain in the colloid center of the thyroid follicles until TSH stimulates endocytosis of colloid back into the follicle cells. There, lysosomal enzymes break apart the thyroglobulin colloid, releasing free T3 and T4, which diffuse across the follicle cell membrane and enter the bloodstream.

In the bloodstream, less than one percent of the circulating T3 and T4 remains unbound. This free T3 and T4 can cross the lipid bilayer of cell membranes and be taken up by cells. The remaining 99 percent of circulating T3 and T4 is bound to specialized transport proteins called thyroxine-binding globulins (TBGs), to albumin, or to other plasma proteins. This “packaging” prevents their free diffusion into body cells. When blood levels of T3 and T4 begin to decline, bound T3 and T4 are released from these plasma proteins and readily cross the membrane of target cells. T3 is more potent than T4, and many cells convert T4 to T3 through the removal of an iodine atom.

Regulation of TH Synthesis

The release of T3 and T4 from the thyroid gland is regulated by thyroid-stimulating hormone (TSH). As shown in Figure 2, low blood levels of T3 and T4 stimulate the release of thyrotropin-releasing hormone (TRH) from the hypothalamus, which triggers secretion of TSH from the anterior pituitary. In turn, TSH stimulates the thyroid gland to secrete T3 and T4. The levels of TRH, TSH, T3, and T4 are regulated by a negative feedback system in which increasing levels of T3 and T4 decrease the production and secretion of TSH.

[caption id="" align="aligncenter" width="520"]This diagram illustrates a negative feedback loop. It shows the general steps of a negative feedback loop at the center (imbalance, hormone release, correction, and negative feedback) using the example of the hormone cascade that regulates metabolic rate. The hypothalamus releases TRH in response to low metabolic rate and or low T three and T four concentrations in the blood (imbalance). This triggers TSH release by the pituitary (hormone release). The TSH travels to the thyroid where it triggers T three and T four release by the thyroid cells. T three and T four increase basal metabolic rate of the body cells and cause a rise in body temperature (the calorigenic effect). T three and T four then feed back to the hypothalamus and inhibits TRH and TSH release. If metabolic rate is high and or T three and T four concentrations are low, then the hypothalamus stops releasing TRH (negative feedback). As a result, the anterior pituitary will not release TSH and no T three or T four will be produced by the thyroid. Figure 2. Classic Negative Feedback Loop. A classic negative feedback loop controls the regulation of thyroid hormone levels.[/caption]

Functions of Thyroid Hormones

The thyroid hormones, T3 and T4, are often referred to as metabolic hormones because their levels influence the body’s basal metabolic rate, the amount of energy used by the body at rest. When T3 and T4 bind to intracellular receptors located on the mitochondria, they cause an increase in nutrient breakdown and the use of oxygen to produce ATP. In addition, T3 and T4 initiate the transcription of genes involved in glucose oxidation. Although these mechanisms prompt cells to produce more ATP, the process is inefficient, and an abnormally increased level of heat is released as a byproduct of these reactions. This so-called calorigenic effect (calor- = “heat”) raises body temperature.

Adequate levels of thyroid hormones are also required for protein synthesis and for fetal and childhood tissue development and growth. They are especially critical for normal development of the nervous system both in utero and in early childhood, and they continue to support neurological function in adults. As noted earlier, these thyroid hormones have a complex interrelationship with reproductive hormones, and deficiencies can influence libido, fertility, and other aspects of reproductive function. Finally, thyroid hormones increase the body’s sensitivity to catecholamines (epinephrine and norepinephrine) from the adrenal medulla by upregulation of receptors in the blood vessels. When levels of T3 and T4 hormones are excessive, this effect accelerates the heart rate, strengthens the heartbeat, and increases blood pressure. Because thyroid hormones regulate metabolism, heat production, protein synthesis, and many other body functions, thyroid disorders can have severe and widespread consequences.

Disorders of the…

Endocrine System: Iodine Deficiency, Hypothyroidism, and Hyperthyroidism As discussed above, dietary iodine is required for the synthesis of T3 and T4. But for much of the world’s population, foods do not provide adequate levels of this mineral, because the amount varies according to the level in the soil in which the food was grown, as well as the irrigation and fertilizers used. Marine fish and shrimp tend to have high levels because they concentrate iodine from seawater, but many people in landlocked regions lack access to seafood. Thus, the primary source of dietary iodine in many countries is iodized salt. Fortification of salt with iodine began in the United States in 1924, and international efforts to iodize salt in the world’s poorest nations continue today.

Dietary iodine deficiency can result in the impaired ability to synthesize T3 and T4, leading to a variety of severe disorders. When T3 and T4 cannot be produced, TSH is secreted in increasing amounts. As a result of this hyperstimulation, thyroglobulin accumulates in the thyroid gland follicles, increasing their deposits of colloid. The accumulation of colloid increases the overall size of the thyroid gland, a condition called a goiter (Figure 3). A goiter is only a visible indication of the deficiency. Other iodine deficiency disorders include impaired growth and development, decreased fertility, and prenatal and infant death. Moreover, iodine deficiency is the primary cause of preventable mental retardation worldwide. Neonatal hypothyroidism (cretinism) is characterized by cognitive deficits, short stature, and sometimes deafness and muteness in children and adults born to mothers who were iodine-deficient during pregnancy.
[caption id="" align="aligncenter" width="380"]This photo shows a woman with a goiter, which is an extreme, irregular swelling on the anterior side of the neck. Figure 3. Goiter. (credit: “Almazi”/Wikimedia Commons)[/caption]

In areas of the world with access to iodized salt, dietary deficiency is rare. Instead, inflammation of the thyroid gland is the more common cause of low blood levels of thyroid hormones. Called hypothyroidism, the condition is characterized by a low metabolic rate, weight gain, cold extremities, constipation, reduced libido, menstrual irregularities, and reduced mental activity. In contrast, hyperthyroidism—an abnormally elevated blood level of thyroid hormones—is often caused by a pituitary or thyroid tumor. In Graves’ disease, the hyperthyroid state results from an autoimmune reaction in which antibodies overstimulate the follicle cells of the thyroid gland. Hyperthyroidism can lead to an increased metabolic rate, excessive body heat and sweating, diarrhea, weight loss, tremors, and increased heart rate. The person’s eyes may bulge (called exophthalmos) as antibodies produce inflammation in the soft tissues of the orbits. The person may also develop a goiter.

Calcitonin

The thyroid gland also secretes a hormone called calcitonin that is produced by the parafollicular cells (also called C cells) that stud the tissue between distinct follicles. Calcitonin is released in response to a rise in blood calcium levels. It appears to have a function in decreasing blood calcium concentrations by:

  • Inhibiting the activity of osteoclasts, bone cells that release calcium into the circulation by degrading bone matrix
  • Increasing osteoblastic activity
  • Decreasing calcium absorption in the intestines
  • Increasing calcium loss in the urine

However, these functions are usually not significant in maintaining calcium homeostasis, so the importance of calcitonin is not entirely understood. Pharmaceutical preparations of calcitonin are sometimes prescribed to reduce osteoclast activity in people with osteoporosis and to reduce the degradation of cartilage in people with osteoarthritis. The hormones secreted by thyroid are summarized in Table 4.

Thyroid Hormones (Table 4)
Associated hormones Chemical class Effect
Thyroxine (T4), triiodothyronine (T3) Amine Stimulate basal metabolic rate
Calcitonin Peptide Reduces blood Ca2+ levels

Of course, calcium is critical for many other biological processes. It is a second messenger in many signaling pathways, and is essential for muscle contraction, nerve impulse transmission, and blood clotting. Given these roles, it is not surprising that blood calcium levels are tightly regulated by the endocrine system. The organs involved in the regulation are the parathyroid glands.

]]> 2603 2585 4 0 17.5 The Parathyroid Glands https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2606 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2606

Learning Objectives

By the end of this section, you will be able to:
  • Describe the location and structure of the parathyroid glands
  • Describe the hormonal control of blood calcium levels
  • Discuss the physiological response of parathyroid dysfunction

The parathyroid glands are tiny, round structures usually found embedded in the posterior surface of the thyroid gland (Figure 1). A thick connective tissue capsule separates the glands from the thyroid tissue. Most people have four parathyroid glands, but occasionally there are more in tissues of the neck or chest. The function of one type of parathyroid cells, the oxyphil cells, is not clear. The primary functional cells of the parathyroid glands are the chief cells. These epithelial cells produce and secrete the parathyroid hormone (PTH), the major hormone involved in the regulation of blood calcium levels.

[caption id="" align="aligncenter" width="550"]Part A of this diagram shows the four, small, disc-shaped parathyroid glands embedded in the posterior surface of the thyroid gland. Part B shows a micrograph of parathyroid tissue. The tissue is largely composed of cube-shaped chief cells encircling a central blood vessel. A few larger and darker-staining oxyphil cells are embedded within the many chief cells. Figure 1. Parathyroid Glands. The small parathyroid glands are embedded in the posterior surface of the thyroid gland. LM × 760. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

The parathyroid glands produce and secrete PTH, a peptide hormone, in response to low blood calcium levels (Figure 2). PTH secretion causes the release of calcium from the bones by stimulating osteoclasts, which secrete enzymes that degrade bone and release calcium into the interstitial fluid. PTH also inhibits osteoblasts, the cells involved in bone deposition, thereby sparing blood calcium. PTH causes increased reabsorption of calcium (and magnesium) in the kidney tubules from the urine filtrate. In addition, PTH initiates the production of the steroid hormone calcitriol (also known as 1,25-dihydroxyvitamin D), which is the active form of vitamin D3, in the kidneys. Calcitriol then stimulates increased absorption of dietary calcium by the intestines. A negative feedback loop regulates the levels of PTH, with rising blood calcium levels inhibiting further release of PTH.

[caption id="" align="aligncenter" width="550"]This diagram shows the role of parathyroid hormone in maintaining blood calcium homeostasis. When blood calcium concentration drops, chief cells of the parathyroid gland release parathyroid hormone (PTH). PTH affects bone, the kidneys and the intestines. In regards to bone, PTH inhibits osteoblasts and stimulates osteoclasts. This results in compact bone being broken down, as illustrated by an osteoclast burrowing into the surface of a bone. The break down releases calcium ions into a nearby blood vessel. The osteoblasts are inactive in this stage. In regards to the kidneys, PTH stimulates kidney tubule cells to recover waste calcium from the urine. PTH also stimulates kidney tubule cells to release calcitrol. This is illustrated with a cross section of a kidney tubule, showing the cells of the tubule wall. Urine is running to the left of the tubule wall cells while an artery is to the right. The right edge of the tubule wall cells and the left edge of the artery are separated by a small region of interstitial space. The cells are removing calcium from the urine and pumping it into the interstitial fluid, after which the calcium enters the artery. The cells are also pumping calcitrol into the blood vessel. In regards to the intestine, PTH stimulates the intestines to absorb calcium from digesting food. A cross section of an intestinal cell is shown, which is cube-shaped but with finger-like projections on the intestinal lumen side (top). Beneath the intestinal cell is an artery. Calcitrol is leaving the artery and entering the intestinal cell, stimulating it to absorb calcium from food in the intestinal lumen. The effects of PTH on bone, the kidneys and the intestines all cause blood calcium levels to increase. High calcium concentrations in the blood stimulate the parafollicular cells in the thyroid to release calcitonin. Calcitonin reverses the effects of PTH by stimulating osteoblasts and inhibiting osteoclasts in bone tissue. This is illustrated by calcium ions leaving a blood vessel and traveling to osteoblasts on a section of compact bone. The osteoblasts are thickening the compact bone layer while, in this stage, the osteoclasts are inactive. Figure 2. Parathyroid Hormone in Maintaining Blood Calcium Homeostasis. Parathyroid hormone increases blood calcium levels when they drop too low. Conversely, calcitonin, which is released from the thyroid gland, decreases blood calcium levels when they become too high. These two mechanisms constantly maintain blood calcium concentration at homeostasis.[/caption]
Abnormally high activity of the parathyroid gland can cause hyperparathyroidism, a disorder caused by an overproduction of PTH that results in excessive calcium reabsorption from bone. Hyperparathyroidism can significantly decrease bone density, leading to spontaneous fractures or deformities. As blood calcium levels rise, cell membrane permeability to sodium is decreased, and the responsiveness of the nervous system is reduced. At the same time, calcium deposits may collect in the body’s tissues and organs, impairing their functioning. In contrast, abnormally low blood calcium levels may be caused by parathyroid hormone deficiency, called hypoparathyroidism, which may develop following injury or surgery involving the thyroid gland. Low blood calcium increases membrane permeability to sodium, resulting in muscle twitching, cramping, spasms, or convulsions. Severe deficits can paralyze muscles, including those involved in breathing, and can be fatal.

When blood calcium levels are high, calcitonin is produced and secreted by the parafollicular cells of the thyroid gland. As discussed earlier, calcitonin inhibits the activity of osteoclasts, reduces the absorption of dietary calcium in the intestine, and signals the kidneys to reabsorb less calcium, resulting in larger amounts of calcium excreted in the urine.

]]> 2606 2585 5 0 17.6 The Adrenal Glands https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2608 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2608

Learning Objectives

By the end of this section, you will be able to:
  • Describe the location and structure of the adrenal glands
  • Identify the hormones produced by the adrenal cortex and adrenal medulla, and summarize their target cells and effects

The adrenal glands are wedges of glandular and neuroendocrine tissue adhering to the top of the kidneys by a fibrous capsule (Figure 1). The adrenal glands have a rich blood supply and experience one of the highest rates of blood flow in the body. They are served by several arteries branching off the aorta, including the suprarenal and renal arteries. Blood flows to each adrenal gland at the adrenal cortex and then drains into the adrenal medulla. Adrenal hormones are released into the circulation via the left and right suprarenal veins.

[caption id="" align="aligncenter" width="550"]This diagram shows the left adrenal gland located atop the left kidney. The gland is composed of an outer cortex and an inner medulla all surrounded by a connective tissue capsule. The cortex can be subdivided into additional zones, all of which produce different types of hormones. The outermost layer is the zona glomerulosa, which releases mineralcorticoids, such as aldosterone, that regulate mineral balance. Underneath this layer is the zona fasciculate, which releases glucocorticoids, such as cortisol, corticosterone and cortisone, that regulate glucose metabolism. Underneath this layer is the zona reticularis, which releases androgens, such as dehydroepiandrosterone, that stimulate masculinization. Below this layer is the adrenal medulla, which releases stress hormones, such as epinephrine and norepinephrine, that stimulate the symphathetic ANS. Figure 1. Adrenal Glands. Both adrenal glands sit atop the kidneys and are composed of an outer cortex and an inner medulla, all surrounded by a connective tissue capsule. The cortex can be subdivided into additional zones, all of which produce different types of hormones. LM × 204. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

The adrenal gland consists of an outer cortex of glandular tissue and an inner medulla of nervous tissue. The cortex itself is divided into three zones: the zona glomerulosa, the zona fasciculata, and the zona reticularis. Each region secretes its own set of hormones.

The adrenal cortex, as a component of the hypothalamic-pituitary-adrenal (HPA) axis, secretes steroid hormones important for the regulation of the long-term stress response, blood pressure and blood volume, nutrient uptake and storage, fluid and electrolyte balance, and inflammation. The HPA axis involves the stimulation of hormone release of adrenocorticotropic hormone (ACTH) from the pituitary by the hypothalamus. ACTH then stimulates the adrenal cortex to produce the hormone cortisol. This pathway will be discussed in more detail below.

The adrenal medulla is neuroendocrine tissue composed of postganglionic sympathetic nervous system (SNS) neurons. It is really an extension of the autonomic nervous system, which regulates homeostasis in the body. The sympathomedullary (SAM) pathway involves the stimulation of the medulla by impulses from the hypothalamus via neurons from the thoracic spinal cord. The medulla is stimulated to secrete the amine hormones epinephrine and norepinephrine.

One of the major functions of the adrenal gland is to respond to stress. Stress can be either physical or psychological or both. Physical stresses include exposing the body to injury, walking outside in cold and wet conditions without a coat on, or malnutrition. Psychological stresses include the perception of a physical threat, a fight with a loved one, or just a bad day at school.

The body responds in different ways to short-term stress and long-term stress following a pattern known as the general adaptation syndrome (GAS). Stage one of GAS is called the alarm reaction. This is short-term stress, the fight-or-flight response, mediated by the hormones epinephrine and norepinephrine from the adrenal medulla via the SAM pathway. Their function is to prepare the body for extreme physical exertion. Once this stress is relieved, the body quickly returns to normal. The section on the adrenal medulla covers this response in more detail.

If the stress is not soon relieved, the body adapts to the stress in the second stage called the stage of resistance. If a person is starving for example, the body may send signals to the gastrointestinal tract to maximize the absorption of nutrients from food.

If the stress continues for a longer term however, the body responds with symptoms quite different than the fight-or-flight response. During the stage of exhaustion, individuals may begin to suffer depression, the suppression of their immune response, severe fatigue, or even a fatal heart attack. These symptoms are mediated by the hormones of the adrenal cortex, especially cortisol, released as a result of signals from the HPA axis.

Adrenal hormones also have several non–stress-related functions, including the increase of blood sodium and glucose levels, which will be described in detail below.

Adrenal Cortex

The adrenal cortex consists of multiple layers of lipid-storing cells that occur in three structurally distinct regions. Each of these regions produces different hormones.

Hormones of the Zona Glomerulosa

The most superficial region of the adrenal cortex is the zona glomerulosa, which produces a group of hormones collectively referred to as mineralocorticoids because of their effect on body minerals, especially sodium and potassium. These hormones are essential for fluid and electrolyte balance.

Aldosterone is the major mineralocorticoid. It is important in the regulation of the concentration of sodium and potassium ions in urine, sweat, and saliva. For example, it is released in response to elevated blood K+, low blood Na+, low blood pressure, or low blood volume. In response, aldosterone increases the excretion of K+ and the retention of Na+, which in turn increases blood volume and blood pressure. Its secretion is prompted when CRH from the hypothalamus triggers ACTH release from the anterior pituitary.

Aldosterone is also a key component of the renin-angiotensin-aldosterone system (RAAS) in which specialized cells of the kidneys secrete the enzyme renin in response to low blood volume or low blood pressure. Renin then catalyzes the conversion of the blood protein angiotensinogen, produced by the liver, to the hormone angiotensin I. Angiotensin I is converted in the lungs to angiotensin II by angiotensin-converting enzyme (ACE). Angiotensin II has three major functions:

  1. Initiating vasoconstriction of the arterioles, decreasing blood flow
  2. Stimulating kidney tubules to reabsorb NaCl and water, increasing blood volume
  3. Signaling the adrenal cortex to secrete aldosterone, the effects of which further contribute to fluid retention, restoring blood pressure and blood volume

For individuals with hypertension, or high blood pressure, drugs are available that block the production of angiotensin II. These drugs, known as ACE inhibitors, block the ACE enzyme from converting angiotensin I to angiotensin II, thus mitigating the latter’s ability to increase blood pressure.

Hormones of the Zona Fasciculata

The intermediate region of the adrenal cortex is the zona fasciculata, named as such because the cells form small fascicles (bundles) separated by tiny blood vessels. The cells of the zona fasciculata produce hormones called glucocorticoids because of their role in glucose metabolism. The most important of these is cortisol, some of which the liver converts to cortisone. A glucocorticoid produced in much smaller amounts is corticosterone. In response to long-term stressors, the hypothalamus secretes CRH, which in turn triggers the release of ACTH by the anterior pituitary. ACTH triggers the release of the glucocorticoids. Their overall effect is to inhibit tissue building while stimulating the breakdown of stored nutrients to maintain adequate fuel supplies. In conditions of long-term stress, for example, cortisol promotes the catabolism of glycogen to glucose, the catabolism of stored triglycerides into fatty acids and glycerol, and the catabolism of muscle proteins into amino acids. These raw materials can then be used to synthesize additional glucose and ketones for use as body fuels. The hippocampus, which is part of the temporal lobe of the cerebral cortices and important in memory formation, is highly sensitive to stress levels because of its many glucocorticoid receptors.

You are probably familiar with prescription and over-the-counter medications containing glucocorticoids, such as cortisone injections into inflamed joints, prednisone tablets and steroid-based inhalers used to manage severe asthma, and hydrocortisone creams applied to relieve itchy skin rashes. These drugs reflect another role of cortisol—the downregulation of the immune system, which inhibits the inflammatory response.

Hormones of the Zona Reticularis

The deepest region of the adrenal cortex is the zona reticularis, which produces small amounts of a class of steroid sex hormones called androgens. During puberty and most of adulthood, androgens are produced in the gonads. The androgens produced in the zona reticularis supplement the gonadal androgens. They are produced in response to ACTH from the anterior pituitary and are converted in the tissues to testosterone or estrogens. In adult women, they may contribute to the sex drive, but their function in adult men is not well understood. In post-menopausal women, as the functions of the ovaries decline, the main source of estrogens becomes the androgens produced by the zona reticularis.

Adrenal Medulla

As noted earlier, the adrenal cortex releases glucocorticoids in response to long-term stress such as severe illness. In contrast, the adrenal medulla releases its hormones in response to acute, short-term stress mediated by the sympathetic nervous system (SNS).

The medullary tissue is composed of unique postganglionic SNS neurons called chromaffin cells, which are large and irregularly shaped, and produce the neurotransmitters epinephrine (also called adrenaline) and norepinephrine (or noradrenaline). Epinephrine is produced in greater quantities—approximately a 4 to 1 ratio with norepinephrine—and is the more powerful hormone. Because the chromaffin cells release epinephrine and norepinephrine into the systemic circulation, where they travel widely and exert effects on distant cells, they are considered hormones. Derived from the amino acid tyrosine, they are chemically classified as catecholamines.

The secretion of medullary epinephrine and norepinephrine is controlled by a neural pathway that originates from the hypothalamus in response to danger or stress (the SAM pathway). Both epinephrine and norepinephrine signal the liver and skeletal muscle cells to convert glycogen into glucose, resulting in increased blood glucose levels. These hormones increase the heart rate, pulse, and blood pressure to prepare the body to fight the perceived threat or flee from it. In addition, the pathway dilates the airways, raising blood oxygen levels. It also prompts vasodilation, further increasing the oxygenation of important organs such as the lungs, brain, heart, and skeletal muscle. At the same time, it triggers vasoconstriction to blood vessels serving less essential organs such as the gastrointestinal tract, kidneys, and skin, and downregulates some components of the immune system. Other effects include a dry mouth, loss of appetite, pupil dilation, and a loss of peripheral vision. The major hormones of the adrenal glands are summarized in Table 5.

Hormones of the Adrenal Glands (Table 5)
Adrenal gland Associated hormones Chemical class Effect
Adrenal cortex Aldosterone Steroid Increases blood Na+ levels
Adrenal cortex Cortisol, corticosterone, cortisone Steroid Increase blood glucose levels
Adrenal medulla Epinephrine, norepinephrine Amine Stimulate fight-or-flight response

Disorders Involving the Adrenal Glands

Several disorders are caused by the dysregulation of the hormones produced by the adrenal glands. For example, Cushing’s disease is a disorder characterized by high blood glucose levels and the accumulation of lipid deposits on the face and neck. It is caused by hypersecretion of cortisol. The most common source of Cushing’s disease is a pituitary tumor that secretes cortisol or ACTH in abnormally high amounts. Other common signs of Cushing’s disease include the development of a moon-shaped face, a buffalo hump on the back of the neck, rapid weight gain, and hair loss. Chronically elevated glucose levels are also associated with an elevated risk of developing type 2 diabetes. In addition to hyperglycemia, chronically elevated glucocorticoids compromise immunity, resistance to infection, and memory, and can result in rapid weight gain and hair loss.

In contrast, the hyposecretion of corticosteroids can result in Addison’s disease, a rare disorder that causes low blood glucose levels and low blood sodium levels. The signs and symptoms of Addison’s disease are vague and are typical of other disorders as well, making diagnosis difficult. They may include general weakness, abdominal pain, weight loss, nausea, vomiting, sweating, and cravings for salty food.

]]> 2608 2585 6 0 17.7 The Pineal Gland https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2609 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2609

Learning Objectives

By the end of this section, you will be able to:
  • Describe the location and structure of the pineal gland
  • Discuss the function of melatonin

Recall that the hypothalamus, part of the diencephalon of the brain, sits inferior and somewhat anterior to the thalamus. Inferior but somewhat posterior to the thalamus is the pineal gland, a tiny endocrine gland whose functions are not entirely clear. The pinealocyte cells that make up the pineal gland are known to produce and secrete the amine hormone melatonin, which is derived from serotonin.

The secretion of melatonin varies according to the level of light received from the environment. When photons of light stimulate the retinas of the eyes, a nerve impulse is sent to a region of the hypothalamus called the suprachiasmatic nucleus (SCN), which is important in regulating biological rhythms. From the SCN, the nerve signal is carried to the spinal cord and eventually to the pineal gland, where the production of melatonin is inhibited. As a result, blood levels of melatonin fall, promoting wakefulness. In contrast, as light levels decline—such as during the evening—melatonin production increases, boosting blood levels and causing drowsiness.

The secretion of melatonin may influence the body’s circadian rhythms, the dark-light fluctuations that affect not only sleepiness and wakefulness, but also appetite and body temperature. Interestingly, children have higher melatonin levels than adults, which may prevent the release of gonadotropins from the anterior pituitary, thereby inhibiting the onset of puberty. Finally, an antioxidant role of melatonin is the subject of current research.

Jet lag occurs when a person travels across several time zones and feels sleepy during the day or wakeful at night. Traveling across multiple time zones significantly disturbs the light-dark cycle regulated by melatonin. It can take up to several days for melatonin synthesis to adjust to the light-dark patterns in the new environment, resulting in jet lag. Some air travelers take melatonin supplements to induce sleep.]]> 2609 2585 7 0 17.8 Gonadal and Placental Hormones https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2610 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2610

Learning Objectives

By the end of this section, you will be able to:
  • Identify the most important hormones produced by the testes and ovaries
  • Name the hormones produced by the placenta and state their functions

This section briefly discusses the hormonal role of the gonads—the male testes and female ovaries—which produce the sex cells (sperm and ova) and secrete the gonadal hormones. The roles of the gonadotropins released from the anterior pituitary (FSH and LH) were discussed earlier.

The primary hormone produced by the male testes is testosterone, a steroid hormone important in the development of the male reproductive system, the maturation of sperm cells, and the development of male secondary sex characteristics such as a deepened voice, body hair, and increased muscle mass. Interestingly, testosterone is also produced in the female ovaries, but at a much reduced level. In addition, the testes produce the peptide hormone inhibin, which inhibits the secretion of FSH from the anterior pituitary gland. FSH stimulates spermatogenesis.

The primary hormones produced by the ovaries are estrogens, which include estradiol, estriol, and estrone. Estrogens play an important role in a larger number of physiological processes, including the development of the female reproductive system, regulation of the menstrual cycle, the development of female secondary sex characteristics such as increased adipose tissue and the development of breast tissue, and the maintenance of pregnancy. Another significant ovarian hormone is progesterone, which contributes to regulation of the menstrual cycle and is important in preparing the body for pregnancy as well as maintaining pregnancy. In addition, the granulosa cells of the ovarian follicles produce inhibin, which—as in males—inhibits the secretion of FSH.During the initial stages of pregnancy, an organ called the placenta develops within the uterus. The placenta supplies oxygen and nutrients to the fetus, excretes waste products, and produces and secretes estrogens and progesterone. The placenta produces human chorionic gonadotropin (hCG) as well. The hCG hormone promotes progesterone synthesis and reduces the mother’s immune function to protect the fetus from immune rejection. It also secretes human placental lactogen (hPL), which plays a role in preparing the breasts for lactation, and relaxin, which is thought to help soften and widen the pubic symphysis in preparation for childbirth. The hormones controlling reproduction are summarized in Table 6.

Reproductive Hormones (Table 6)
Gonad Associated hormones Chemical class Effect
Testes Testosterone Steroid Stimulates development of male secondary sex characteristics and sperm production
Testes Inhibin Protein Inhibits FSH release from pituitary
Ovaries Estrogens and progesterone Steroid Stimulate development of female secondary sex characteristics and prepare the body for childbirth
Placenta Human chorionic gonadotropin Protein Promotes progesterone synthesis during pregnancy and inhibits immune response against fetus
Everyday Connections

Anabolic SteroidsThe endocrine system can be exploited for illegal or unethical purposes. A prominent example of this is the use of steroid drugs by professional athletes.

Commonly used for performance enhancement, anabolic steroids are synthetic versions of the male sex hormone, testosterone. By boosting natural levels of this hormone, athletes experience increased muscle mass. Synthetic versions of human growth hormone are also used to build muscle mass.

The use of performance-enhancing drugs is banned by all major collegiate and professional sports organizations in the United States because they impart an unfair advantage to athletes who take them. In addition, the drugs can cause significant and dangerous side effects. For example, anabolic steroid use can increase cholesterol levels, raise blood pressure, and damage the liver. Altered testosterone levels (both too low or too high) have been implicated in causing structural damage to the heart, and increasing the risk for cardiac arrhythmias, heart attacks, congestive heart failure, and sudden death. Paradoxically, steroids can have a feminizing effect in males, including shriveled testicles and enlarged breast tissue. In females, their use can cause masculinizing effects such as an enlarged clitoris and growth of facial hair. In both sexes, their use can promote increased aggression (commonly known as “roid-rage”), depression, sleep disturbances, severe acne, and infertility.

]]> 2610 2585 8 0 17.9 The Endocrine Pancreas https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2613 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2613

Learning Objectives

By the end of this section, you will be able to:
  • Describe the location and structure of the pancreas, and the morphology and function of the pancreatic islets
  • Compare and contrast the functions of insulin and glucagon

The pancreas is a long, slender organ, most of which is located posterior to the bottom half of the stomach (Figure 1). Although it is primarily an exocrine gland, secreting a variety of digestive enzymes, the pancreas has an endocrine function. Its pancreatic islets—clusters of cells formerly known as the islets of Langerhans—secrete the hormones glucagon, insulin, somatostatin, and pancreatic polypeptide (PP).

[caption id="" align="aligncenter" width="500"]This diagram shows the anatomy of the pancreas. The left, larger side of the pancreas is seated within the curve of the duodenum of the small intestine. The smaller, rightmost tip of the pancreas is located near the spleen. The splenic artery is seen travelling to the spleen, however, it has several branches connecting to the pancreas. An interior view of the pancreas shows that the pancreatic duct is a large tube running through the center of the pancreas. It branches throughout its length in to several horseshoe- shaped pockets of acinar cells. These cells secrete digestive enzymes, which travel down the bile duct and into the small intestine. There are also small pancreatic islets scattered throughout the pancreas. The pancreatic islets secrete the pancreatic hormones insulin and glucagon into the splenic artery. An inset micrograph shows that the pancreatic islets are small discs of tissue consisting of a thin, outer ring called the exocrine acinus, a thicker, inner ring of beta cells and a central circle of alpha cells. Figure 1. Pancreas. The pancreatic exocrine function involves the acinar cells secreting digestive enzymes that are transported into the small intestine by the pancreatic duct. Its endocrine function involves the secretion of insulin (produced by beta cells) and glucagon (produced by alpha cells) within the pancreatic islets. These two hormones regulate the rate of glucose metabolism in the body. The micrograph reveals pancreatic islets. LM × 760. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Cells and Secretions of the Pancreatic Islets

The pancreatic islets each contain four varieties of cells:

  • The alpha cell produces the hormone glucagon and makes up approximately 20 percent of each islet. Glucagon plays an important role in blood glucose regulation; low blood glucose levels stimulate its release.
  • The beta cell produces the hormone insulin and makes up approximately 75 percent of each islet. Elevated blood glucose levels stimulate the release of insulin.
  • The delta cell accounts for four percent of the islet cells and secretes the peptide hormone somatostatin. Recall that somatostatin is also released by the hypothalamus (as GHIH), and the stomach and intestines also secrete it. An inhibiting hormone, pancreatic somatostatin inhibits the release of both glucagon and insulin.
  • The PP cell accounts for about one percent of islet cells and secretes the pancreatic polypeptide hormone. It is thought to play a role in appetite, as well as in the regulation of pancreatic exocrine and endocrine secretions. Pancreatic polypeptide released following a meal may reduce further food consumption; however, it is also released in response to fasting.

Regulation of Blood Glucose Levels by Insulin and Glucagon

Glucose is required for cellular respiration and is the preferred fuel for all body cells. The body derives glucose from the breakdown of the carbohydrate-containing foods and drinks we consume. Glucose not immediately taken up by cells for fuel can be stored by the liver and muscles as glycogen, or converted to triglycerides and stored in the adipose tissue. Hormones regulate both the storage and the utilization of glucose as required. Receptors located in the pancreas sense blood glucose levels, and subsequently the pancreatic cells secrete glucagon or insulin to maintain normal levels.

Glucagon

Receptors in the pancreas can sense the decline in blood glucose levels, such as during periods of fasting or during prolonged labor or exercise (Figure 2). In response, the alpha cells of the pancreas secrete the hormone glucagon, which has several effects:

  • It stimulates the liver to convert its stores of glycogen back into glucose. This response is known as glycogenolysis. The glucose is then released into the circulation for use by body cells.
  • It stimulates the liver to take up amino acids from the blood and convert them into glucose. This response is known as gluconeogenesis.
  • It stimulates lipolysis, the breakdown of stored triglycerides into free fatty acids and glycerol. Some of the free glycerol released into the bloodstream travels to the liver, which converts it into glucose. This is also a form of gluconeogenesis.

Taken together, these actions increase blood glucose levels. The activity of glucagon is regulated through a negative feedback mechanism; rising blood glucose levels inhibit further glucagon production and secretion.

[caption id="" align="aligncenter" width="550"]This diagram shows the homeostatic regulation of blood glucose levels. Blood glucose concentration is tightly maintained between 70 milligrams per deciliter and 110 milligrams per deciliter. If blood glucose concentration rises above this range (hyperglycemia), insulin is released from the pancreas. Insulin triggers body cells to take up glucose from the blood and utilize it in cellular respiration. Insulin also inhibits glycogenolysis, in that glucose is removed from the blood and stored as glycogen in the liver. Insulin also inhibits gluconeogenesis, in that amino acids and free glycerol are not converted to glucose in the ER. If blood glucose concentration drops below this range, glucagon is released, which stimulates body cells to release glucose into the blood. All of these actions cause blood glucose concentration to decrease. When blood glucose concentration is low (hypoglycemia), alpha cells of the pancreas release glucagon. Glucagon inhibits body cells from taking up glucose from the blood and utilizing it in cellular respiration. Glucagon also stimulates glycogenolysis, in that glycogen in the liver is broken down into glucose and released into the blood. Glucagon also stimulates glucogenogenesis, in that amino acids and free glycerol are converted to glucose in the ER and released into the blood. All of these actions cause blood glucose concentrations to increase. Figure 2. Homeostatic Regulation of Blood Glucose Levels. Blood glucose concentration is tightly maintained between 70 mg/dL and 110 mg/dL. If blood glucose concentration rises above this range, insulin is released, which stimulates body cells to remove glucose from the blood. If blood glucose concentration drops below this range, glucagon is released, which stimulates body cells to release glucose into the blood.[/caption]

Insulin

The primary function of insulin is to facilitate the uptake of glucose into body cells. Red blood cells, as well as cells of the brain, liver, kidneys, and the lining of the small intestine, do not have insulin receptors on their cell membranes and do not require insulin for glucose uptake. Although all other body cells do require insulin if they are to take glucose from the bloodstream, skeletal muscle cells and adipose cells are the primary targets of insulin.

The presence of food in the intestine triggers the release of gastrointestinal tract hormones such as glucose-dependent insulinotropic peptide (previously known as gastric inhibitory peptide). This is in turn the initial trigger for insulin production and secretion by the beta cells of the pancreas. Once nutrient absorption occurs, the resulting surge in blood glucose levels further stimulates insulin secretion.

Precisely how insulin facilitates glucose uptake is not entirely clear. However, insulin appears to activate a tyrosine kinase receptor, triggering the phosphorylation of many substrates within the cell. These multiple biochemical reactions converge to support the movement of intracellular vesicles containing facilitative glucose transporters to the cell membrane. In the absence of insulin, these transport proteins are normally recycled slowly between the cell membrane and cell interior. Insulin triggers the rapid movement of a pool of glucose transporter vesicles to the cell membrane, where they fuse and expose the glucose transporters to the extracellular fluid. The transporters then move glucose by facilitated diffusion into the cell interior.

Insulin also reduces blood glucose levels by stimulating glycolysis, the metabolism of glucose for generation of ATP. Moreover, it stimulates the liver to convert excess glucose into glycogen for storage, and it inhibits enzymes involved in glycogenolysis and gluconeogenesis. Finally, insulin promotes triglyceride and protein synthesis. The secretion of insulin is regulated through a negative feedback mechanism. As blood glucose levels decrease, further insulin release is inhibited. The pancreatic hormones are summarized in Table 7.

Hormones of the Pancreas (Table 7)
Associated hormones Chemical class Effect
Insulin (beta cells) Protein Reduces blood glucose levels
Glucagon (alpha cells) Protein Increases blood glucose levels
Somatostatin (delta cells) Protein Inhibits insulin and glucagon release
Pancreatic polypeptide (PP cells) Protein Role in appetite
Disorders of the…

Endocrine System: Diabetes Mellitus Dysfunction of insulin production and secretion, as well as the target cells’ responsiveness to insulin, can lead to a condition called diabetes mellitus. An increasingly common disease, diabetes mellitus has been diagnosed in more than 18 million adults in the United States, and more than 200,000 children. It is estimated that up to 7 million more adults have the condition but have not been diagnosed. In addition, approximately 79 million people in the US are estimated to have pre-diabetes, a condition in which blood glucose levels are abnormally high, but not yet high enough to be classified as diabetes.

There are two main forms of diabetes mellitus. Type 1 diabetes is an autoimmune disease affecting the beta cells of the pancreas. Certain genes are recognized to increase susceptibility. The beta cells of people with type 1 diabetes do not produce insulin; thus, synthetic insulin must be administered by injection or infusion. This form of diabetes accounts for less than five percent of all diabetes cases.

Type 2 diabetes accounts for approximately 95 percent of all cases. It is acquired, and lifestyle factors such as poor diet, inactivity, and the presence of pre-diabetes greatly increase a person’s risk. About 80 to 90 percent of people with type 2 diabetes are overweight or obese. In type 2 diabetes, cells become resistant to the effects of insulin. In response, the pancreas increases its insulin secretion, but over time, the beta cells become exhausted. In many cases, type 2 diabetes can be reversed by moderate weight loss, regular physical activity, and consumption of a healthy diet; however, if blood glucose levels cannot be controlled, the diabetic will eventually require insulin.

Two of the early manifestations of diabetes are excessive urination and excessive thirst. They demonstrate how the out-of-control levels of glucose in the blood affect kidney function. The kidneys are responsible for filtering glucose from the blood. Excessive blood glucose draws water into the urine, and as a result the person eliminates an abnormally large quantity of sweet urine. The use of body water to dilute the urine leaves the body dehydrated, and so the person is unusually and continually thirsty. The person may also experience persistent hunger because the body cells are unable to access the glucose in the bloodstream.

Over time, persistently high levels of glucose in the blood injure tissues throughout the body, especially those of the blood vessels and nerves. Inflammation and injury of the lining of arteries lead to atherosclerosis and an increased risk of heart attack and stroke. Damage to the microscopic blood vessels of the kidney impairs kidney function and can lead to kidney failure. Damage to blood vessels that serve the eyes can lead to blindness. Blood vessel damage also reduces circulation to the limbs, whereas nerve damage leads to a loss of sensation, called neuropathy, particularly in the hands and feet. Together, these changes increase the risk of injury, infection, and tissue death (necrosis), contributing to a high rate of toe, foot, and lower leg amputations in people with diabetes. Uncontrolled diabetes can also lead to a dangerous form of metabolic acidosis called ketoacidosis. Deprived of glucose, cells increasingly rely on fat stores for fuel. However, in a glucose-deficient state, the liver is forced to use an alternative lipid metabolism pathway that results in the increased production of ketone bodies (or ketones), which are acidic. The build-up of ketones in the blood causes ketoacidosis, which—if left untreated—may lead to a life-threatening “diabetic coma.” Together, these complications make diabetes the seventh leading cause of death in the United States.

Diabetes is diagnosed when lab tests reveal that blood glucose levels are higher than normal, a condition called hyperglycemia. The treatment of diabetes depends on the type, the severity of the condition, and the ability of the patient to make lifestyle changes. As noted earlier, moderate weight loss, regular physical activity, and consumption of a healthful diet can reduce blood glucose levels. Some patients with type 2 diabetes may be unable to control their disease with these lifestyle changes, and will require medication. Historically, the first-line treatment of type 2 diabetes was insulin. Research advances have resulted in alternative options, including medications that enhance pancreatic function.

]]> 2613 2585 9 0 17.10 Organs with Secondary Endocrine Functions https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2614 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2614

Learning Objectives

By the end of this section, you will be able to:
  • Identify the organs with a secondary endocrine function, the hormone they produce, and its effects

In your study of anatomy and physiology, you have already encountered a few of the many organs of the body that have secondary endocrine functions. Here, you will learn about the hormone-producing activities of the heart, gastrointestinal tract, kidneys, skeleton, adipose tissue, skin, and thymus.

Heart

When the body experiences an increase in blood volume or pressure, the cells of the heart’s atrial wall stretch. In response, specialized cells in the wall of the atria produce and secrete the peptide hormone atrial natriuretic peptide (ANP). ANP signals the kidneys to reduce sodium reabsorption, thereby decreasing the amount of water reabsorbed from the urine filtrate and reducing blood volume. Other actions of ANP include the inhibition of renin secretion and the initiation of the renin-angiotensin-aldosterone system (RAAS) and vasodilation. Therefore, ANP aids in decreasing blood pressure, blood volume, and blood sodium levels.

Gastrointestinal Tract

The endocrine cells of the GI tract are located in the mucosa of the stomach and small intestine. Some of these hormones are secreted in response to eating a meal and aid in digestion. An example of a hormone secreted by the stomach cells is gastrin, a peptide hormone secreted in response to stomach distention that stimulates the release of hydrochloric acid. Secretin is a peptide hormone secreted by the small intestine as acidic chyme (partially digested food and fluid) moves from the stomach. It stimulates the release of bicarbonate from the pancreas, which buffers the acidic chyme, and inhibits the further secretion of hydrochloric acid by the stomach. Cholecystokinin (CCK) is another peptide hormone released from the small intestine. It promotes the secretion of pancreatic enzymes and the release of bile from the gallbladder, both of which facilitate digestion. Other hormones produced by the intestinal cells aid in glucose metabolism, such as by stimulating the pancreatic beta cells to secrete insulin, reducing glucagon secretion from the alpha cells, or enhancing cellular sensitivity to insulin.

Kidneys

The kidneys participate in several complex endocrine pathways and produce certain hormones. A decline in blood flow to the kidneys stimulates them to release the enzyme renin, triggering the renin-angiotensin-aldosterone (RAAS) system, and stimulating the reabsorption of sodium and water. The reabsorption increases blood flow and blood pressure. The kidneys also play a role in regulating blood calcium levels through the production of calcitriol from vitamin D3, which is released in response to the secretion of parathyroid hormone (PTH). In addition, the kidneys produce the hormone erythropoietin (EPO) in response to low oxygen levels. EPO stimulates the production of red blood cells (erythrocytes) in the bone marrow, thereby increasing oxygen delivery to tissues. You may have heard of EPO as a performance-enhancing drug (in a synthetic form).

Skeleton

Although bone has long been recognized as a target for hormones, only recently have researchers recognized that the skeleton itself produces at least two hormones. Fibroblast growth factor 23 (FGF23) is produced by bone cells in response to increased blood levels of vitamin D3 or phosphate. It triggers the kidneys to inhibit the formation of calcitriol from vitamin D3 and to increase phosphorus excretion. Osteocalcin, produced by osteoblasts, stimulates the pancreatic beta cells to increase insulin production. It also acts on peripheral tissues to increase their sensitivity to insulin and their utilization of glucose.

Adipose Tissue

Adipose tissue produces and secretes several hormones involved in lipid metabolism and storage. One important example is leptin, a protein manufactured by adipose cells that circulates in amounts directly proportional to levels of body fat. Leptin is released in response to food consumption and acts by binding to brain neurons involved in energy intake and expenditure. Binding of leptin produces a feeling of satiety after a meal, thereby reducing appetite. It also appears that the binding of leptin to brain receptors triggers the sympathetic nervous system to regulate bone metabolism, increasing deposition of cortical bone. Adiponectin—another hormone synthesized by adipose cells—appears to reduce cellular insulin resistance and to protect blood vessels from inflammation and atherosclerosis. Its levels are lower in people who are obese, and rise following weight loss.

Skin

The skin functions as an endocrine organ in the production of the inactive form of vitamin D3, cholecalciferol. When cholesterol present in the epidermis is exposed to ultraviolet radiation, it is converted to cholecalciferol, which then enters the blood. In the liver, cholecalciferol is converted to an intermediate that travels to the kidneys and is further converted to calcitriol, the active form of vitamin D3. Vitamin D is important in a variety of physiological processes, including intestinal calcium absorption and immune system function. In some studies, low levels of vitamin D have been associated with increased risks of cancer, severe asthma, and multiple sclerosis. Vitamin D deficiency in children causes rickets, and in adults, osteomalacia—both of which are characterized by bone deterioration.

Thymus

The thymus is an organ of the immune system that is larger and more active during infancy and early childhood, and begins to atrophy as we age. Its endocrine function is the production of a group of hormones called thymosins that contribute to the development and differentiation of T lymphocytes, which are immune cells. Although the role of thymosins is not yet well understood, it is clear that they contribute to the immune response. Thymosins have been found in tissues other than the thymus and have a wide variety of functions, so the thymosins cannot be strictly categorized as thymic hormones.

Liver

The liver is responsible for secreting at least four important hormones or hormone precursors: insulin-like growth factor (somatomedin), angiotensinogen, thrombopoetin, and hepcidin. Insulin-like growth factor-1 is the immediate stimulus for growth in the body, especially of the bones. Angiotensinogen is the precursor to angiotensin, mentioned earlier, which increases blood pressure. Thrombopoetin stimulates the production of the blood’s platelets. Hepcidins block the release of iron from cells in the body, helping to regulate iron homeostasis in our body fluids. The major hormones of these other organs are summarized in Table 8.

Organs with Secondary Endocrine Functions and Their Major Hormones (Table 8)
Organ Major hormones Effects
Heart Atrial natriuretic peptide (ANP) Reduces blood volume, blood pressure, and Na+ concentration
Gastrointestinal tract Gastrin, secretin, and cholecystokinin Aid digestion of food and buffering of stomach acids
Gastrointestinal tract Glucose-dependent insulinotropic peptide (GIP) and glucagon-like peptide 1 (GLP-1) Stimulate beta cells of the pancreas to release insulin
Kidneys Renin Stimulates release of aldosterone
Kidneys Calcitriol Aids in the absorption of Ca2+
Kidneys Erythropoietin Triggers the formation of red blood cells in the bone marrow
Skeleton FGF23 Inhibits production of calcitriol and increases phosphate excretion
Skeleton Osteocalcin Increases insulin production
Adipose tissue Leptin Promotes satiety signals in the brain
Adipose tissue Adiponectin Reduces insulin resistance
Skin Cholecalciferol Modified to form vitamin D
Thymus (and other organs) Thymosins Among other things, aids in the development of T lymphocytes of the immune system
Liver Insulin-like growth factor-1 Stimulates bodily growth
Liver Angiotensinogen Raises blood pressure
Liver Thrombopoetin Causes increase in platelets
Liver Hepcidin Blocks release of iron into body fluids
]]> 2614 2585 10 0 24.2 Carbohydrate Metabolism https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2646 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2646

Learning Objectives

By the end of this section, you will be able to:
  • Explain the processes of glycolysis
  • Describe the pathway of a pyruvate molecule through the Krebs cycle
  • Explain the transport of electrons through the electron transport chain
  • Describe the process of ATP production through oxidative phosphorylation
  • Summarize the process of gluconeogenesis

Carbohydrates are organic molecules composed of carbon, hydrogen, and oxygen atoms. The family of carbohydrates includes both simple and complex sugars. Glucose and fructose are examples of simple sugars, and starch, glycogen, and cellulose are all examples of complex sugars. The complex sugars are also called polysaccharides and are made of multiple monosaccharide molecules. Polysaccharides serve as energy storage (e.g., starch and glycogen) and as structural components (e.g., chitin in insects and cellulose in plants).

During digestion, carbohydrates are broken down into simple, soluble sugars that can be transported across the intestinal wall into the circulatory system to be transported throughout the body. Carbohydrate digestion begins in the mouth with the action of salivary amylase on starches and ends with monosaccharides being absorbed across the epithelium of the small intestine. Once the absorbed monosaccharides are transported to the tissues, the process of cellular respiration begins (Figure 1). This section will focus first on glycolysis, a process where the monosaccharide glucose is oxidized, releasing the energy stored in its bonds to produce ATP.

[caption id="" align="aligncenter" width="520"]This figure shows the different pathways of cellular respiration. The pathways shown are glycolysis, the pyruvic acid cycle, the Krebs cycle, and oxidative phosphorylation. Figure 1. Cellular Respiration. Cellular respiration oxidizes glucose molecules through glycolysis, the Krebs cycle, and oxidative phosphorylation to produce ATP.[/caption]

Glycolysis

Glucose is the body’s most readily available source of energy. After digestive processes break polysaccharides down into monosaccharides, including glucose, the monosaccharides are transported across the wall of the small intestine and into the circulatory system, which transports them to the liver. In the liver, hepatocytes either pass the glucose on through the circulatory system or store excess glucose as glycogen. Cells in the body take up the circulating glucose in response to insulin and, through a series of reactions called glycolysis, transfer some of the energy in glucose to ADP to form ATP (Figure 2). The last step in glycolysis produces the product pyruvate.

Glycolysis begins with the phosphorylation of glucose by hexokinase to form glucose-6-phosphate. This step uses one ATP, which is the donor of the phosphate group. Under the action of phosphofructokinase, glucose-6-phosphate is converted into fructose-6-phosphate. At this point, a second ATP donates its phosphate group, forming fructose-1,6-bisphosphate. This six-carbon sugar is split to form two phosphorylated three-carbon molecules, glyceraldehyde-3-phosphate and dihydroxyacetone phosphate, which are both converted into glyceraldehyde-3-phosphate. The glyceraldehyde-3-phosphate is further phosphorylated with groups donated by dihydrogen phosphate present in the cell to form the three-carbon molecule 1,3-bisphosphoglycerate. The energy of this reaction comes from the oxidation of (removal of electrons from) glyceraldehyde-3-phosphate. In a series of reactions leading to pyruvate, the two phosphate groups are then transferred to two ADPs to form two ATPs. Thus, glycolysis uses two ATPs but generates four ATPs, yielding a net gain of two ATPs and two molecules of pyruvate. In the presence of oxygen, pyruvate continues on to the Krebs cycle (also called the citric acid cycle or tricarboxylic acid cycle (TCA), where additional energy is extracted and passed on.

[caption id="" align="aligncenter" width="520"]This flowchart shows the different steps in glycolysis in detail. The top panel shows the energy-consuming phase, the middle panel shows the coupling of phosphorylation with oxidation, and the bottom panel shows the energy-releasing phase. Figure 2. Glycolysis Overview. During the energy-consuming phase of glycolysis, two ATPs are consumed, transferring two phosphates to the glucose molecule. The glucose molecule then splits into two three-carbon compounds, each containing a phosphate. During the second phase, an additional phosphate is added to each of the three-carbon compounds. The energy for this endergonic reaction is provided by the removal (oxidation) of two electrons from each three-carbon compound. During the energy-releasing phase, the phosphates are removed from both three-carbon compounds and used to produce four ATP molecules.[/caption]
[caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this video to learn about glycolysis.[/caption]

Glycolysis can be divided into two phases: energy consuming (also called chemical priming) and energy yielding. The first phase is the energy-consuming phase, so it requires two ATP molecules to start the reaction for each molecule of glucose. However, the end of the reaction produces four ATPs, resulting in a net gain of two ATP energy molecules.

Glycolysis can be expressed as the following equation:

Glucose + 2ATP + 2NAD+ + 4ADP + 2Pi → 2 Pyruvate + 4ATP + 2NADH + 2H+

This equation states that glucose, in combination with ATP (the energy source), NAD+ (a coenzyme that serves as an electron acceptor), and inorganic phosphate, breaks down into two pyruvate molecules, generating four ATP molecules—for a net yield of two ATP—and two energy-containing NADH coenzymes. The NADH that is produced in this process will be used later to produce ATP in the mitochondria. Importantly, by the end of this process, one glucose molecule generates two pyruvate molecules, two high-energy ATP molecules, and two electron-carrying NADH molecules.

The following discussions of glycolysis include the enzymes responsible for the reactions. When glucose enters a cell, the enzyme hexokinase (or glucokinase, in the liver) rapidly adds a phosphate to convert it into glucose-6-phosphate. A kinase is a type of enzyme that adds a phosphate molecule to a substrate (in this case, glucose, but it can be true of other molecules also). This conversion step requires one ATP and essentially traps the glucose in the cell, preventing it from passing back through the plasma membrane, thus allowing glycolysis to proceed. It also functions to maintain a concentration gradient with higher glucose levels in the blood than in the tissues. By establishing this concentration gradient, the glucose in the blood will be able to flow from an area of high concentration (the blood) into an area of low concentration (the tissues) to be either used or stored. Hexokinase is found in nearly every tissue in the body. Glucokinase, on the other hand, is expressed in tissues that are active when blood glucose levels are high, such as the liver. Hexokinase has a higher affinity for glucose than glucokinase and therefore is able to convert glucose at a faster rate than glucokinase. This is important when levels of glucose are very low in the body, as it allows glucose to travel preferentially to those tissues that require it more.

In the next step of the first phase of glycolysis, the enzyme glucose-6-phosphate isomerase converts glucose-6-phosphate into fructose-6-phosphate. Like glucose, fructose is also a six carbon-containing sugar. The enzyme phosphofructokinase-1 then adds one more phosphate to convert fructose-6-phosphate into fructose-1-6-bisphosphate, another six-carbon sugar, using another ATP molecule. Aldolase then breaks down this fructose-1-6-bisphosphate into two three-carbon molecules, glyceraldehyde-3-phosphate and dihydroxyacetone phosphate. The triosephosphate isomerase enzyme then converts dihydroxyacetone phosphate into a second glyceraldehyde-3-phosphate molecule. Therefore, by the end of this chemical-priming or energy-consuming phase, one glucose molecule is broken down into two glyceraldehyde-3-phosphate molecules.

The second phase of glycolysis, the energy-yielding phase, creates the energy that is the product of glycolysis. Glyceraldehyde-3-phosphate dehydrogenase converts each three-carbon glyceraldehyde-3-phosphate produced during the energy-consuming phase into 1,3-bisphosphoglycerate. This reaction releases an electron that is then picked up by NAD+ to create an NADH molecule. NADH is a high-energy molecule, like ATP, but unlike ATP, it is not used as energy currency by the cell. Because there are two glyceraldehyde-3-phosphate molecules, two NADH molecules are synthesized during this step. Each 1,3-bisphosphoglycerate is subsequently dephosphorylated (i.e., a phosphate is removed) by phosphoglycerate kinase into 3-phosphoglycerate. Each phosphate released in this reaction can convert one molecule of ADP into one high-energy ATP molecule, resulting in a gain of two ATP molecules.

The enzyme phosphoglycerate mutase then converts the 3-phosphoglycerate molecules into 2-phosphoglycerate. The enolase enzyme then acts upon the 2-phosphoglycerate molecules to convert them into phosphoenolpyruvate molecules. The last step of glycolysis involves the dephosphorylation of the two phosphoenolpyruvate molecules by pyruvate kinase to create two pyruvate molecules and two ATP molecules.

In summary, one glucose molecule breaks down into two pyruvate molecules, and creates two net ATP molecules and two NADH molecules by glycolysis. Therefore, glycolysis generates energy for the cell and creates pyruvate molecules that can be processed further through the aerobic Krebs cycle (also called the citric acid cycle or tricarboxylic acid cycle); converted into lactic acid or alcohol (in yeast) by fermentation; or used later for the synthesis of glucose through gluconeogenesis.

Anaerobic Respiration

When oxygen is limited or absent, pyruvate enters an anaerobic pathway. In these reactions, pyruvate can be converted into lactic acid. In addition to generating an additional ATP, this pathway serves to keep the pyruvate concentration low so glycolysis continues, and it oxidizes NADH into the NAD+ needed by glycolysis. In this reaction, lactic acid replaces oxygen as the final electron acceptor. Anaerobic respiration occurs in most cells of the body when oxygen is limited or mitochondria are absent or nonfunctional. For example, because erythrocytes (red blood cells) lack mitochondria, they must produce their ATP from anaerobic respiration. This is an effective pathway of ATP production for short periods of time, ranging from seconds to a few minutes. The lactic acid produced diffuses into the plasma and is carried to the liver, where it is converted back into pyruvate or glucose via the Cori cycle. Similarly, when a person exercises, muscles use ATP faster than oxygen can be delivered to them. They depend on glycolysis and lactic acid production for rapid ATP production.

Aerobic Respiration

In the presence of oxygen, pyruvate can enter the Krebs cycle where additional energy is extracted as electrons are transferred from the pyruvate to the receptors NAD+, GDP, and FAD, with carbon dioxide being a “waste product” (Figure 3). The NADH and FADH2 pass electrons on to the electron transport chain, which uses the transferred energy to produce ATP. As the terminal step in the electron transport chain, oxygen is the terminal electron acceptor and creates water inside the mitochondria.

[caption id="" align="aligncenter" width="550"]This flowchart shows the processes of anaerobic and aerobic respiration. The top image shows the energy consuming phase of glycolysis. This branches into aerobic respiration on the left and anaerobic respiration on the right. Figure 3. Aerobic versus Anaerobic Respiration. The process of anaerobic respiration converts glucose into two lactate molecules in the absence of oxygen or within erythrocytes that lack mitochondria. During aerobic respiration, glucose is oxidized into two pyruvate molecules.[/caption]

Krebs Cycle/Citric Acid Cycle/Tricarboxylic Acid Cycle

The pyruvate molecules generated during glycolysis are transported across the mitochondrial membrane into the inner mitochondrial matrix, where they are metabolized by enzymes in a pathway called the Krebs cycle (Figure 4). The Krebs cycle is also commonly called the citric acid cycle or the tricarboxylic acid (TCA) cycle. During the Krebs cycle, high-energy molecules, including ATP, NADH, and FADH2, are created. NADH and FADH2 then pass electrons through the electron transport chain in the mitochondria to generate more ATP molecules.

[caption id="" align="aligncenter" width="520"]The top panel of this figure shows the transformation of pyruvate to acetyl-CoA, and the bottom panel shows the steps in Krebs cycle. Figure 4. Krebs Cycle. During the Krebs cycle, each pyruvate that is generated by glycolysis is converted into a two-carbon acetyl CoA molecule. The acetyl CoA is systematically processed through the cycle and produces high-energy NADH, FADH2, and ATP molecules.[/caption]

The three-carbon pyruvate molecule generated during glycolysis moves from the cytoplasm into the mitochondrial matrix, where it is converted by the enzyme pyruvate dehydrogenase into a two-carbon acetyl coenzyme A (acetyl CoA) molecule. This reaction is an oxidative decarboxylation reaction. It converts the three-carbon pyruvate into a two-carbon acetyl CoA molecule, releasing carbon dioxide and transferring two electrons that combine with NAD+ to form NADH. Acetyl CoA enters the Krebs cycle by combining with a four-carbon molecule, oxaloacetate, to form the six-carbon molecule citrate, or citric acid, at the same time releasing the coenzyme A molecule.

The six-carbon citrate molecule is systematically converted to a five-carbon molecule and then a four-carbon molecule, ending with oxaloacetate, the beginning of the cycle. Along the way, each citrate molecule will produce one ATP, one FADH2, and three NADH. The FADH2 and NADH will enter the oxidative phosphorylation system located in the inner mitochondrial membrane. In addition, the Krebs cycle supplies the starting materials to process and break down proteins and fats.

To start the Krebs cycle, citrate synthase combines acetyl CoA and oxaloacetate to form a six-carbon citrate molecule; CoA is subsequently released and can combine with another pyruvate molecule to begin the cycle again. The aconitase enzyme converts citrate into isocitrate. In two successive steps of oxidative decarboxylation, two molecules of CO2 and two NADH molecules are produced when isocitrate dehydrogenase converts isocitrate into the five-carbon α-ketoglutarate, which is then catalyzed and converted into the four-carbon succinyl CoA by α-ketoglutarate dehydrogenase. The enzyme succinyl CoA dehydrogenase then converts succinyl CoA into succinate and forms the high-energy molecule GTP, which transfers its energy to ADP to produce ATP. Succinate dehydrogenase then converts succinate into fumarate, forming a molecule of FADH2. Fumarase then converts fumarate into malate, which malate dehydrogenase then converts back into oxaloacetate while reducing NAD+ to NADH. Oxaloacetate is then ready to combine with the next acetyl CoA to start the Krebs cycle again (see Figure 4). For each turn of the cycle, three NADH, one ATP (through GTP), and one FADH2 are created. Each carbon of pyruvate is converted into CO2, which is released as a byproduct of oxidative (aerobic) respiration.

Oxidative Phosphorylation and the Electron Transport Chain

The electron transport chain (ETC) uses the NADH and FADH2 produced by the Krebs cycle to generate ATP. Electrons from NADH and FADH2 are transferred through protein complexes embedded in the inner mitochondrial membrane by a series of enzymatic reactions. The electron transport chain consists of a series of four enzyme complexes (Complex I – Complex IV) and two coenzymes (ubiquinone and Cytochrome c), which act as electron carriers and proton pumps used to transfer H+ ions into the space between the inner and outer mitochondrial membranes (Figure 5). The ETC couples the transfer of electrons between a donor (like NADH) and an electron acceptor (like O2) with the transfer of protons (H+ ions) across the inner mitochondrial membrane, enabling the process of oxidative phosphorylation. In the presence of oxygen, energy is passed, stepwise, through the electron carriers to collect gradually the energy needed to attach a phosphate to ADP and produce ATP. The role of molecular oxygen, O2, is as the terminal electron acceptor for the ETC. This means that once the electrons have passed through the entire ETC, they must be passed to another, separate molecule. These electrons, O2, and H+ ions from the matrix combine to form new water molecules. This is the basis for your need to breathe in oxygen. Without oxygen, electron flow through the ETC ceases.

[caption id="" align="aligncenter" width="550"]This image shows the mitochondrial membrane with proton pumps and ATP synthase embedded in the membrane. Arrows show the direction of flow of proteins and electrons across the membrane. Figure 5. Electron Transport Chain. The electron transport chain is a series of electron carriers and ion pumps that are used to pump H+ ions out of the inner mitochondrial matrix.[/caption]

The electrons released from NADH and FADH2 are passed along the chain by each of the carriers, which are reduced when they receive the electron and oxidized when passing it on to the next carrier. Each of these reactions releases a small amount of energy, which is used to pump H+ ions across the inner membrane. The accumulation of these protons in the space between the membranes creates a proton gradient with respect to the mitochondrial matrix.

Also embedded in the inner mitochondrial membrane is an amazing protein pore complex called ATP synthase. Effectively, it is a turbine that is powered by the flow of H+ ions across the inner membrane down a gradient and into the mitochondrial matrix. As the H+ ions traverse the complex, the shaft of the complex rotates. This rotation enables other portions of ATP synthase to encourage ADP and Pi to create ATP. In accounting for the total number of ATP produced per glucose molecule through aerobic respiration, it is important to remember the following points:

  • A net of two ATP are produced through glycolysis (four produced and two consumed during the energy-consuming stage). However, these two ATP are used for transporting the NADH produced during glycolysis from the cytoplasm into the mitochondria. Therefore, the net production of ATP during glycolysis is zero.
  • In all phases after glycolysis, the number of ATP, NADH, and FADH2 produced must be multiplied by two to reflect how each glucose molecule produces two pyruvate molecules.
  • In the ETC, about three ATP are produced for every oxidized NADH. However, only about two ATP are produced for every oxidized FADH2. The electrons from FADH2 produce less ATP, because they start at a lower point in the ETC (Complex II) compared to the electrons from NADH (Complex I) (see Figure 5).

Therefore, for every glucose molecule that enters aerobic respiration, a net total of 36 ATPs are produced (Figure 6).

[caption id="" align="aligncenter" width="450"]This figure shows the different steps in which carbohydrates are metabolized and lists the number of ATP molecules produced in each step. The different steps shown are glycolysis, transformation of pyruvate to acetyl-CoA, the Krebs cycle, and the electron transport chain. Figure 6. Carbohydrate Metabolism. Carbohydrate metabolism involves glycolysis, the Krebs cycle, and the electron transport chain.[/caption]

Gluconeogenesis

Gluconeogenesis is the synthesis of new glucose molecules from pyruvate, lactate, glycerol, or the amino acids alanine or glutamine. This process takes place primarily in the liver during periods of low glucose, that is, under conditions of fasting, starvation, and low carbohydrate diets. So, the question can be raised as to why the body would create something it has just spent a fair amount of effort to break down? Certain key organs, including the brain, can use only glucose as an energy source; therefore, it is essential that the body maintain a minimum blood glucose concentration. When the blood glucose concentration falls below that certain point, new glucose is synthesized by the liver to raise the blood concentration to normal.

Gluconeogenesis is not simply the reverse of glycolysis. There are some important differences (Figure 7). Pyruvate is a common starting material for gluconeogenesis. First, the pyruvate is converted into oxaloacetate. Oxaloacetate then serves as a substrate for the enzyme phosphoenolpyruvate carboxykinase (PEPCK), which transforms oxaloacetate into phosphoenolpyruvate (PEP). From this step, gluconeogenesis is nearly the reverse of glycolysis. PEP is converted back into 2-phosphoglycerate, which is converted into 3-phosphoglycerate. Then, 3-phosphoglycerate is converted into 1,3 bisphosphoglycerate and then into glyceraldehyde-3-phosphate. Two molecules of glyceraldehyde-3-phosphate then combine to form fructose-1-6-bisphosphate, which is converted into fructose 6-phosphate and then into glucose-6-phosphate. Finally, a series of reactions generates glucose itself. In gluconeogenesis (as compared to glycolysis), the enzyme hexokinase is replaced by glucose-6-phosphatase, and the enzyme phosphofructokinase-1 is replaced by fructose-1,6-bisphosphatase. This helps the cell to regulate glycolysis and gluconeogenesis independently of each other.

As will be discussed as part of lipolysis, fats can be broken down into glycerol, which can be phosphorylated to form dihydroxyacetone phosphate or DHAP. DHAP can either enter the glycolytic pathway or be used by the liver as a substrate for gluconeogenesis.

[caption id="" align="aligncenter" width="550"]This figure shows the different steps in gluconeogenesis, where pyruvate is converted to glucose. Figure 8. Gluconeogenesis. Gluconeogenesis is the synthesis of glucose from pyruvate, lactate, glycerol, alanine, or glutamate.[/caption]
Watch this CrashCourse video on ATP and cellular respiration.
]]> 2646 2634 2 0 24.3 Lipid Metabolism https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2653 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2653

Learning Objectives

By the end of this section, you will be able to:
  • Explain how energy can be derived from fat
  • Explain the purpose and process of ketogenesis
  • Describe the process of ketone body oxidation
  • Explain the purpose and the process of lipogenesis

Fats (or triglycerides) within the body are ingested as food or synthesized by adipocytes or hepatocytes from carbohydrate precursors (Figure 1). Lipid metabolism entails the oxidation of fatty acids to either generate energy or synthesize new lipids from smaller constituent molecules. Lipid metabolism is associated with carbohydrate metabolism, as products of glucose (such as acetyl CoA) can be converted into lipids.

[caption id="" align="aligncenter" width="450"]The top image shows the chemical formula for a triglyceride, and the bottom panel shows the formula for a monoglyceride. Figure 1. Triglyceride Broken Down into a Monoglyceride A triglyceride molecule (a) breaks down into a monoglyceride (b).[/caption]

Lipid metabolism begins in the intestine where ingested triglycerides are broken down into smaller chain fatty acids and subsequently into monoglyceride molecules (see Figure 1b) by pancreatic lipases, enzymes that break down fats after they are emulsified by bile salts. When food reaches the small intestine in the form of chyme, a digestive hormone called cholecystokinin (CCK) is released by intestinal cells in the intestinal mucosa. CCK stimulates the release of pancreatic lipase from the pancreas and stimulates the contraction of the gallbladder to release stored bile salts into the intestine. CCK also travels to the brain, where it can act as a hunger suppressant.

Together, the pancreatic lipases and bile salts break down triglycerides into free fatty acids. These fatty acids can be transported across the intestinal membrane. However, once they cross the membrane, they are recombined to again form triglyceride molecules. Within the intestinal cells, these triglycerides are packaged along with cholesterol molecules in phospholipid vesicles called chylomicrons (Figure 2). The chylomicrons enable fats and cholesterol to move within the aqueous environment of your lymphatic and circulatory systems. Chylomicrons leave the enterocytes by exocytosis and enter the lymphatic system via lacteals in the villi of the intestine. From the lymphatic system, the chylomicrons are transported to the circulatory system. Once in the circulation, they can either go to the liver or be stored in fat cells (adipocytes) that comprise adipose (fat) tissue found throughout the body.

[caption id="" align="aligncenter" width="380"]This figure shows a chylomicron containing triglycerides and cholesterol molecules as well as other lipids. Figure 2. Chylomicrons. Chylomicrons contain triglycerides, cholesterol molecules, and other apolipoproteins (protein molecules). They function to carry these water-insoluble molecules from the intestine, through the lymphatic system, and into the bloodstream, which carries the lipids to adipose tissue for storage.[/caption]

Lipolysis

To obtain energy from fat, triglycerides must first be broken down by hydrolysis into their two principal components, fatty acids and glycerol. This process, called lipolysis, takes place in the cytoplasm. The resulting fatty acids are oxidized by β-oxidation into acetyl CoA, which is used by the Krebs cycle. The glycerol that is released from triglycerides after lipolysis directly enters the glycolysis pathway as DHAP. Because one triglyceride molecule yields three fatty acid molecules with as much as 16 or more carbons in each one, fat molecules yield more energy than carbohydrates and are an important source of energy for the human body. Triglycerides yield more than twice the energy per unit mass when compared to carbohydrates and proteins. Therefore, when glucose levels are low, triglycerides can be converted into acetyl CoA molecules and used to generate ATP through aerobic respiration.

The breakdown of fatty acids, called fatty acid oxidation or beta (β)-oxidation, begins in the cytoplasm, where fatty acids are converted into fatty acyl CoA molecules. This fatty acyl CoA combines with carnitine to create a fatty acyl carnitine molecule, which helps to transport the fatty acid across the mitochondrial membrane. Once inside the mitochondrial matrix, the fatty acyl carnitine molecule is converted back into fatty acyl CoA and then into acetyl CoA (Figure 3). The newly formed acetyl CoA enters the Krebs cycle and is used to produce ATP in the same way as acetyl CoA derived from pyruvate.

[caption id="" align="aligncenter" width="500"]This figure shows the reactions that break down fatty acids. The top panel shows the conversion of fatty acids into carnitine. The bottom panel shows the conversion of carnitine into acetyl-CoA. Figure 3. Breakdown of Fatty Acids. During fatty acid oxidation, triglycerides can be broken down into acetyl CoA molecules and used for energy when glucose levels are low.[/caption]

Ketogenesis

If excessive acetyl CoA is created from the oxidation of fatty acids and the Krebs cycle is overloaded and cannot handle it, the acetyl CoA is diverted to create ketone bodies. These ketone bodies can serve as a fuel source if glucose levels are too low in the body. Ketones serve as fuel in times of prolonged starvation or when patients suffer from uncontrolled diabetes and cannot utilize most of the circulating glucose. In both cases, fat stores are liberated to generate energy through the Krebs cycle and will generate ketone bodies when too much acetyl CoA accumulates.

In this ketone synthesis reaction, excess acetyl CoA is converted into hydroxymethylglutaryl CoA (HMG CoA). HMG CoA is a precursor of cholesterol and is an intermediate that is subsequently converted into β-hydroxybutyrate, the primary ketone body in the blood (Figure 4).

[caption id="" align="aligncenter" width="580"]This pathway shows the production of beta-hydroxybutyrate from acetyl-CoA. Figure 4. Ketogenesis. Excess acetyl CoA is diverted from the Krebs cycle to the ketogenesis pathway. This reaction occurs in the mitochondria of liver cells. The result is the production of β-hydroxybutyrate, the primary ketone body found in the blood.[/caption]

Ketone Body Oxidation

Organs that have classically been thought to be dependent solely on glucose, such as the brain, can actually use ketones as an alternative energy source. This keeps the brain functioning when glucose is limited. When ketones are produced faster than they can be used, they can be broken down into CO2 and acetone. The acetone is removed by exhalation. One symptom of ketogenesis is that the patient’s breath smells sweet like alcohol. This effect provides one way of telling if a diabetic is properly controlling the disease. The carbon dioxide produced can acidify the blood, leading to diabetic ketoacidosis, a dangerous condition in diabetics.

Ketones oxidize to produce energy for the brain. beta (β)-hydroxybutyrate is oxidized to acetoacetate and NADH is released. An HS-CoA molecule is added to acetoacetate, forming acetoacetyl CoA. The carbon within the acetoacetyl CoA that is not bonded to the CoA then detaches, splitting the molecule in two. This carbon then attaches to another free HS-CoA, resulting in two acetyl CoA molecules. These two acetyl CoA molecules are then processed through the Krebs cycle to generate energy (Figure 5).

[caption id="" align="aligncenter" width="520"]This figure shows the reactions in which ketone is oxidized to acetyl-CoA. Figure 5. Ketone Oxidation. When glucose is limited, ketone bodies can be oxidized to produce acetyl CoA to be used in the Krebs cycle to generate energy.[/caption]

Lipogenesis

When glucose levels are plentiful, the excess acetyl CoA generated by glycolysis can be converted into fatty acids, triglycerides, cholesterol, steroids, and bile salts. This process, called lipogenesis, creates lipids (fat) from the acetyl CoA and takes place in the cytoplasm of adipocytes (fat cells) and hepatocytes (liver cells). When you eat more glucose or carbohydrates than your body needs, your system uses acetyl CoA to turn the excess into fat. Although there are several metabolic sources of acetyl CoA, it is most commonly derived from glycolysis. Acetyl CoA availability is significant, because it initiates lipogenesis. Lipogenesis begins with acetyl CoA and advances by the subsequent addition of two carbon atoms from another acetyl CoA; this process is repeated until fatty acids are the appropriate length. Because this is a bond-creating anabolic process, ATP is consumed. However, the creation of triglycerides and lipids is an efficient way of storing the energy available in carbohydrates. Triglycerides and lipids, high-energy molecules, are stored in adipose tissue until they are needed.

Although lipogenesis occurs in the cytoplasm, the necessary acetyl CoA is created in the mitochondria and cannot be transported across the mitochondrial membrane. To solve this problem, pyruvate is converted into both oxaloacetate and acetyl CoA. Two different enzymes are required for these conversions. Oxaloacetate forms via the action of pyruvate carboxylase, whereas the action of pyruvate dehydrogenase creates acetyl CoA. Oxaloacetate and acetyl CoA combine to form citrate, which can cross the mitochondrial membrane and enter the cytoplasm. In the cytoplasm, citrate is converted back into oxaloacetate and acetyl CoA. Oxaloacetate is converted into malate and then into pyruvate. Pyruvate crosses back across the mitochondrial membrane to wait for the next cycle of lipogenesis. The acetyl CoA is converted into malonyl CoA that is used to synthesize fatty acids. Figure 6 summarizes the pathways of lipid metabolism.

[caption id="" align="aligncenter" width="550"]This figure shows the different reactions that take place for lipid metabolism. Figure 6. Lipid Metabolism. Lipids may follow one of several pathways during metabolism. Glycerol and fatty acids follow different pathways.[/caption]
]]> 2653 2634 3 0 24.4 Protein Metabolism https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2658 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2658

Learning Objectives

By the end of this section, you will be able to:
  • Describe how the body digests proteins
  • Explain how the urea cycle prevents toxic concentrations of nitrogen
  • Differentiate between glucogenic and ketogenic amino acids
  • Explain how protein can be used for energy

Much of the body is made of protein, and these proteins take on a myriad of forms. They represent cell signaling receptors, signaling molecules, structural members, enzymes, intracellular trafficking components, extracellular matrix scaffolds, ion pumps, ion channels, oxygen and CO2 transporters (hemoglobin). That is not even the complete list! There is protein in bones (collagen), muscles, and tendons; the hemoglobin that transports oxygen; and enzymes that catalyze all biochemical reactions. Protein is also used for growth and repair. Amid all these necessary functions, proteins also hold the potential to serve as a metabolic fuel source. Proteins are not stored for later use, so excess proteins must be converted into glucose or triglycerides, and used to supply energy or build energy reserves. Although the body can synthesize proteins from amino acids, food is an important source of those amino acids, especially because humans cannot synthesize all of the 20 amino acids used to build proteins.

The digestion of proteins begins in the stomach. When protein-rich foods enter the stomach, they are greeted by a mixture of the enzyme pepsin and hydrochloric acid (HCl; 0.5 percent). The latter produces an environmental pH of 1.5–3.5 that denatures proteins within food. Pepsin cuts proteins into smaller polypeptides and their constituent amino acids. When the food-gastric juice mixture (chyme) enters the small intestine, the pancreas releases sodium bicarbonate to neutralize the HCl. This helps to protect the lining of the intestine. The small intestine also releases digestive hormones, including secretin and CCK, which stimulate digestive processes to break down the proteins further. Secretin also stimulates the pancreas to release sodium bicarbonate. The pancreas releases most of the digestive enzymes, including the proteases trypsin, chymotrypsin, and elastase, which aid protein digestion. Together, all of these enzymes break complex proteins into smaller individual amino acids (Figure 1), which are then transported across the intestinal mucosa to be used to create new proteins, or to be converted into fats or acetyl CoA and used in the Krebs cycle.

[caption id="" align="aligncenter" width="500"]The left panel shows the main organs of the digestive system, and the right panel shows a magnified view of the intestine. Text callouts indicate the different protein digesting enzymes produced in different organs. Figure 1. Digestive Enzymes and Hormones. Enzymes in the stomach and small intestine break down proteins into amino acids. HCl in the stomach aids in proteolysis, and hormones secreted by intestinal cells direct the digestive processes.[/caption]

In order to avoid breaking down the proteins that make up the pancreas and small intestine, pancreatic enzymes are released as inactive proenzymes that are only activated in the small intestine. In the pancreas, vesicles store trypsin and chymotrypsin as trypsinogen and chymotrypsinogen. Once released into the small intestine, an enzyme found in the wall of the small intestine, called enterokinase, binds to trypsinogen and converts it into its active form, trypsin. Trypsin then binds to chymotrypsinogen to convert it into the active chymotrypsin. Trypsin and chymotrypsin break down large proteins into smaller peptides, a process called proteolysis. These smaller peptides are catabolized into their constituent amino acids, which are transported across the apical surface of the intestinal mucosa in a process that is mediated by sodium-amino acid transporters. These transporters bind sodium and then bind the amino acid to transport it across the membrane. At the basal surface of the mucosal cells, the sodium and amino acid are released. The sodium can be reused in the transporter, whereas the amino acids are transferred into the bloodstream to be transported to the liver and cells throughout the body for protein synthesis.

Freely available amino acids are used to create proteins. If amino acids exist in excess, the body has no capacity or mechanism for their storage; thus, they are converted into glucose or ketones, or they are decomposed. Amino acid decomposition results in hydrocarbons and nitrogenous waste. However, high concentrations of nitrogen are toxic. The urea cycle processes nitrogen and facilitates its excretion from the body.

Urea Cycle

The urea cycle is a set of biochemical reactions that produces urea from ammonium ions in order to prevent a toxic level of ammonium in the body. It occurs primarily in the liver and, to a lesser extent, in the kidney. Prior to the urea cycle, ammonium ions are produced from the breakdown of amino acids. In these reactions, an amine group, or ammonium ion, from the amino acid is exchanged with a keto group on another molecule. This transamination event creates a molecule that is necessary for the Krebs cycle and an ammonium ion that enters into the urea cycle to be eliminated.

In the urea cycle, ammonium is combined with CO2, resulting in urea and water. The urea is eliminated through the kidneys in the urine (Figure 2).

[caption id="" align="aligncenter" width="520"]This image shows the reactions of the urea cycle and the organelles in which they take place. Figure 2. Urea Cycle. Nitrogen is transaminated, creating ammonia and intermediates of the Krebs cycle. Ammonia is processed in the urea cycle to produce urea that is eliminated through the kidneys.[/caption]

Amino acids can also be used as a source of energy, especially in times of starvation. Because the processing of amino acids results in the creation of metabolic intermediates, including pyruvate, acetyl CoA, acetoacyl CoA, oxaloacetate, and α-ketoglutarate, amino acids can serve as a source of energy production through the Krebs cycle (Figure 3). Figure 4 summarizes the pathways of catabolism and anabolism for carbohydrates, lipids, and proteins.

[caption id="" align="aligncenter" width="550"]This figure shows the different reactions in which products of carbohydrate breakdown are converted into different amino acids. Figure 3. Energy from Amino Acids. Amino acids can be broken down into precursors for glycolysis or the Krebs cycle. Amino acids (in bold) can enter the cycle through more than one pathway.[/caption]
[caption id="attachment_1819" align="aligncenter" width="500"]This diagram shows the different metabolic pathways, and how they are connected. Figure 4. Catabolic and Anabolic Pathways. Nutrients follow a complex pathway from ingestion through anabolism and catabolism to energy production.[/caption]
]]> 2658 2634 4 0 24.5 Metabolic States of the Body https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2661 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2661

Learning Objectives

By the end of this section, you will be able to:
  • Describe what defines each of the three metabolic states
  • Describe the processes that occur during the absorptive state of metabolism
  • Describe the processes that occur during the postabsorptive state of metabolism
  • Explain how the body processes glucose when the body is starved of fuel

You eat periodically throughout the day; however, your organs, especially the brain, need a continuous supply of glucose. How does the body meet this constant demand for energy? Your body processes the food you eat both to use immediately and, importantly, to store as energy for later demands. If there were no method in place to store excess energy, you would need to eat constantly in order to meet energy demands. Distinct mechanisms are in place to facilitate energy storage, and to make stored energy available during times of fasting and starvation.

The Absorptive State

The absorptive state, or the fed state, occurs after a meal when your body is digesting the food and absorbing the nutrients (anabolism exceeds catabolism). Digestion begins the moment you put food into your mouth, as the food is broken down into its constituent parts to be absorbed through the intestine. The digestion of carbohydrates begins in the mouth, whereas the digestion of proteins and fats begins in the stomach and small intestine. The constituent parts of these carbohydrates, fats, and proteins are transported across the intestinal wall and enter the bloodstream (sugars and amino acids) or the lymphatic system (fats). From the intestines, these systems transport them to the liver, adipose tissue, or muscle cells that will process and use, or store, the energy.

Depending on the amounts and types of nutrients ingested, the absorptive state can linger for up to 4 hours. The ingestion of food and the rise of glucose concentrations in the bloodstream stimulate pancreatic beta cells to release insulin into the bloodstream, where it initiates the absorption of blood glucose by liver hepatocytes, and by adipose and muscle cells. Once inside these cells, glucose is immediately converted into glucose-6-phosphate. By doing this, a concentration gradient is established where glucose levels are higher in the blood than in the cells. This allows for glucose to continue moving from the blood to the cells where it is needed. Insulin also stimulates the storage of glucose as glycogen in the liver and muscle cells where it can be used for later energy needs of the body. Insulin also promotes the synthesis of protein in muscle. As you will see, muscle protein can be catabolized and used as fuel in times of starvation.

If energy is exerted shortly after eating, the dietary fats and sugars that were just ingested will be processed and used immediately for energy. If not, the excess glucose is stored as glycogen in the liver and muscle cells, or as fat in adipose tissue; excess dietary fat is also stored as triglycerides in adipose tissues.

Figure 1 summarizes the metabolic processes occurring in the body during the absorptive state.

[caption id="" align="aligncenter" width="560"]This figure shows how nutrients are absorbed by the body. The diagram shows digested nutrients entering the blood stream and being absorbed by liver cells, muscle cells, and adipose cells. Underneath each panel, text details the process taking place in each cell type. Figure 1. Absorptive State. During the absorptive state, the body digests food and absorbs the nutrients.[/caption]

The Postabsorptive State

The postabsorptive state, or the fasting state, occurs when the food has been digested, absorbed, and stored. You commonly fast overnight, but skipping meals during the day puts your body in the postabsorptive state as well. During this state, the body must rely initially on stored glycogen. Glucose levels in the blood begin to drop as it is absorbed and used by the cells. In response to the decrease in glucose, insulin levels also drop. Glycogen and triglyceride storage slows. However, due to the demands of the tissues and organs, blood glucose levels must be maintained in the normal range of 80–120 mg/dL. In response to a drop in blood glucose concentration, the hormone glucagon is released from the alpha cells of the pancreas. Glucagon acts upon the liver cells, where it inhibits the synthesis of glycogen and stimulates the breakdown of stored glycogen back into glucose. This glucose is released from the liver to be used by the peripheral tissues and the brain. As a result, blood glucose levels begin to rise. Gluconeogenesis will also begin in the liver to replace the glucose that has been used by the peripheral tissues.

After ingestion of food, fats and proteins are processed as described previously; however, the glucose processing changes a bit. The peripheral tissues preferentially absorb glucose. The liver, which normally absorbs and processes glucose, will not do so after a prolonged fast. The gluconeogenesis that has been ongoing in the liver will continue after fasting to replace the glycogen stores that were depleted in the liver. After these stores have been replenished, excess glucose that is absorbed by the liver will be converted into triglycerides and fatty acids for long-term storage. Figure 2 summarizes the metabolic processes occurring in the body during the postabsorptive state.

[caption id="" align="aligncenter" width="550"]This figure shows the postabsorptive stage where no nutrients enter the blood stream from the digestive system and its effects of liver cells, muscle cells, and adipose cells. Figure 2. Postabsorptive State. During the postabsorptive state, the body must rely on stored glycogen for energy.[/caption]

Starvation

When the body is deprived of nourishment for an extended period of time, it goes into “survival mode.” The first priority for survival is to provide enough glucose or fuel for the brain. The second priority is the conservation of amino acids for proteins. Therefore, the body uses ketones to satisfy the energy needs of the brain and other glucose-dependent organs, and to maintain proteins in the cells (see Chapter 24.1 Figure 1). Because glucose levels are very low during starvation, glycolysis will shut off in cells that can use alternative fuels. For example, muscles will switch from using glucose to fatty acids as fuel. As previously explained, fatty acids can be converted into acetyl CoA and processed through the Krebs cycle to make ATP. Pyruvate, lactate, and alanine from muscle cells are not converted into acetyl CoA and used in the Krebs cycle, but are exported to the liver to be used in the synthesis of glucose. As starvation continues, and more glucose is needed, glycerol from fatty acids can be liberated and used as a source for gluconeogenesis. After several days of starvation, ketone bodies become the major source of fuel for the heart and other organs. As starvation continues, fatty acids and triglyceride stores are used to create ketones for the body. This prevents the continued breakdown of proteins that serve as carbon sources for gluconeogenesis. Once these stores are fully depleted, proteins from muscles are released and broken down for glucose synthesis. Overall survival is dependent on the amount of fat and protein stored in the body.
]]> 2661 2634 5 0 24.7 Nutrition and Diet https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2663 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2663

Learning Objectives

By the end of this section, you will be able to:
  • Explain how different foods can affect metabolism
  • Describe a healthy diet, as recommended by the U.S. Department of Agriculture (USDA)
  • List reasons why vitamins and minerals are critical to a healthy diet

The carbohydrates, lipids, and proteins in the foods you eat are used for energy to power molecular, cellular, and organ system activities. Importantly, the energy is stored primarily as fats. The quantity and quality of food that is ingested, digested, and absorbed affects the amount of fat that is stored as excess calories. Diet—both what you eat and how much you eat—has a dramatic impact on your health. Eating too much or too little food can lead to serious medical issues, including cardiovascular disease, cancer, anorexia, and diabetes, among others. Combine an unhealthy diet with unhealthy environmental conditions, such as smoking, and the potential medical complications increase significantly.

Food and Metabolism

The amount of energy that is needed or ingested per day is measured in calories. The nutritional Calorie (C) is the amount of heat it takes to raise 1 kg (1000 g) of water by 1 °C. This is different from the calorie (c) used in the physical sciences, which is the amount of heat it takes to raise 1 g of water by 1 °C. When we refer to "calorie," we are referring to the nutritional Calorie.

On average, a person needs 1500 to 2000 calories per day to sustain (or carry out) daily activities. The total number of calories needed by one person is dependent on their body mass, age, height, gender, activity level, and the amount of exercise per day. If exercise is regular part of one’s day, more calories are required. As a rule, people underestimate the number of calories ingested and overestimate the amount they burn through exercise. This can lead to ingestion of too many calories per day. The accumulation of an extra 3500 calories adds one pound of weight. If an excess of 200 calories per day is ingested, one extra pound of body weight will be gained every 18 days. At that rate, an extra 20 pounds can be gained over the course of a year. Of course, this increase in calories could be offset by increased exercise. Running or jogging one mile burns almost 100 calories.

The type of food ingested also affects the body’s metabolic rate. Processing of carbohydrates requires less energy than processing of proteins. In fact, the breakdown of carbohydrates requires the least amount of energy, whereas the processing of proteins demands the most energy. In general, the amount of calories ingested and the amount of calories burned determines the overall weight. To lose weight, the number of calories burned per day must exceed the number ingested. Calories are in almost everything you ingest, so when considering calorie intake, beverages must also be considered.

To help provide guidelines regarding the types and quantities of food that should be eaten every day, the USDA has updated their food guidelines from MyPyramid to MyPlate. They have put the recommended elements of a healthy meal into the context of a place setting of food. MyPlate categorizes food into the standard six food groups: fruits, vegetables, grains, protein foods, dairy, and oils. The accompanying website gives clear recommendations regarding quantity and type of each food that you should consume each day, as well as identifying which foods belong in each category. The accompanying graphic (Figure 1) gives a clear visual with general recommendations for a healthy and balanced meal. The guidelines recommend to “Make half your plate fruits and vegetables.” The other half is grains and protein, with a slightly higher quantity of grains than protein. Dairy products are represented by a drink, but the quantity can be applied to other dairy products as well.

[caption id="" align="aligncenter" width="350"]The figure shows a plate with different food groups assigned different portion sizes. Figure 1. MyPlate. The U.S. Department of Agriculture developed food guidelines called MyPlate to help demonstrate how to maintain a healthy lifestyle.[/caption]

ChooseMyPlate.gov provides extensive online resources for planning a healthy diet and lifestyle, including offering weight management tips and recommendations for physical activity. It also includes the SuperTracker, a web-based application to help you analyze your own diet and physical activity.

Everyday Connections

Metabolism and Obesity Obesity in the United States is epidemic. The rate of obesity has been steadily rising since the 1980s. In the 1990s, most states reported that less than 10 percent of their populations was obese, and the state with the highest rate reported that only 15 percent of their population was considered obese. By 2010, the U.S. Centers for Disease Control and Prevention reported that nearly 36 percent of adults over 20 years old were obese and an additional 33 percent were overweight, leaving only about 30 percent of the population at a healthy weight. These studies find the highest levels of obesity are concentrated in the southern states. They also find the level of childhood obesity is rising.

Obesity is defined by the body mass index (BMI), which is a measure of an individual’s weight-to-height ratio. The normal, or healthy, BMI range is between 18 and 24.9 kg/m2. Overweight is defined as a BMI of 25 to 29.9 kg/m2, and obesity is considered to be a BMI greater than 30 kg/m2. Obesity can arise from a number of factors, including overeating, poor diet, sedentary lifestyle, limited sleep, genetic factors, and even diseases or drugs. Severe obesity (morbid obesity) or long-term obesity can result in serious medical conditions, including coronary heart disease; type 2 diabetes; endometrial, breast, or colon cancer; hypertension (high blood pressure); dyslipidemia (high cholesterol or elevated triglycerides); stroke; liver disease; gall bladder disease; sleep apnea or respiratory diseases; osteoarthritis; and infertility. Research has shown that losing weight can help reduce or reverse the complications associated with these conditions.

Vitamins

Vitamins are organic compounds found in foods and are a necessary part of the biochemical reactions in the body. They are involved in a number of processes, including mineral and bone metabolism, and cell and tissue growth, and they act as cofactors for energy metabolism. The B vitamins play the largest role of any vitamins in metabolism (Table 3 and Table 4).

You get most of your vitamins through your diet, although some can be formed from the precursors absorbed during digestion. For example, the body synthesizes vitamin A from the β-carotene in orange vegetables like carrots and sweet potatoes. Vitamins are either fat-soluble or water-soluble. Fat-soluble vitamins A, D, E, and K, are absorbed through the intestinal tract with lipids in chylomicrons. Vitamin D is also synthesized in the skin through exposure to sunlight. Because they are carried in lipids, fat-soluble vitamins can accumulate in the lipids stored in the body. If excess vitamins are retained in the lipid stores in the body, hypervitaminosis can result.

Water-soluble vitamins, including the eight B vitamins and vitamin C, are absorbed with water in the gastrointestinal tract. These vitamins move easily through bodily fluids, which are water based, so they are not stored in the body. Excess water-soluble vitamins are excreted in the urine. Therefore, hypervitaminosis of water-soluble vitamins rarely occurs, except with an excess of vitamin supplements.

Fat-soluble Vitamins (Table 3)
Vitamin and alternative name Sources Recommended daily allowance Function Problems associated with deficiency
A
retinal or β-carotene
 Yellow and orange fruits and vegetables, dark green leafy vegetables, eggs, milk, liver 700–900 µg Eye and bone development, immune function Night blindness, epithelial changes, immune system deficiency
D
cholecalciferol
 Dairy products, egg yolks; also synthesized in the skin from exposure to sunlight 5–15 µg Aids in calcium absorption, promoting bone growth Rickets, bone pain, muscle weakness, increased risk of death from cardiovascular disease, cognitive impairment, asthma in children, cancer
E
tocopherols
 Seeds, nuts, vegetable oils, avocados, wheat germ 15 mg Antioxidant Anemia
K
phylloquinone
 Dark green leafy vegetables, broccoli, Brussels sprouts, cabbage 90–120 µg Blood clotting, bone health Hemorrhagic disease of newborn in infants; uncommon in adults
Water-soluble Vitamins (Table 4)
Vitamin and alternative name Sources Recommended daily allowance Function Problems associated with deficiency
B1
thiamine
 Whole grains, enriched bread and cereals, milk, meat 1.1–1.2 mg Carbohydrate metabolism Beriberi, Wernicke-Korsikoff syndrome
B2
riboflavin
 Brewer’s yeast, almonds, milk, organ meats, legumes, enriched breads and cereals, broccoli, asparagus 1.1–1.3 mg Synthesis of FAD for metabolism, production of red blood cells Fatigue, slowed growth, digestive problems, light sensitivity, epithelial problems like cracks in the corners of the mouth
B3
niacin
 Meat, fish, poultry, enriched breads and cereals, peanuts 14–16 mg Synthesis of NAD, nerve function, cholesterol production Cracked, scaly skin; dementia; diarrhea; also known as pellagra
B5
pantothenic acid
 Meat, poultry, potatoes, oats, enriched breads and cereals, tomatoes 5 mg Synthesis of coenzyme A in fatty acid metabolism Rare: symptoms may include fatigue, insomnia, depression, irritability
B6
pyridoxine
 Potatoes, bananas, beans, seeds, nuts, meat, poultry, fish, eggs, dark green leafy vegetables, soy, organ meats 1.3–1.5 mg Sodium and potassium balance, red blood cell synthesis, protein metabolism Confusion, irritability, depression, mouth and tongue sores
B7
biotin
 Liver, fruits, meats 30 µg Cell growth, metabolism of fatty acids, production of blood cells Rare in developed countries; symptoms include dermatitis, hair loss, loss of muscular coordination
B9
folic acid
 Liver, legumes, dark green leafy vegetables, enriched breads and cereals, citrus fruits 400 µg DNA/protein synthesis Poor growth, gingivitis, appetite loss, shortness of breath, gastrointestinal problems, mental deficits
B12
cyanocobalamin
 Fish, meat, poultry, dairy products, eggs 2.4 µg Fatty acid oxidation, nerve cell function, red blood cell production Pernicious anemia, leading to nerve cell damage
C
ascorbic acid
 Citrus fruits, red berries, peppers, tomatoes, broccoli, dark green leafy vegetables 75–90 mg Necessary to produce collagen for formation of connective tissue and teeth, and for wound healing Dry hair, gingivitis, bleeding gums, dry and scaly skin, slow wound healing, easy bruising, compromised immunity; can lead to scurvy

Minerals

Minerals in food are inorganic compounds that work with other nutrients to ensure the body functions properly. Minerals cannot be made in the body; they come from the diet. The amount of minerals in the body is small—only 4 percent of the total body mass—and most of that consists of the minerals that the body requires in moderate quantities: potassium, sodium, calcium, phosphorus, magnesium, and chloride.

The most common minerals in the body are calcium and phosphorous, both of which are stored in the skeleton and necessary for the hardening of bones. Most minerals are ionized, and their ionic forms are used in physiological processes throughout the body. Sodium and chloride ions are electrolytes in the blood and extracellular tissues, and iron ions are critical to the formation of hemoglobin. There are additional trace minerals that are still important to the body’s functions, but their required quantities are much lower.

Like vitamins, minerals can be consumed in toxic quantities (although it is rare). A healthy diet includes most of the minerals your body requires, so supplements and processed foods can add potentially toxic levels of minerals. Table 5 and Table 6 provide a summary of minerals and their function in the body.

Major Minerals (Table 5)
Mineral Sources Recommended daily allowance Function Problems associated with deficiency
Potassium Meats, some fish, fruits, vegetables, legumes, dairy products 4700 mg Nerve and muscle function; acts as an electrolyte Hypokalemia: weakness, fatigue, muscle cramping, gastrointestinal problems, cardiac problems
Sodium Table salt, milk, beets, celery, processed foods 2300 mg Blood pressure, blood volume, muscle and nerve function Rare
Calcium Dairy products, dark green leafy vegetables, blackstrap molasses, nuts, brewer’s yeast, some fish 1000 mg Bone structure and health; nerve and muscle functions, especially cardiac function Slow growth, weak and brittle bones
Phosphorous Meat, milk 700 mg Bone formation, metabolism, ATP production Rare
Magnesium Whole grains, nuts, leafy green vegetables 310–420 mg Enzyme activation, production of energy, regulation of other nutrients Agitation, anxiety, sleep problems, nausea and vomiting, abnormal heart rhythms, low blood pressure, muscular problems
Chloride Most foods, salt, vegetables, especially seaweed, tomatoes, lettuce, celery, olives 2300 mg Balance of body fluids, digestion Loss of appetite, muscle cramps
Trace Minerals (Table 6)
Mineral Sources Recommended daily allowance Function Problems associated with deficiency
Iron Meat, poultry, fish, shellfish, legumes, nuts, seeds, whole grains, dark leafy green vegetables 8–18 mg Transport of oxygen in blood, production of ATP Anemia, weakness, fatigue
Zinc Meat, fish, poultry, cheese, shellfish 8–11 mg Immunity, reproduction, growth, blood clotting, insulin and thyroid function Loss of appetite, poor growth, weight loss, skin problems, hair loss, vision problems, lack of taste or smell
Copper Seafood, organ meats, nuts, legumes, chocolate, enriched breads and cereals, some fruits and vegetables 900 µg Red blood cell production, nerve and immune system function, collagen formation, acts as an antioxidant Anemia, low body temperature, bone fractures, low white blood cell concentration, irregular heartbeat, thyroid problems
Iodine Fish, shellfish, garlic, lima beans, sesame seeds, soybeans, dark leafy green vegetables 150 µg Thyroid function Hypothyroidism: fatigue, weight gain, dry skin, temperature sensitivity
Sulfur Eggs, meat, poultry, fish, legumes None Component of amino acids Protein deficiency
Fluoride Fluoridated water 3–4 mg Maintenance of bone and tooth structure Increased cavities, weak bones and teeth
Manganese Nuts, seeds, whole grains, legumes 1.8–2.3 mg Formation of connective tissue and bones, blood clotting, sex hormone development, metabolism, brain and nerve function Infertility, bone malformation, weakness, seizures
Cobalt Fish, nuts, leafy green vegetables, whole grains None Component of B12 None
Selenium Brewer’s yeast, wheat germ, liver, butter, fish, shellfish, whole grains 55 µg Antioxidant, thyroid function, immune system function Muscle pain
Chromium Whole grains, lean meats, cheese, black pepper, thyme, brewer’s yeast 25–35 µg Insulin function High blood sugar, triglyceride, and cholesterol levels
Molybdenum Legumes, whole grains, nuts 45 µg Cofactor for enzymes Rare

]]> 2663 2634 6 0 Ch. 23 Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2666 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2666 [caption id="" align="aligncenter" width="600"]This photograph shows two women eating apples. Eating Apples Eating may be one of the simple pleasures in life, but digesting even one apple requires the coordinated work of many organs. (credit: “Aimanness Photography”/Flickr)[/caption]

Learning Objectives

After studying this chapter, you will be able to:
  • List and describe the functional anatomy of the organs and accessory organs of the digestive system
  • Discuss the processes and control of ingestion, propulsion, mechanical digestion, chemical digestion, absorption, and defecation
  • Discuss the roles of the liver, pancreas, and gallbladder in digestion
  • Compare and contrast the digestion of the three macronutrients
The digestive system is continually at work, yet people seldom appreciate the complex tasks it performs in a choreographed biologic symphony. Consider what happens when you eat an apple. Of course, you enjoy the apple’s taste as you chew it, but in the hours that follow, unless something goes amiss and you get a stomachache, you don’t notice that your digestive system is working. You may be taking a walk or studying or sleeping, having forgotten all about the apple, but your stomach and intestines are busy digesting it and absorbing its vitamins and other nutrients. By the time any waste material is excreted, the body has appropriated all it can use from the apple. In short, whether you pay attention or not, the organs of the digestive system perform their specific functions, allowing you to use the food you eat to keep you going. This chapter examines the structure and functions of these organs, and explores the mechanics and chemistry of the digestive processes.]]> 2666 2664 1 0 23.3 The Mouth, Pharynx, and Esophagus https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2684 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2684

Learning Objectives

By the end of this section, you will be able to:
  • Describe the structures of the mouth, including its three accessory digestive organs
  • Group the 32 adult teeth according to name, location, and function
  • Describe the process of swallowing, including the roles of the tongue, upper esophageal sphincter, and epiglottis
  • Trace the pathway food follows from ingestion into the mouth through release into the stomach

In this section, you will examine the anatomy and functions of the three main organs of the upper alimentary canal—the mouth, pharynx, and esophagus—as well as three associated accessory organs—the tongue, salivary glands, and teeth.

The Mouth

The cheeks, tongue, and palate frame the mouth, which is also called the oral cavity (or buccal cavity). The structures of the mouth are illustrated in Figure 1.

At the entrance to the mouth are the lips, or labia (singular = labium). Their outer covering is skin, which transitions to a mucous membrane in the mouth proper. Lips are very vascular with a thin layer of keratin; hence, the reason they are "red." They have a huge representation on the cerebral cortex, which probably explains the human fascination with kissing! The lips cover the orbicularis oris muscle, which regulates what comes in and goes out of the mouth. The labial frenulum is a midline fold of mucous membrane that attaches the inner surface of each lip to the gum. The cheeks make up the oral cavity’s sidewalls. While their outer covering is skin, their inner covering is mucous membrane. This membrane is made up of non-keratinized, stratified squamous epithelium. Between the skin and mucous membranes are connective tissue and buccinator muscles. The next time you eat some food, notice how the buccinator muscles in your cheeks and the orbicularis oris muscle in your lips contract, helping you keep the food from falling out of your mouth. Additionally, notice how these muscles work when you are speaking.

The pocket-like part of the mouth that is framed on the inside by the gums and teeth, and on the outside by the cheeks and lips is called the oral vestibule. Moving farther into the mouth, the opening between the oral cavity and throat (oropharynx) is called the fauces (like the kitchen "faucet"). The main open area of the mouth, or oral cavity proper, runs from the gums and teeth to the fauces.

When you are chewing, you do not find it difficult to breathe simultaneously. The next time you have food in your mouth, notice how the arched shape of the roof of your mouth allows you to handle both digestion and respiration at the same time. This arch is called the palate. The anterior region of the palate serves as a wall (or septum) between the oral and nasal cavities as well as a rigid shelf against which the tongue can push food. It is created by the maxillary and palatine bones of the skull and, given its bony structure, is known as the hard palate. If you run your tongue along the roof of your mouth, you’ll notice that the hard palate ends in the posterior oral cavity, and the tissue becomes fleshier. This part of the palate, known as the soft palate, is composed mainly of skeletal muscle. You can therefore manipulate, subconsciously, the soft palate—for instance, to yawn, swallow, or sing (see Figure 1).

[caption id="" align="aligncenter" width="480"]This diagram shows the structure of the mouth. The teeth, lips, tongue, gums and many other parts are labeled. Figure 1. Mouth. The mouth includes the lips, tongue, palate, gums, and teeth.[/caption]

A fleshy bead of tissue called the uvula drops down from the center of the posterior edge of the soft palate. Although some have suggested that the uvula is a vestigial organ, it serves an important purpose. When you swallow, the soft palate and uvula move upward, helping to keep foods and liquid from entering the nasal cavity. Unfortunately, it can also contribute to the sound produced by snoring. Two muscular folds extend downward from the soft palate, on either side of the uvula. Toward the front, the palatoglossal arch lies next to the base of the tongue; behind it, the palatopharyngeal arch forms the superior and lateral margins of the fauces. Between these two arches are the palatine tonsils, clusters of lymphoid tissue that protect the pharynx. The lingual tonsils are located at the base of the tongue.

The Tongue

Perhaps you have heard it said that the tongue is the strongest muscle in the body. Those who stake this claim cite its strength proportionate to its size. Although it is difficult to quantify the relative strength of different muscles, it remains indisputable that the tongue is a workhorse, facilitating ingestion, mechanical digestion, chemical digestion (lingual lipase), sensation (of taste, texture, and temperature of food), swallowing, and vocalization.

The tongue is attached to the mandible, the styloid processes of the temporal bones, and the hyoid bone. The hyoid is unique in that it only distantly/indirectly articulates with other bones. The tongue is positioned over the floor of the oral cavity. A medial septum extends the entire length of the tongue, dividing it into symmetrical halves.

Beneath its mucous membrane covering, each half of the tongue is composed of the same number and type of intrinsic and extrinsic skeletal muscles. The intrinsic muscles (those within the tongue) are the longitudinalis inferior, longitudinalis superior, transversus linguae, and verticalis linguae muscles. These allow you to change the size and shape of your tongue, as well as to stick it out, if you wish. Having such a flexible tongue facilitates both swallowing and speech.

As you learned in your study of the muscular system, the extrinsic muscles of the tongue are the mylohyoid, hyoglossus, styloglossus, and genioglossus muscles. These muscles originate outside the tongue and insert into connective tissues within the tongue. The mylohyoid is responsible for raising the tongue, the hyoglossus pulls it down and back, the styloglossus pulls it up and back, and the genioglossus pulls it forward. Working in concert, these muscles perform three important digestive functions in the mouth: (1) position food for optimal chewing, (2) gather food into a bolus (rounded mass), and (3) position food so it can be swallowed.

The top and sides of the tongue are studded with papillae, extensions of lamina propria of the mucosa, which are covered in stratified squamous epithelium (Figure 2). Fungiform papillae, which are mushroom shaped, cover a large area of the tongue; they tend to be larger toward the rear of the tongue and smaller on the tip and sides. In contrast, filiform papillae are long and thin. Fungiform papillae contain taste buds, and filiform papillae have touch receptors that help the tongue move food around in the mouth. The filiform papillae create an abrasive surface that performs mechanically, much like a cat’s rough tongue that is used for grooming. Lingual glands in the lamina propria of the tongue secrete mucus and a watery serous fluid that contains the enzyme lingual lipase, which plays a minor role in breaking down triglycerides but does not begin working until it is activated in the stomach. A fold of mucous membrane on the underside of the tongue, the lingual frenulum, tethers the tongue to the floor of the mouth. People with the congenital anomaly ankyloglossia, also known by the non-medical term “tongue tie,” have a lingual frenulum that is too short or otherwise malformed. Severe ankyloglossia can impair speech and must be corrected with surgery.

[caption id="" align="aligncenter" width="330"]This diagram shows the structure of the tongue and different parts of the tongue are labeled. Figure 2. Tongue. This superior view of the tongue shows the locations and types of lingual papillae.[/caption]

The Salivary Glands

Many small salivary glands are housed within the mucous membranes of the mouth and tongue. These minor exocrine glands are constantly secreting saliva, either directly into the oral cavity or indirectly through ducts, even while you sleep. In fact, an average of 1 to 1.5 liters of saliva is secreted each day. Usually just enough saliva is present to moisten the mouth and teeth. Secretion increases when you eat, because saliva is essential to moisten food and initiate the chemical breakdown of carbohydrates. Small amounts of saliva are also secreted by the labial glands in the lips. In addition, the buccal glands in the cheeks, palatal glands in the palate, and lingual glands in the tongue help ensure that all areas of the mouth are supplied with adequate saliva.

The Major Salivary Glands

Outside the oral mucosa are three pairs of major salivary glands, which secrete the majority of saliva into ducts that open into the mouth:

  • The submandibular glands, which are in the floor of the mouth, secrete saliva into the mouth through the submandibular ducts.
  • The sublingual glands, which lie below the tongue, use the lesser sublingual ducts to secrete saliva into the oral cavity.
  • The parotid glands lie between the skin and the masseter muscle, near the ears. They secrete saliva into the mouth through the parotid duct, which is located near the second upper molar tooth (Figure 3).

Saliva

Saliva is essentially (95.5 percent) water. The remaining 4.5 percent is a complex mixture of ions, glycoproteins, enzymes, growth factors, and waste products. Perhaps the most important ingredient in salvia from the perspective of digestion is the enzyme salivary amylase, which initiates the breakdown of carbohydrates. Food does not spend enough time in the mouth to allow all the carbohydrates to break down, but salivary amylase continues acting until it is inactivated by stomach acids. Bicarbonate and phosphate ions function as chemical buffers, maintaining saliva at a pH between 6.35 and 6.85. Salivary mucus helps lubricate food, facilitating movement in the mouth, bolus formation, and swallowing. Saliva contains immunoglobulin A, which prevents microbes from penetrating the epithelium, and lysozyme, which makes saliva antimicrobial. Saliva also contains epidermal growth factor, which might have given rise to the adage “a mother’s kiss can heal a wound.”

Each of the major salivary glands secretes a unique formulation of saliva according to its cellular makeup. For example, the parotid glands secrete a watery solution that contains salivary amylase. The submandibular glands have cells similar to those of the parotid glands, as well as mucus-secreting cells. Therefore, saliva secreted by the submandibular glands also contains amylase but in a liquid thickened with mucus. The sublingual glands contain mostly mucous cells, and they secrete the thickest saliva with the least amount of salivary amylase.

[caption id="" align="aligncenter" width="300"]This image shows the location of the salivary glands with reference to the teeth. The different salivary glands are labeled. Figure 3. Salivary glands. The major salivary glands are located outside the oral mucosa and deliver saliva into the mouth through ducts.[/caption]

Regulation of Salivation

The autonomic nervous system regulates salivation (the secretion of saliva). In the absence of food, parasympathetic stimulation keeps saliva flowing at just the right level for comfort as you speak, swallow, sleep, and generally go about life. Over-salivation can occur, for example, if you are stimulated by the smell of food, but that food is not available for you to eat. Drooling is an extreme instance of the overproduction of saliva. During times of stress, such as before speaking in public, sympathetic stimulation takes over, reducing salivation and producing the symptom of dry mouth often associated with anxiety. When you are dehydrated, salivation is reduced, causing the mouth to feel dry and prompting you to take action to quench your thirst.

Salivation can be stimulated by the sight, smell, and taste of food. It can even be stimulated by thinking about food. You might notice whether reading about food and salivation right now has had any effect on your production of saliva.

How does the salivation process work while you are eating? Food contains chemicals that stimulate taste receptors on the tongue, which send impulses to the superior and inferior salivatory nuclei in the brain stem. These two nuclei then send back parasympathetic impulses through fibers in the glossopharyngeal and facial nerves, which stimulate salivation. Even after you swallow food, salivation is increased to cleanse the mouth and to water down and neutralize any irritating chemical remnants, such as that hot sauce in your burrito. Most saliva is swallowed along with food and is reabsorbed, so that fluid is not lost.

The Teeth

The teeth, or dentes (singular = dens), are organs similar to bones that you use to tear, grind, and otherwise mechanically break down food.

Types of Teeth

During the course of your lifetime, you have two sets of teeth (one set of teeth is a dentition). Your 20 deciduous teeth, or baby teeth, first begin to appear at about 6 months of age. Between approximately age 6 and 12, these teeth are replaced by 32 permanent teeth. Moving from the center of the mouth toward the side, these are as follows (Figure 4):

  • The eight incisors, four top and four bottom, are the sharp front teeth you use for biting into food.
  • The four cuspids (or canines) flank the incisors and have a pointed edge (cusp) to tear up food. These fang-like teeth are superb for piercing tough or fleshy foods.
  • Posterior to the cuspids are the eight premolars (or bicuspids), which have an overall flatter shape with two rounded cusps useful for mashing foods.
  • The most posterior and largest are the 12 molars, which have several pointed cusps used to crush food so it is ready for swallowing. The third members of each set of three molars, top and bottom, are commonly referred to as the wisdom teeth, because their eruption is commonly delayed until early adulthood. It is not uncommon for wisdom teeth to fail to erupt; that is, they remain impacted. In these cases, the teeth are typically removed by orthodontic surgery.
[caption id="" align="aligncenter" width="350"]This diagram shows the arrangement of permanent and deciduous teeth in human. The permanent teeth are labeled along with the average age at which they emerge. An inset shows the arrangement of the deciduous teeth, with the age at which they emerge listed. Figure 4. Permanent and Deciduous Teeth. This figure of two human dentitions shows the arrangement of teeth in the maxilla and mandible, and the relationship between the deciduous and permanent teeth.[/caption]

Anatomy of a Tooth

The teeth are secured in the alveolar processes (sockets) of the maxilla and the mandible. Gingivae (commonly called the gums) are soft tissues that line the alveolar processes and surround the necks of the teeth. Teeth are also held in their sockets by a connective tissue called the periodontal ligament.

The two main parts of a tooth are the crown, which is the portion projecting above the gum line, and the root, which is embedded within the maxilla and mandible. Both parts contain an inner pulp cavity, containing loose connective tissue through which run nerves and blood vessels. The region of the pulp cavity that runs through the root of the tooth is called the root canal. Surrounding the pulp cavity is dentin, a bone-like tissue. In the root of each tooth, the dentin is covered by an even harder bone-like layer called cementum. In the crown of each tooth, the dentin is covered by an outer layer of enamel, the hardest substance in the body (Figure 5).

Although enamel protects the underlying dentin and pulp cavity, it is still nonetheless susceptible to mechanical and chemical erosion, or what is known as tooth decay. The most common form, dental caries (cavities) develops when colonies of bacteria feeding on sugars in the mouth release acids that cause soft tissue inflammation and degradation of the calcium crystals of the enamel. The digestive functions of the mouth are summarized in Table 4.

[caption id="" align="aligncenter" width="350"]This diagram shows a cross-section of a human tooth elucidating its structure. The major parts of the tooth along with the blood vessels are shown. Figure 5. The Structure of the Tooth. This longitudinal section through a molar in its alveolar socket shows the relationships between enamel, dentin, and pulp.[/caption]
Digestive Functions of the Mouth (Table 4)
Structure Action Outcome
Lips and cheeks Confine food between teeth
  • Food is chewed evenly during mastication
Salivary glands Secrete saliva
  • Moisten and lubricate the lining of the mouth and pharynx
  • Moisten, soften, and dissolve food
  • Clean the mouth and teeth
  • Salivary amylase breaks down starch
Tongue’s extrinsic muscles Move tongue sideways, and in and out
  • Manipulate food for chewing
  • Shape food into a bolus
  • Manipulate food for swallowing
Tongue’s intrinsic muscles Change tongue shape
  • Manipulate food for swallowing
Taste buds Sense food in mouth and sense taste
  • Nerve impulses from taste buds are conducted to salivary nuclei in the brain stem and then to salivary glands, stimulating saliva secretion
Lingual glands Secrete lingual lipase
  • Activated in the stomach
  • Break down triglycerides into fatty acids and diglycerides
Teeth Shred and crush food
  • Break down solid food into smaller particles for deglutition

The Pharynx

The pharynx (throat) is involved in both digestion and respiration. It receives food and air from the mouth, and air from the nasal cavities. When food enters the pharynx, involuntary muscle contractions close off the air passageways.

A short tube of skeletal muscle lined with a mucous membrane, the pharynx runs from the posterior oral and nasal cavities to the opening of the esophagus and larynx. It has three subdivisions. The most superior, the nasopharynx, is involved only in breathing and speech. The other two subdivisions, the oropharynx and the laryngopharynx, are used for both breathing and digestion. The oropharynx begins inferior to the nasopharynx and is continuous below with the laryngopharynx (Figure 6). The inferior border of the laryngopharynx connects to the esophagus, whereas the anterior portion connects to the larynx, allowing air to flow into the bronchial tree.

[caption id="" align="aligncenter" width="380"]This diagram shows the cross-section of a human face and highlights the location of the pharynx, which runs from the nostrils to the esophagus and the larynx. Figure 6. Pharynx. The pharynx runs from the nostrils to the esophagus and the larynx.[/caption]

Histologically, the wall of the oropharynx is similar to that of the oral cavity. The mucosa includes a stratified squamous epithelium that is endowed with mucus-producing glands. During swallowing, the elevator skeletal muscles of the pharynx contract, raising and expanding the pharynx to receive the bolus of food. Once received, these muscles relax and the constrictor muscles of the pharynx contract, forcing the bolus into the esophagus and initiating peristalsis.

Usually during swallowing, the soft palate and uvula rise reflexively to close off the entrance to the nasopharynx. At the same time, the larynx is pulled superiorly and the cartilaginous epiglottis, its most superior structure, folds inferiorly, covering the glottis (the opening to the larynx); this process effectively blocks access to the trachea and bronchi. When the food “goes down the wrong way,” it goes into the trachea. When food enters the trachea, the reaction is to cough, which usually forces the food up and out of the trachea, and back into the pharynx.

The Esophagus

The esophagus is a muscular tube that connects the pharynx to the stomach. It is approximately 25.4 cm (10 in) in length, located posterior to the trachea, and remains in a collapsed form when not engaged in swallowing. As you can see in Figure 7, the esophagus runs a mainly straight route through the mediastinum of the thorax. To enter the abdomen, the esophagus penetrates the diaphragm through an opening called the esophageal hiatus.

Passage of Food through the Esophagus

The upper esophageal sphincter, which is continuous with the inferior pharyngeal constrictor, controls the movement of food from the pharynx into the esophagus. The upper two-thirds of the esophagus consists of both smooth and skeletal muscle fibers, with the latter fading out in the bottom third of the esophagus. Rhythmic waves of peristalsis, which begin in the upper esophagus, propel the bolus of food toward the stomach. Meanwhile, secretions from the esophageal mucosa lubricate the esophagus and food. Food passes from the esophagus into the stomach at the lower esophageal sphincter (also called the gastroesophageal or cardiac sphincter). Recall that sphincters are muscles that surround tubes and serve as valves, closing the tube when the sphincters contract and opening it when they relax. The lower esophageal sphincter relaxes to let food pass into the stomach, and then contracts to prevent stomach acids from backing up into the esophagus. Surrounding this sphincter is the muscular diaphragm, which helps close off the sphincter when no food is being swallowed. When the lower esophageal sphincter does not completely close, the stomach’s contents can reflux (that is, back up into the esophagus), causing heartburn or gastroesophageal reflux disease (GERD).

[caption id="" align="aligncenter" width="320"]This diagram shows the esophagus, going from the mouth to the stomach. The upper and the lower esophageal sphincter are labeled. Figure 7. Esophagus. The upper esophageal sphincter controls the movement of food from the pharynx to the esophagus. The lower esophageal sphincter controls the movement of food from the esophagus to the stomach.[/caption]

Histology of the Esophagus

The mucosa of the esophagus is made up of an epithelial lining that contains non-keratinized, stratified squamous epithelium, with a layer of basal and parabasal cells. This epithelium protects against erosion from food particles. The mucosa’s lamina propria contains mucus-secreting glands. The muscularis layer changes according to location: In the upper third of the esophagus, the muscularis is skeletal muscle. In the middle third, it is both skeletal and smooth muscle. In the lower third, it is smooth muscle. As mentioned previously, the most superficial layer of the esophagus is called the adventitia, not the serosa. In contrast to the stomach and intestines, the loose connective tissue of the adventitia is not covered by a fold of visceral peritoneum. The digestive functions of the esophagus are identified in Table 5.

Digestive Functions of the Esophagus (Table 5)
Action Outcome
Upper esophageal sphincter relaxation Allows the bolus to move from the laryngopharynx to the esophagus
Peristalsis Propels the bolus through the esophagus
Lower esophageal sphincter relaxation Allows the bolus to move from the esophagus into the stomach and prevents chime from entering the esophagus
Mucus secretion Lubricates the esophagus, allowing easy passage of the bolus

Deglutition

Deglutition is another word for swallowing—the movement of food from the mouth to the stomach. The entire process takes about 4 to 8 seconds for solid or semisolid food, and about 1 second for very soft food and liquids. Although this sounds quick and effortless, deglutition is, in fact, a complex process that involves both the skeletal muscle of the tongue and the muscles of the pharynx and esophagus. It is aided by the presence of mucus and saliva. There are three stages in deglutition: the voluntary phase, the pharyngeal phase, and the esophageal phase (Figure 8). The autonomic nervous system controls the latter two phases.

[caption id="" align="aligncenter" width="520"]This figure shows the three different phases of deglutition. The left panel shows the voluntary phase, the middle panel shows the pharyngeal phase and the right panel’s shows the esophageal phase. Figure 8. Deglutition. Deglutition includes the voluntary phase and two involuntary phases: the pharyngeal phase and the esophageal phase.[/caption]

The Voluntary Phase

The voluntary phase of deglutition (also known as the oral or buccal phase) is so called because you can control when you swallow food. In this phase, chewing has been completed and swallowing is set in motion. The tongue moves upward and backward against the palate, pushing the bolus to the back of the oral cavity and into the oropharynx. Other muscles keep the mouth closed and prevent food from falling out. At this point, the two involuntary phases of swallowing begin.

The Pharyngeal Phase

In the pharyngeal phase, stimulation of receptors in the oropharynx sends impulses to the deglutition center (a collection of neurons that controls swallowing) in the medulla oblongata. Impulses are then sent back to the uvula and soft palate, causing them to move upward and close off the nasopharynx. The laryngeal muscles also constrict to prevent aspiration of food into the trachea. At this point, deglutition apnea takes place, which means that breathing ceases for a very brief time. Contractions of the pharyngeal constrictor muscles move the bolus through the oropharynx and laryngopharynx. Relaxation of the upper esophageal sphincter then allows food to enter the esophagus.

The Esophageal Phase

The entry of food into the esophagus marks the beginning of the esophageal phase of deglutition and the initiation of peristalsis. As in the previous phase, the complex neuromuscular actions are controlled by the medulla oblongata. Peristalsis propels the bolus through the esophagus and toward the stomach. The circular muscle layer of the muscularis contracts, pinching the esophageal wall and forcing the bolus forward. At the same time, the longitudinal muscle layer of the muscularis also contracts, shortening this area and pushing out its walls to receive the bolus. In this way, a series of contractions keeps moving food toward the stomach. When the bolus nears the stomach, distention of the esophagus initiates a short reflex relaxation of the lower esophageal sphincter that allows the bolus to pass into the stomach. During the esophageal phase, esophageal glands secrete mucus that lubricates the bolus and minimizes friction.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this animation to see how swallowing is a complex process that involves the nervous system to coordinate the actions of upper respiratory and digestive activities.[/caption]

]]> 2684 2664 4 0 23.4 The Stomach https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2689 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2689

Learning Objectives

By the end of this section, you will be able to:
  • Label on a diagram the four main regions of the stomach, its curvatures, and its sphincter
  • Identify the four main types of secreting cells in gastric glands, and their important products
  • Explain why the stomach does not digest itself
  • Describe the mechanical and chemical digestion of food entering the stomach

Although a minimal amount of carbohydrate digestion occurs in the mouth, chemical digestion really gets underway in the stomach. An expansion of the alimentary canal that lies immediately inferior to the esophagus, the stomach links the esophagus to the first part of the small intestine (the duodenum) and is relatively fixed in place at its esophageal and duodenal ends. In between, however, it can be a highly active structure, contracting and continually changing position and size. These contractions provide mechanical assistance to digestion. The empty stomach is only about the size of your fist, but can stretch to hold as much as 4 liters of food and fluid, or more than 75 times its empty volume, and then return to its resting size when empty. Although you might think that the size of a person’s stomach is related to how much food that individual consumes, body weight does not correlate with stomach size. Rather, when you eat greater quantities of food—such as at holiday dinner—you stretch the stomach more than when you eat less.

Popular culture tends to refer to the stomach as the location where all digestion takes place. Of course, this is not true. An important function of the stomach is to serve as a temporary holding chamber. You can ingest a meal far more quickly than it can be digested and absorbed by the small intestine. Thus, the stomach holds food and parses only small amounts into the small intestine at a time. Foods are not processed in the order they are eaten; rather, they are mixed together with digestive juices in the stomach until they are converted into chyme, which is released into the small intestine.

As you will see in the sections that follow, the stomach plays several important roles in chemical digestion, including the continued digestion of carbohydrates and the initial digestion of proteins and triglycerides. Little if any nutrient absorption occurs in the stomach, with the exception of the negligible amount of nutrients in alcohol.

Structure

There are four main regions in the stomach: the cardia, fundus, body, and pylorus (Figure 1). The cardia (or cardiac region) is the point where the esophagus connects to the stomach and through which food passes into the stomach. Located inferior to the diaphragm, above and to the left of the cardia, is the dome-shaped fundus. Below the fundus is the body, the main part of the stomach. The funnel-shaped pylorus connects the stomach to the duodenum. The wider end of the funnel, the pyloric antrum, connects to the body of the stomach. The narrower end is called the pyloric canal, which connects to the duodenum. The smooth muscle pyloric sphincter is located at this latter point of connection and controls stomach emptying. In the absence of food, the stomach deflates inward, and its mucosa and submucosa fall into a large fold called a ruga.

[caption id="" align="aligncenter" width="520"]This image shows a cross-section of the stomach, and the major parts: the cardia, fundus, body and pylorus are labeled. Figure 1. Stomach. The stomach has four major regions: the cardia, fundus, body, and pylorus. The addition of an inner oblique smooth muscle layer gives the muscularis the ability to vigorously churn and mix food.[/caption]

The convex lateral surface of the stomach is called the greater curvature; the concave medial border is the lesser curvature. The stomach is held in place by the lesser omentum, which extends from the liver to the lesser curvature, and the greater omentum, which runs from the greater curvature to the posterior abdominal wall.

Histology

The wall of the stomach is made of the same four layers as most of the rest of the alimentary canal, but with adaptations to the mucosa and muscularis for the unique functions of this organ. In addition to the typical circular and longitudinal smooth muscle layers, the muscularis has an inner oblique smooth muscle layer (Figure 2). As a result, in addition to moving food through the canal, the stomach can vigorously churn food, mechanically breaking it down into smaller particles.

[caption id="" align="aligncenter" width="500"]This diagram shows the histological cross-section of the stomach. The left panel shows the stomach and the center panel shows a magnified view of a small region including the epithelium and the gastric glands. The right panel shows a further magnification of the mucosa and the different cell types are labeled. Figure 2. Histology of the Stomach. The stomach wall is adapted for the functions of the stomach. In the epithelium, gastric pits lead to gastric glands that secrete gastric juice. The gastric glands (one gland is shown enlarged on the right) contain different types of cells that secrete a variety of enzymes, including hydrochloride acid, which activates the protein-digesting enzyme pepsin.[/caption]

The stomach mucosa’s epithelial lining consists only of surface mucus cells, which secrete a protective coat of alkaline mucus. A vast number of gastric pits dot the surface of the epithelium, giving it the appearance of a well-used pincushion, and mark the entry to each gastric gland, which secretes a complex digestive fluid referred to as gastric juice.

Although the walls of the gastric pits are made up primarily of mucus cells, the gastric glands are made up of different types of cells. The glands of the cardia and pylorus are composed primarily of mucus-secreting cells. Cells that make up the pyloric antrum secrete mucus and a number of hormones, including the majority of the stimulatory hormone, gastrin. The much larger glands of the fundus and body of the stomach, the site of most chemical digestion, produce most of the gastric secretions. These glands are made up of a variety of secretory cells. These include parietal cells, chief cells, mucous neck cells, and enteroendocrine cells.

Parietal cells—Located primarily in the middle region of the gastric glands are parietal cells, which are among the most highly differentiated of the body’s epithelial cells. These relatively large cells produce both hydrochloric acid (HCl) and intrinsic factor. HCl is responsible for the high acidity (pH 1.5 to 3.5) of the stomach contents and is needed to activate the protein-digesting enzyme, pepsin. The acidity also kills much of the bacteria you ingest with food and helps to denature proteins, making them more available for enzymatic digestion. Intrinsic factor is a glycoprotein necessary for the absorption of vitamin B12 in the small intestine.

Chief cells—Located primarily in the basal regions of gastric glands are chief cells, which secrete pepsinogen, the inactive proenzyme form of pepsin. HCl is necessary for the conversion of pepsinogen to pepsin.

Mucous neck cells—Gastric glands in the upper part of the stomach contain mucous neck cells that secrete thin, acidic mucus that is much different from the mucus secreted by the goblet cells of the surface epithelium. The role of this mucus is not currently known.

Enteroendocrine cells—Finally, enteroendocrine cells found in the gastric glands secrete various hormones into the interstitial fluid of the lamina propria. These include gastrin, which is released mainly by enteroendocrine G cells.

Table 6 describes the digestive functions of important hormones secreted by the stomach.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this animation that depicts the structure of the stomach and how this structure functions in the initiation of protein digestion.[/caption]
Hormones Secreted by the Stomach (Table 6)
Hormone Production site Production stimulus Target organ Action
Gastrin Stomach mucosa, mainly G cells of the pyloric antrum Presence of peptides and amino acids in stomach Stomach Increases secretion by gastric glands; promotes gastric emptying
Gastrin Stomach mucosa, mainly G cells of the pyloric antrum Presence of peptides and amino acids in stomach Small intestine Promotes intestinal muscle contraction
Gastrin Stomach mucosa, mainly G cells of the pyloric antrum Presence of peptides and amino acids in stomach Ileocecal valve Relaxes valve
Gastrin Stomach mucosa, mainly G cells of the pyloric antrum Presence of peptides and amino acids in stomach Large intestine Triggers mass movements
Ghrelin Stomach mucosa, mainly fundus Fasting state (levels increase just prior to meals) Hypothalamus Regulates food intake, primarily by stimulating hunger and satiety
Histamine Stomach mucosa Presence of food in the stomach Stomach Stimulates parietal cells to release HCl
Serotonin Stomach mucosa Presence of food in the stomach Stomach Contracts stomach muscle
Somatostatin Mucosa of stomach, especially pyloric antrum; also duodenum Presence of food in the stomach; sympathetic axon stimulation Stomach Restricts all gastric secretions, gastric motility, and emptying
Somatostatin Mucosa of stomach, especially pyloric antrum; also duodenum Presence of food in the stomach; sympathetic axon stimulation Pancreas Restricts pancreatic secretions
Somatostatin Mucosa of stomach, especially pyloric antrum; also duodenum Presence of food in the stomach; sympathetic axon stimulation Small intestine Reduces intestinal absorption by reducing blood flow

Gastric Secretion

The secretion of gastric juice is controlled by both nerves and hormones. Stimuli in the brain, stomach, and small intestine activate or inhibit gastric juice production. This is why the three phases of gastric secretion are called the cephalic, gastric, and intestinal phases (Figure 3). However, once gastric secretion begins, all three phases can occur simultaneously.
[caption id="" align="aligncenter" width="450"]This flowchart shows the three different phases of gastric secretion. The top panel shows the cephalic phase, the middle panel shows the gastric phase and the bottom panel shows the intestinal phase. Figure 3. The Three Phases of Gastric Secretion. Gastric secretion occurs in three phases: cephalic, gastric, and intestinal. During each phase, the secretion of gastric juice can be stimulated or inhibited.[/caption]

The cephalic phase (reflex phase) of gastric secretion, which is relatively brief, takes place before food enters the stomach. The smell, taste, sight, or thought of food triggers this phase. For example, when you bring a piece of sushi to your lips, impulses from receptors in your taste buds or the nose are relayed to your brain, which returns signals that increase gastric secretion to prepare your stomach for digestion. This enhanced secretion is a conditioned reflex, meaning it occurs only if you like or want a particular food. Depression and loss of appetite can suppress the cephalic reflex.

The gastric phase of secretion lasts 3 to 4 hours, and is set in motion by local neural and hormonal mechanisms triggered by the entry of food into the stomach. For example, when your sushi reaches the stomach, it creates distention that activates the stretch receptors. This stimulates parasympathetic neurons to release acetylcholine, which then provokes increased secretion of gastric juice. Partially digested proteins, caffeine, and rising pH stimulate the release of gastrin from enteroendocrine G cells, which in turn induces parietal cells to increase their production of HCl, which is needed to create an acidic environment for the conversion of pepsinogen to pepsin, and protein digestion. Additionally, the release of gastrin activates vigorous smooth muscle contractions. However, it should be noted that the stomach does have a natural means of avoiding excessive acid secretion and potential heartburn. Whenever pH levels drop too low, cells in the stomach react by suspending HCl secretion and increasing mucous secretions.

The intestinal phase of gastric secretion has both excitatory and inhibitory elements. The duodenum has a major role in regulating the stomach and its emptying. When partially digested food fills the duodenum, intestinal mucosal cells release a hormone called intestinal (enteric) gastrin, which further excites gastric juice secretion. This stimulatory activity is brief, however, because when the intestine distends with chyme, the enterogastric reflex inhibits secretion. One of the effects of this reflex is to close the pyloric sphincter, which blocks additional chyme from entering the duodenum.

The Mucosal Barrier

The mucosa of the stomach is exposed to the highly corrosive acidity of gastric juice. Gastric enzymes that can digest protein can also digest the stomach itself. The stomach is protected from self-digestion by the mucosal barrier. This barrier has several components. First, the stomach wall is covered by a thick coating of bicarbonate-rich mucus. This mucus forms a physical barrier, and its bicarbonate ions neutralize acid. Second, the epithelial cells of the stomach's mucosa meet at tight junctions, which block gastric juice from penetrating the underlying tissue layers. Finally, stem cells located where gastric glands join the gastric pits quickly replace damaged epithelial mucosal cells, when the epithelial cells are shed. In fact, the surface epithelium of the stomach is completely replaced every 3 to 6 days.

Homeostatic Imbalances

Ulcers: When the Mucosal Barrier Breaks Down As effective as the mucosal barrier is, it is not a “fail-safe” mechanism. Sometimes, gastric juice eats away at the superficial lining of the stomach mucosa, creating erosions, which mostly heal on their own. Deeper and larger erosions are called ulcers.

Why does the mucosal barrier break down? A number of factors can interfere with its ability to protect the stomach lining. The majority of all ulcers are caused by either excessive intake of non-steroidal anti-inflammatory drugs (NSAIDs), including aspirin, or Helicobacter pylori infection.

Antacids help relieve symptoms of ulcers such as “burning” pain and indigestion. When ulcers are caused by NSAID use, switching to other classes of pain relievers allows healing. When caused by H. pylori infection, antibiotics are effective.

A potential complication of ulcers is perforation: Perforated ulcers create a hole in the stomach wall, resulting in peritonitis (inflammation of the peritoneum). These ulcers must be repaired surgically.

Digestive Functions of the Stomach

The stomach participates in virtually all the digestive activities with the exception of ingestion and defecation. Although almost all absorption takes place in the small intestine, the stomach does absorb some nonpolar substances, such as alcohol and aspirin.

Mechanical Digestion

Within a few moments after food after enters your stomach, mixing waves begin to occur at intervals of approximately 20 seconds. A mixing wave is a unique type of peristalsis that mixes and softens the food with gastric juices to create chyme. The initial mixing waves are relatively gentle, but these are followed by more intense waves, starting at the body of the stomach and increasing in force as they reach the pylorus. It is fair to say that long before your sushi exits through the pyloric sphincter, it bears little resemblance to the sushi you ate. The pylorus, which holds around 30 mL (1 fluid ounce) of chyme, acts as a filter, permitting only liquids and small food particles to pass through the mostly, but not fully, closed pyloric sphincter. In a process called gastric emptying, rhythmic mixing waves force about 3 mL of chyme at a time through the pyloric sphincter and into the duodenum. Release of a greater amount of chyme at one time would overwhelm the capacity of the small intestine to handle it. The rest of the chyme is pushed back into the body of the stomach, where it continues mixing. This process is repeated when the next mixing waves force more chyme into the duodenum.

Gastric emptying is regulated by both the stomach and the duodenum. The presence of chyme in the duodenum activates receptors that inhibit gastric secretion. This prevents additional chyme from being released by the stomach before the duodenum is ready to process it.

Chemical Digestion

The fundus plays an important role, because it stores both undigested food and gases that are released during the process of chemical digestion. Food may sit in the fundus of the stomach for a while before being mixed with the chyme. While the food is in the fundus, the digestive activities of salivary amylase continue until the food begins mixing with the acidic chyme. Ultimately, mixing waves incorporate this food with the chyme, the acidity of which inactivates salivary amylase and activates lingual lipase. Lingual lipase then begins breaking down triglycerides into free fatty acids, and mono- and diglycerides.

The breakdown of protein begins in the stomach through the actions of HCl and the enzyme pepsin. During infancy, gastric glands also produce rennin, an enzyme that helps digest milk protein.

Its numerous digestive functions notwithstanding, there is only one stomach function necessary to life: the production of intrinsic factor. The intestinal absorption of vitamin B12, which is necessary for both the production of mature red blood cells and normal neurological functioning, cannot occur without intrinsic factor. People who undergo total gastrectomy (stomach removal)—for life-threatening stomach cancer, for example—can survive with minimal digestive dysfunction if they receive vitamin B12 injections.

The contents of the stomach are completely emptied into the duodenum within 2 to 4 hours after you eat a meal. Different types of food take different amounts of time to process. Foods heavy in carbohydrates empty fastest, followed by high-protein foods. Meals with a high triglyceride content remain in the stomach the longest. Since enzymes in the small intestine digest fats slowly, food can stay in the stomach for 6 hours or longer when the duodenum is processing fatty chyme. However, note that this is still a fraction of the 24 to 72 hours that full digestion typically takes from start to finish.
]]> 2689 2664 5 0 23.6 Accessory Organs in Digestion: The Liver, Pancreas, and Gallbladder https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2703 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2703

Learning Objectives

By the end of this section, you will be able to:
  • State the main digestive roles of the liver, pancreas, and gallbladder
  • Identify three main features of liver histology that are critical to its function
  • Discuss the composition and function of bile
  • Identify the major types of enzymes and buffers present in pancreatic juice

Chemical digestion in the small intestine relies on the activities of three accessory digestive organs: the liver, pancreas, and gallbladder (Figure 1). The digestive role of the liver is to produce bile and export it to the duodenum. The gallbladder primarily stores, concentrates, and releases bile. The pancreas produces pancreatic juice, which contains digestive enzymes and bicarbonate ions, and delivers it to the duodenum.

[caption id="" align="aligncenter" width="390"]This diagram shows the accessory organs of the digestive system. The liver, spleen, pancreas, gallbladder and their major parts are shown. Figure 1. Accessory Organs. The liver, pancreas, and gallbladder are considered accessory digestive organs, but their roles in the digestive system are vital.[/caption]

The Liver

The liver is the largest gland in the body, weighing about three pounds in an adult. It is also one of the most important organs. In addition to being an accessory digestive organ, it plays a number of roles in metabolism and regulation. The liver lies inferior to the diaphragm in the right upper quadrant of the abdominal cavity and receives protection from the surrounding ribs.

The liver is divided into two primary lobes: a large right lobe and a much smaller left lobe. In the right lobe, some anatomists also identify an inferior quadrate lobe and a posterior caudate lobe, which are defined by internal features. The liver is connected to the abdominal wall and diaphragm by five peritoneal folds referred to as ligaments. These are the falciform ligament, the coronary ligament, two lateral ligaments, and the ligamentum teres hepatis. The falciform ligament and ligamentum teres hepatis are actually remnants of the umbilical vein, and separate the right and left lobes anteriorly. The lesser omentum tethers the liver to the lesser curvature of the stomach.

The porta hepatis (“gate to the liver”) is where the hepatic artery and hepatic portal vein enter the liver. These two vessels, along with the common hepatic duct, run behind the lateral border of the lesser omentum on the way to their destinations. As shown in Figure 2, the hepatic artery delivers oxygenated blood from the heart to the liver. The hepatic portal vein delivers partially deoxygenated blood containing nutrients absorbed from the small intestine and actually supplies more oxygen to the liver than do the much smaller hepatic arteries. In addition to nutrients, drugs and toxins are also absorbed. After processing the bloodborne nutrients and toxins, the liver releases nutrients needed by other cells back into the blood, which drains into the central vein and then through the hepatic vein to the inferior vena cava. With this hepatic portal circulation, all blood from the alimentary canal passes through the liver. This largely explains why the liver is the most common site for the metastasis of cancers that originate in the alimentary canal.

[caption id="" align="aligncenter" width="420"]This image shows the microscopic anatomy of the liver. The top panel shows the liver; the center panel shows a magnified view of the connective tissue and the lobules. The bottom panel shows a further magnified view of a lobule, identifying the veins, bile duct and the sinusoids. Figure 2. Microscopic Anatomy of the Liver. The liver receives oxygenated blood from the hepatic artery and nutrient-rich deoxygenated blood from the hepatic portal vein.[/caption]

Histology

The liver has three main components: hepatocytes, bile canaliculi, and hepatic sinusoids. A hepatocyte is the liver’s main cell type, accounting for around 80 percent of the liver's volume. These cells play a role in a wide variety of secretory, metabolic, and endocrine functions. Plates of hepatocytes called hepatic laminae radiate outward from the portal vein in each hepatic lobule.

Between adjacent hepatocytes, grooves in the cell membranes provide room for each bile canaliculus (plural = canaliculi). These small ducts accumulate the bile produced by hepatocytes. From here, bile flows first into bile ductules and then into bile ducts. The bile ducts unite to form the larger right and left hepatic ducts, which themselves merge and exit the liver as the common hepatic duct. This duct then joins with the cystic duct from the gallbladder, forming the common bile duct through which bile flows into the small intestine. A hepatic sinusoid is an open, porous blood space formed by fenestrated capillaries from nutrient-rich hepatic portal veins and oxygen-rich hepatic arteries. Hepatocytes are tightly packed around the fenestrated endothelium of these spaces, giving them easy access to the blood. From their central position, hepatocytes process the nutrients, toxins, and waste materials carried by the blood. Materials such as bilirubin are processed and excreted into the bile canaliculi. Other materials including proteins, lipids, and carbohydrates are processed and secreted into the sinusoids or just stored in the cells until called upon. The hepatic sinusoids combine and send blood to a central vein. Blood then flows through a hepatic vein into the inferior vena cava. This means that blood and bile flow in opposite directions. The hepatic sinusoids also contain star-shaped reticuloendothelial cells (Kupffer cells), phagocytes that remove dead red and white blood cells, bacteria, and other foreign material that enter the sinusoids. The portal triad is a distinctive arrangement around the perimeter of hepatic lobules, consisting of three basic structures: a bile duct, a hepatic artery branch, and a hepatic portal vein branch.

Bile

Recall that lipids are hydrophobic, that is, they do not dissolve in water. Thus, before they can be digested in the watery environment of the small intestine, large lipid globules must be broken down into smaller lipid globules, a process called emulsification. Bile is a mixture secreted by the liver to accomplish the emulsification of lipids in the small intestine.

Hepatocytes secrete about one liter of bile each day. A yellow-brown or yellow-green alkaline solution (pH 7.6 to 8.6), bile is a mixture of water, bile salts, bile pigments, phospholipids (such as lecithin), electrolytes, cholesterol, and triglycerides. The components most critical to emulsification are bile salts and phospholipids, which have a nonpolar (hydrophobic) region as well as a polar (hydrophilic) region. The hydrophobic region interacts with the large lipid molecules, whereas the hydrophilic region interacts with the watery chyme in the intestine. This results in the large lipid globules being pulled apart into many tiny lipid fragments of about 1 µm in diameter. This change dramatically increases the surface area available for lipid-digesting enzyme activity. This is the same way dish soap works on fats mixed with water.

Bile salts act as emulsifying agents, so they are also important for the absorption of digested lipids. While most constituents of bile are eliminated in feces, bile salts are reclaimed by the enterohepatic circulation. Once bile salts reach the ileum, they are absorbed and returned to the liver in the hepatic portal blood. The hepatocytes then excrete the bile salts into newly formed bile. Thus, this precious resource is recycled.

Bilirubin, the main bile pigment, is a waste product produced when the spleen removes old or damaged red blood cells from the circulation. These breakdown products, including proteins, iron, and toxic bilirubin, are transported to the liver via the splenic vein of the hepatic portal system. In the liver, proteins and iron are recycled, whereas bilirubin is excreted in the bile. It accounts for the green color of bile. Bilirubin is eventually transformed by intestinal bacteria into stercobilin, a brown pigment that gives your stool its characteristic color! In some disease states, bile does not enter the intestine, resulting in white (‘acholic’) stool with a high fat content, since virtually no fats are broken down or absorbed.

Hepatocytes work non-stop, but bile production increases when fatty chyme enters the duodenum and stimulates the secretion of the gut hormone secretin. Between meals, bile is produced but conserved. The valve-like hepatopancreatic ampulla closes, allowing bile to divert to the gallbladder, where it is concentrated and stored until the next meal.

The Pancreas

The soft, oblong, glandular pancreas lies transversely in the retroperitoneum behind the stomach. Its head is nestled into the “c-shaped” curvature of the duodenum with the body extending to the left about 15.2 cm (6 in) and ending as a tapering tail in the hilum of the spleen. It is a curious mix of exocrine (secreting digestive enzymes) and endocrine (releasing hormones into the blood) functions (Figure 3).

[caption id="" align="aligncenter" width="350"]This figure shows the pancreas and its major parts. A magnified view of a small region of the pancreas shows the pancreatic islet cells, the acinar cells and the pancreatic duct. Figure 3. Exocrine and Endocrine Pancreas. The pancreas has a head, a body, and a tail. It delivers pancreatic juice to the duodenum through the pancreatic duct.[/caption]

The exocrine part of the pancreas arises as little grape-like cell clusters, each called an acinus (plural = acini), located at the terminal ends of pancreatic ducts. These acinar cells secrete enzyme-rich pancreatic juice into tiny merging ducts that form two dominant ducts. The larger duct fuses with the common bile duct (carrying bile from the liver and gallbladder) just before entering the duodenum via a common opening (the hepatopancreatic ampulla). The smooth muscle sphincter of the hepatopancreatic ampulla controls the release of pancreatic juice and bile into the small intestine. The second and smaller pancreatic duct, the accessory duct (duct of Santorini), runs from the pancreas directly into the duodenum, approximately 1 inch above the hepatopancreatic ampulla. When present, it is a persistent remnant of pancreatic development.

Scattered through the sea of exocrine acini are small islands of endocrine cells, the islets of Langerhans. These vital cells produce the hormones pancreatic polypeptide, insulin, glucagon, and somatostatin.

Pancreatic Juice

The pancreas produces over a liter of pancreatic juice each day. Unlike bile, it is clear and composed mostly of water along with some salts, sodium bicarbonate, and several digestive enzymes. Sodium bicarbonate is responsible for the slight alkalinity of pancreatic juice (pH 7.1 to 8.2), which serves to buffer the acidic gastric juice in chyme, inactivate pepsin from the stomach, and create an optimal environment for the activity of pH-sensitive digestive enzymes in the small intestine. Pancreatic enzymes are active in the digestion of sugars, proteins, and fats.

The pancreas produces protein-digesting enzymes in their inactive forms. These enzymes are activated in the duodenum. If produced in an active form, they would digest the pancreas (which is exactly what occurs in the disease, pancreatitis). The intestinal brush border enzyme enteropeptidase stimulates the activation of trypsin from trypsinogen of the pancreas, which in turn changes the pancreatic enzymes procarboxypeptidase and chymotrypsinogen into their active forms, carboxypeptidase and chymotrypsin.

The enzymes that digest starch (amylase), fat (lipase), and nucleic acids (nuclease) are secreted in their active forms, since they do not attack the pancreas as do the protein-digesting enzymes.

Pancreatic Secretion

Regulation of pancreatic secretion is the job of hormones and the parasympathetic nervous system. The entry of acidic chyme into the duodenum stimulates the release of secretin, which in turn causes the duct cells to release bicarbonate-rich pancreatic juice. The presence of proteins and fats in the duodenum stimulates the secretion of CCK, which then stimulates the acini to secrete enzyme-rich pancreatic juice and enhances the activity of secretin. Parasympathetic regulation occurs mainly during the cephalic and gastric phases of gastric secretion, when vagal stimulation prompts the secretion of pancreatic juice.

Usually, the pancreas secretes just enough bicarbonate to counterbalance the amount of HCl produced in the stomach. Hydrogen ions enter the blood when bicarbonate is secreted by the pancreas. Thus, the acidic blood draining from the pancreas neutralizes the alkaline blood draining from the stomach, maintaining the pH of the venous blood that flows to the liver.

The Gallbladder

The gallbladder is 8–10 cm (~3–4 in) long and is nested in a shallow area on the posterior aspect of the right lobe of the liver. This muscular sac stores, concentrates, and, when stimulated, propels the bile into the duodenum via the common bile duct. It is divided into three regions. The fundus is the widest portion and tapers medially into the body, which in turn narrows to become the neck. The neck angles slightly superiorly as it approaches the hepatic duct. The cystic duct is 1–2 cm (less than 1 in) long and turns inferiorly as it bridges the neck and hepatic duct.

The simple columnar epithelium of the gallbladder mucosa is organized in rugae, similar to those of the stomach. There is no submucosa in the gallbladder wall. The wall’s middle, muscular coat is made of smooth muscle fibers. When these fibers contract, the gallbladder’s contents are ejected through the cystic duct and into the bile duct (Figure 4). Visceral peritoneum reflected from the liver capsule holds the gallbladder against the liver and forms the outer coat of the gallbladder. The gallbladder's mucosa absorbs water and ions from bile, concentrating it by up to 10-fold.

[caption id="" align="aligncenter" width="380"]This figure shows the gallbladder and its major parts are labeled. Figure 4. Gallbladder. The gallbladder stores and concentrates bile, and releases it into the two-way cystic duct when it is needed by the small intestine.[/caption]
]]> 2703 2664 7 0 23.7 Chemical Digestion and Absorption: A Closer Look https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2710 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2710

Learning Objectives

By the end of this section, you will be able to:
  • Identify the locations and primary secretions involved in the chemical digestion of carbohydrates, proteins, lipids, and nucleic acids
  • Compare and contrast absorption of the hydrophilic and hydrophobic nutrients
As you have learned, the process of mechanical digestion is relatively simple. It involves the physical breakdown of food but does not alter its chemical makeup. Chemical digestion, on the other hand, is a complex process that reduces food into its chemical building blocks, which are then absorbed to nourish the cells of the body (Figure 1). In this section, you will look more closely at the processes of chemical digestion and absorption.
[caption id="" align="aligncenter" width="350"]This diagram identifies the functions of mechanical and chemical digestion and absorption at each organ. Next to each organ, a callout identifies which steps of digestion take place in that particular organ. Figure 1. Digestion and Absorption. Digestion begins in the mouth and continues as food travels through the small intestine. Most absorption occurs in the small intestine.[/caption]

Chemical Digestion

Large food molecules (for example, proteins, lipids, nucleic acids, and starches) must be broken down into subunits that are small enough to be absorbed by the lining of the alimentary canal. This is accomplished by enzymes through hydrolysis. The many enzymes involved in chemical digestion are summarized in Table 8.

*These enzymes have been activated by other substances.
The Digestive Enzymes (Table 8)
Enzyme Category Enzyme Name Source Substrate Product
Salivary Enzymes Lingual lipase Lingual glands Triglycerides Free fatty acids, and mono- and diglycerides
Salivary Enzymes Salivary amylase Salivary glands Polysaccharides Disaccharides and trisaccharides
Gastric enzymes Gastric lipase Chief cells Triglycerides Fatty acids and monoacylglycerides
Gastric enzymes Pepsin* Chief cells Proteins Peptides
Brush border enzymes α-Dextrinase Small intestine α-Dextrins Glucose
Brush border enzymes Enteropeptidase Small intestine Trypsinogen Trypsin
Brush border enzymes Lactase Small intestine Lactose Glucose and galactose
Brush border enzymes Maltase Small intestine Maltose Glucose
Brush border enzymes Nucleosidases and phosphatases Small intestine Nucleotides Phosphates, nitrogenous bases, and pentoses
Brush border enzymes Peptidases Small intestine
  • Aminopeptidase: amino acids at the amino end of peptides
  • Dipeptidase: dipeptides
  • Aminopeptidase: amino acids and peptides
  • Dipeptidase: amino acids
Brush border enzymes Sucrase Small intestine Sucrose Glucose and fructose
Pancreatic enzymes Carboxy-peptidase* Pancreatic acinar cells Amino acids at the carboxyl end of peptides Amino acids and peptides
Pancreatic enzymes Chymotrypsin* Pancreatic acinar cells Proteins Peptides
Pancreatic enzymes Elastase* Pancreatic acinar cells Proteins Peptides
Pancreatic enzymes Nucleases Pancreatic acinar cells
  • Ribonuclease: ribonucleic acids
  • Deoxyribonuclease: deoxyribonucleic acids
 Nucleotides
Pancreatic enzymes Pancreatic amylase Pancreatic acinar cells Polysaccharides (starches) α-Dextrins, disaccharides (maltose), trisaccharides (maltotriose)
Pancreatic enzymes Pancreatic lipase Pancreatic acinar cells Triglycerides that have been emulsified by bile salts Fatty acids and monoacylglycerides
Pancreatic enzymes Trypsin* Pancreatic acinar cells Proteins Peptides

Carbohydrate Digestion

The average American diet is about 50 percent carbohydrates, which may be classified according to the number of monomers they contain of simple sugars (monosaccharides and disaccharides) and/or complex sugars (polysaccharides). Glucose, galactose, and fructose are the three monosaccharides that are commonly consumed and are readily absorbed. Your digestive system is also able to break down the disaccharide sucrose (regular table sugar: glucose + fructose), lactose (milk sugar: glucose + galactose), and maltose (grain sugar: glucose + glucose), and the polysaccharides glycogen and starch (chains of monosaccharides). Your bodies do not produce enzymes that can break down most fibrous polysaccharides, such as cellulose. While indigestible polysaccharides do not provide any nutritional value, they do provide dietary fiber, which helps propel food through the alimentary canal.

The chemical digestion of starches begins in the mouth and has been reviewed above.

In the small intestine, pancreatic amylase does the ‘heavy lifting’ for starch and carbohydrate digestion (Figure 2). After amylases break down starch into smaller fragments, the brush border enzyme α-dextrinase starts working on α-dextrin, breaking off one glucose unit at a time. Three brush border enzymes hydrolyze sucrose, lactose, and maltose into monosaccharides. Sucrase splits sucrose into one molecule of fructose and one molecule of glucose; maltase breaks down maltose and maltotriose into two and three glucose molecules, respectively; and lactase breaks down lactose into one molecule of glucose and one molecule of galactose. Insufficient lactase can lead to lactose intolerance.

[caption id="" align="aligncenter" width="280"]This flow chart shows the steps in digestion of carbohydrates. The different levels shown are starch and glycogen, disaccharides and monosaccharides. Under each type of sugar, examples and the enzymes responsible for digestion are listed. Figure 2. Carbohydrate Digestion Flow Chart. Carbohydrates are broken down into their monomers in a series of steps.[/caption]

Protein Digestion

Proteins are polymers composed of amino acids linked by peptide bonds to form long chains. Digestion reduces them to their constituent amino acids. You usually consume about 15 to 20 percent of your total calorie intake as protein.

The digestion of protein starts in the stomach, where HCl and pepsin break proteins into smaller polypeptides, which then travel to the small intestine (Figure 3). Chemical digestion in the small intestine is continued by pancreatic enzymes, including chymotrypsin and trypsin, each of which act on specific bonds in amino acid sequences. At the same time, the cells of the brush border secrete enzymes such as aminopeptidase and dipeptidase, which further break down peptide chains. This results in molecules small enough to enter the bloodstream (Figure 4).
[caption id="" align="aligncenter" width="380"]This diagrams shows the human digestive system and identifies the role of each organ in protein digestion. A text call-out next to each organ details the specific function. Figure 3. Digestion of Protein. The digestion of protein begins in the stomach and is completed in the small intestine.[/caption]
[caption id="" align="aligncenter" width="180"]This flow chart shows the different steps in the digestion of protein. The four steps shown are protein, large polypeptides, short peptides and amino acids and amino acids. Figure 4. Digestion of Protein Flow Chart. Proteins are successively broken down into their amino acid components.[/caption]

Lipid Digestion

A healthy diet limits lipid intake to 35 percent of total calorie intake. The most common dietary lipids are triglycerides, which are made up of a glycerol molecule bound to three fatty acid chains. Small amounts of dietary cholesterol and phospholipids are also consumed.

The three lipases responsible for lipid digestion are lingual lipase, gastric lipase, and pancreatic lipase. However, because the pancreas is the only consequential source of lipase, virtually all lipid digestion occurs in the small intestine. Pancreatic lipase breaks down each triglyceride into two free fatty acids and a monoglyceride. The fatty acids include both short-chain (less than 10 to 12 carbons) and long-chain fatty acids.

Nucleic Acid Digestion

The nucleic acids DNA and RNA are found in most of the foods you eat. Two types of pancreatic nuclease are responsible for their digestion: deoxyribonuclease, which digests DNA, and ribonuclease, which digests RNA. The nucleotides produced by this digestion are further broken down by two intestinal brush border enzymes (nucleosidase and phosphatase) into pentoses, phosphates, and nitrogenous bases, which can be absorbed through the alimentary canal wall. The large food molecules that must be broken down into subunits are summarized Table 9
Absorbable Food Substances (Table 9)
Source Substance
Carbohydrates Monosaccharides: glucose, galactose, and fructose
Proteins Single amino acids, dipeptides, and tripeptides
Triglycerides Monoacylglycerides, glycerol, and free fatty acids
Nucleic acids Pentose sugars, phosphates, and nitrogenous bases

Absorption

The mechanical and digestive processes have one goal: to convert food into molecules small enough to be absorbed by the epithelial cells of the intestinal villi. The absorptive capacity of the alimentary canal is almost endless. Each day, the alimentary canal processes up to 10 liters of food, liquids, and GI secretions, yet less than one liter enters the large intestine. Almost all ingested food, 80 percent of electrolytes, and 90 percent of water are absorbed in the small intestine. Although the entire small intestine is involved in the absorption of water and lipids, most absorption of carbohydrates and proteins occurs in the jejunum. Notably, bile salts and vitamin B12 are absorbed in the terminal ileum. By the time chyme passes from the ileum into the large intestine, it is essentially indigestible food residue (mainly plant fibers like cellulose), some water, and millions of bacteria (Figure 5).
[caption id="" align="aligncenter" width="380"]This image shows the human digestive system. Next to each organ, a text callout identifies how water and digestive secretions such as saliva and bile are processed. Figure 5. Digestive Secretions and Absorption of Water. Absorption is a complex process, in which nutrients from digested food are harvested.[/caption]

Absorption can occur through five mechanisms: (1) active transport, (2) passive diffusion, (3) facilitated diffusion, (4) co-transport (or secondary active transport), and (5) endocytosis. As you will recall from Chapter 3, active transport refers to the movement of a substance across a cell membrane going from an area of lower concentration to an area of higher concentration (up the concentration gradient). In this type of transport, proteins within the cell membrane act as “pumps,” using cellular energy (ATP) to move the substance. Passive diffusion refers to the movement of substances from an area of higher concentration to an area of lower concentration, while facilitated diffusion refers to the movement of substances from an area of higher to an area of lower concentration using a carrier protein in the cell membrane. Co-transport uses the movement of one molecule through the membrane from higher to lower concentration to power the movement of another from lower to higher. Finally, endocytosis is a transportation process in which the cell membrane engulfs material. It requires energy, generally in the form of ATP.

Because the cell’s plasma membrane is made up of hydrophobic phospholipids, water-soluble nutrients must use transport molecules embedded in the membrane to enter cells. Moreover, substances cannot pass between the epithelial cells of the intestinal mucosa because these cells are bound together by tight junctions. Thus, substances can only enter blood capillaries by passing through the apical surfaces of epithelial cells and into the interstitial fluid. Water-soluble nutrients enter the capillary blood in the villi and travel to the liver via the hepatic portal vein.

In contrast to the water-soluble nutrients, lipid-soluble nutrients can diffuse through the plasma membrane. Once inside the cell, they are packaged for transport via the base of the cell and then enter the lacteals of the villi to be transported by lymphatic vessels to the systemic circulation via the thoracic duct. The absorption of most nutrients through the mucosa of the intestinal villi requires active transport fueled by ATP. The routes of absorption for each food category are summarized in Table 10.

Absorption in the Alimentary Canal (Table 10)
Food Breakdown products Absorption mechanism Entry to bloodstream Destination
Carbohydrates Glucose Co-transport with sodium ions Capillary blood in villi Liver via hepatic portal vein
Carbohydrates Galactose Co-transport with sodium ions Capillary blood in villi Liver via hepatic portal vein
Carbohydrates Fructose Facilitated diffusion Capillary blood in villi Liver via hepatic portal vein
Protein Amino acids Co-transport with sodium ions Capillary blood in villi Liver via hepatic portal vein
Lipids Long-chain fatty acids Diffusion into intestinal cells, where they are combined with proteins to create chylomicrons Lacteals of villi Systemic circulation via lymph entering thoracic duct
Lipids Monoacylglycerides Diffusion into intestinal cells, where they are combined with proteins to create chylomicrons Lacteals of villi Systemic circulation via lymph entering thoracic duct
Lipids Short-chain fatty acids Simple diffusion Capillary blood in villi Liver via hepatic portal vein
Lipids Glycerol Simple diffusion Capillary blood in villi Liver via hepatic portal vein
Lipids Nucleic acid digestion products Active transport via membrane carriers Capillary blood in villi Liver via hepatic portal vein

Carbohydrate Absorption

All carbohydrates are absorbed in the form of monosaccharides. The small intestine is highly efficient at this, absorbing monosaccharides at an estimated rate of 120 grams per hour. All normally digested dietary carbohydrates are absorbed; indigestible fibers are eliminated in the feces. The monosaccharides glucose and galactose are transported into the epithelial cells by common protein carriers via secondary active transport (that is, co-transport with sodium ions). The monosaccharides leave these cells via facilitated diffusion and enter the capillaries through intercellular clefts. The monosaccharide fructose (which is in fruit) is absorbed and transported by facilitated diffusion alone. The monosaccharides combine with the transport proteins immediately after the disaccharides are broken down.

Protein Absorption

Active transport mechanisms, primarily in the duodenum and jejunum, absorb most proteins as their breakdown products, amino acids. Almost all (95 to 98 percent) protein is digested and absorbed in the small intestine. The type of carrier that transports an amino acid varies. Most carriers are linked to the active transport of sodium. Short chains of two amino acids (dipeptides) or three amino acids (tripeptides) are also transported actively. However, after they enter the absorptive epithelial cells, they are broken down into their amino acids before leaving the cell and entering the capillary blood via diffusion.

Lipid Absorption

About 95 percent of lipids are absorbed in the small intestine. Bile salts not only speed up lipid digestion, they are also essential to the absorption of the end products of lipid digestion. Short-chain fatty acids are relatively water soluble and can enter the absorptive cells (enterocytes) directly. Despite being hydrophobic, the small size of short-chain fatty acids enables them to be absorbed by enterocytes via simple diffusion, and then take the same path as monosaccharides and amino acids into the blood capillary of a villus. The large and hydrophobic long-chain fatty acids and monoacylglycerides are not so easily suspended in the watery intestinal chyme. However, bile salts and lecithin resolve this issue by enclosing them in a micelle, which is a tiny sphere with polar (hydrophilic) ends facing the watery environment and hydrophobic tails turned to the interior, creating a receptive environment for the long-chain fatty acids. The core also includes cholesterol and fat-soluble vitamins. Without micelles, lipids would sit on the surface of chyme and never come in contact with the absorptive surfaces of the epithelial cells. Micelles can easily squeeze between microvilli and get very near the luminal cell surface. At this point, lipid substances exit the micelle and are absorbed via simple diffusion. The free fatty acids and monoacylglycerides that enter the epithelial cells are reincorporated into triglycerides. The triglycerides are mixed with phospholipids and cholesterol, and surrounded with a protein coat. This new complex, called a chylomicron, is a water-soluble lipoprotein. After being processed by the Golgi apparatus, chylomicrons are released from the cell (Figure 6). Too big to pass through the basement membranes of blood capillaries, chylomicrons instead enter the large pores of lacteals. The lacteals come together to form the lymphatic vessels. The chylomicrons are transported in the lymphatic vessels and empty through the thoracic duct into the subclavian vein of the circulatory system. Once in the bloodstream, the enzyme lipoprotein lipase breaks down the triglycerides of the chylomicrons into free fatty acids and glycerol. These breakdown products then pass through capillary walls to be used for energy by cells or stored in adipose tissue as fat. Liver cells combine the remaining chylomicron remnants with proteins, forming lipoproteins that transport cholesterol in the blood.
[caption id="" align="aligncenter" width="320"]This diagram shows how lipids are absorbed from the lumen of the intestine into the lacteals. The fatty acid micelles are shown to enter the epithelial cell and form chylomicrons inside the Golgi apparatus. Then, the chylomicrons are extruded from the epithelial cell and are taken up by the lacteals. Figure 6. Lipid Absorption. Unlike amino acids and simple sugars, lipids are transformed as they are absorbed through epithelial cells.[/caption]

Nucleic Acid Absorption

The products of nucleic acid digestion—pentose sugars, nitrogenous bases, and phosphate ions—are transported by carriers across the villus epithelium via active transport. These products then enter the bloodstream.

Mineral Absorption

The electrolytes absorbed by the small intestine are from both GI secretions and ingested foods. Since electrolytes dissociate into ions in water, most are absorbed via active transport throughout the entire small intestine. During absorption, co-transport mechanisms result in the accumulation of sodium ions inside the cells, whereas anti-port mechanisms reduce the potassium ion concentration inside the cells. To restore the sodium-potassium gradient across the cell membrane, a sodium-potassium pump requiring ATP pumps sodium out and potassium in.

In general, all minerals that enter the intestine are absorbed, whether you need them or not. Iron and calcium are exceptions; they are absorbed in the duodenum in amounts that meet the body’s current requirements, as follows: Iron—The ionic iron needed for the production of hemoglobin is absorbed into mucosal cells via active transport. Once inside mucosal cells, ionic iron binds to the protein ferritin, creating iron-ferritin complexes that store iron until needed. When the body has enough iron, most of the stored iron is lost when worn-out epithelial cells slough off. When the body needs iron because, for example, it is lost during acute or chronic bleeding, there is increased uptake of iron from the intestine and accelerated release of iron into the bloodstream. Since women experience significant iron loss during menstruation, they have around four times as many iron transport proteins in their intestinal epithelial cells as do men.

Calcium—Blood levels of ionic calcium determine the absorption of dietary calcium. When blood levels of ionic calcium drop, parathyroid hormone (PTH) secreted by the parathyroid glands stimulates the release of calcium ions from bone matrices and increases the reabsorption of calcium by the kidneys. PTH also upregulates the activation of vitamin D in the kidney, which then facilitates intestinal calcium ion absorption.

Vitamin Absorption

The small intestine absorbs the vitamins that occur naturally in food and supplements. Fat-soluble vitamins (A, D, E, and K) are absorbed along with dietary lipids in micelles via simple diffusion. This is why you are advised to eat some fatty foods when you take fat-soluble vitamin supplements. Most water-soluble vitamins (including most B vitamins and vitamin C) also are absorbed by simple diffusion. An exception is vitamin B12, which is a very large molecule. Intrinsic factor secreted in the stomach binds to vitamin B12, preventing its digestion and creating a complex that binds to mucosal receptors in the terminal ileum, where it is taken up by endocytosis.

Water Absorption

Each day, about nine liters of fluid enter the small intestine. About 2.3 liters are ingested in foods and beverages, and the rest is from GI secretions. About 90 percent of this water is absorbed in the small intestine. Water absorption is driven by the concentration gradient of the water: The concentration of water is higher in chyme than it is in epithelial cells. Thus, water moves down its concentration gradient from the chyme into cells. As noted earlier, much of the remaining water is then absorbed in the colon.
]]> 2710 2664 8 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/chapter/introduction/ Thu, 13 Jul 2017 23:38:16 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1814 This photo shows a group of people climbing a mountain. Figure 1. Mountain Climbers. The thin air at high elevations can strain the human respiratory system. (credit: “bortescristian”/flickr.com)[/caption]

Chapter Objectives

After studying this chapter, you will be able to:
  • List the structures of the respiratory system
  • List the major functions of the respiratory system
  • Outline the forces that allow for air movement into and out of the lungs
  • Outline the process of gas exchange
  • Summarize the process of oxygen and carbon dioxide transport within the respiratory system
  • Create a flow chart illustrating how respiration is controlled
  • Discuss how the respiratory system responds to exercise
  • Describe the development of the respiratory system in the embryo
Hold your breath. Really! See how long you can hold your breath as you continue reading…How long can you do it? Chances are you are feeling uncomfortable already. A typical human cannot survive without breathing for more than 3 minutes, and even if you wanted to hold your breath longer, your autonomic nervous system would take control. This is because every cell in the body needs to run the oxidative stages of cellular respiration, the process by which energy is produced in the form of adenosine triphosphate (ATP). For oxidative phosphorylation to occur, oxygen is used as a reactant and carbon dioxide is released as a waste product. You may be surprised to learn that although oxygen is a critical need for cells, it is actually the accumulation of carbon dioxide that primarily drives your need to breathe. Carbon dioxide is exhaled and oxygen is inhaled through the respiratory system, which includes muscles to move air into and out of the lungs, passageways through which air moves, and microscopic gas exchange surfaces covered by capillaries. The circulatory system transports gases from the lungs to tissues throughout the body and vice versa. A variety of diseases can affect the respiratory system, such as asthma, emphysema, chronic obstruction pulmonary disorder (COPD), and lung cancer. All of these conditions affect the gas exchange process and result in labored breathing and other difficulties.]]> 1814 2352 1 0 3.2 The Cytoplasm and Cellular Organelles https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1955 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1955

By the end of this section, you will be able to:
  • Describe the structure and function of the cellular organelles associated with the endomembrane system, including the endoplasmic reticulum, Golgi apparatus, and lysosomes
  • Describe the structure and function of mitochondria and peroxisomes
  • Explain the three components of the cytoskeleton, including their composition and functions
[caption id="" align="alignleft" width="450"]This diagram shows an animal cell with all the intracellular organelles labeled. Figure 1. Prototypical Human Cell. While this image is not indicative of any one particular human cell, it is a prototypical example of a cell containing the primary organelles and internal structures.[/caption]

Now that you have learned that the cell membrane surrounds all cells, you can dive inside of a prototypical human cell to learn about its internal components and their functions. All living cells in multicellular organisms contain an internal cytoplasmic compartment, and a nucleus within the cytoplasm. Cytosol, the jelly-like substance within the cell, provides the fluid medium necessary for biochemical reactions. Eukaryotic cells, including all animal cells, also contain various cellular organelles. An organelle (“little organ”) is one of several different types of membrane-enclosed bodies in the cell, each performing a unique function. Just as the various bodily organs work together in harmony to perform all of a human’s functions, the many different cellular organelles work together to keep the cell healthy and performing all of its important functions. The organelles and cytosol, taken together, compose the cell’s cytoplasm. The nucleus is a cell’s central organelle, which contains the cell’s DNA (Figure 1).

Organelles of the Endomembrane System

A set of three major organelles together form a system within the cell called the endomembrane system. These organelles work together to perform various cellular jobs, including the task of producing, packaging, and exporting certain cellular products. The organelles of the endomembrane system include the endoplasmic reticulum, Golgi apparatus, and vesicles.

Endoplasmic Reticulum

The endoplasmic reticulum (ER) is a system of channels that is continuous with the nuclear membrane (or “envelope”) covering the nucleus and composed of the same lipid bilayer material. The ER can be thought of as a series of winding thoroughfares similar to the waterway canals in Venice. The ER provides passages throughout much of the cell that function in transporting, synthesizing, and storing materials. The winding structure of the ER results in a large membranous surface area that supports its many functions (Figure 2).

[caption id="" align="alignleft" width="550"]This figure shows structure of the endoplasmic reticulum. The diagram highlights the rough and smooth endoplasmic reticulum and the nucleus is labeled. Two micrographs show the structure of the endoplasmic reticulum in detail. The left micrograph shows the rough endoplasmic reticulum in a pancreatic cell and the right micrograph shows a smooth endoplasmic reticulum. Figure 2. Endoplasmic Reticulum (ER). (a) The ER is a winding network of thin membranous sacs found in close association with the cell nucleus. The smooth and rough endoplasmic reticula are very different in appearance and function (source: mouse tissue). (b) Rough ER is studded with numerous ribosomes, which are sites of protein synthesis (source: mouse tissue). EM × 110,000. (c) Smooth ER synthesizes phospholipids, steroid hormones, regulates the concentration of cellular Ca++,metabolizes some carbohydrates, and breaks down certain toxins (source: mouse tissue). EM × 110,510. (Micrographs provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Endoplasmic reticulum can exist in two forms: rough ER and smooth ER. These two types of ER perform some very different functions and can be found in very different amounts depending on the type of cell. Rough ER (RER) is so-called because its membrane is dotted with embedded granules—organelles called ribosomes, giving the RER a bumpy appearance. A ribosome is an organelle that serves as the site of protein synthesis. It is composed of two ribosomal RNA subunits that wrap around mRNA to start the process of translation, followed by protein synthesis. Smooth ER (SER) lacks these ribosomes.

One of the main functions of the smooth ER is in the synthesis of lipids. The smooth ER synthesizes phospholipids, the main component of biological membranes, as well as steroid hormones. For this reason, cells that produce large quantities of such hormones, such as those of the female ovaries and male testes, contain large amounts of smooth ER. In addition to lipid synthesis, the smooth ER also sequesters (i.e., stores) and regulates the concentration of cellular Ca++, a function extremely important in cells of the nervous system where Ca++ is the trigger for neurotransmitter release. The smooth ER additionally metabolizes some carbohydrates and performs a detoxification role, breaking down certain toxins.

In contrast with the smooth ER, the primary job of the rough ER is the synthesis and modification of proteins destined for the cell membrane or for export from the cell. For this protein synthesis, many ribosomes attach to the ER (giving it the studded appearance of rough ER). Typically, a protein is synthesized within the ribosome and released inside the channel of the rough ER, where sugars can be added to it (by a process called glycosylation) before it is transported within a vesicle to the next stage in the packaging and shipping process: the Golgi apparatus.

The Golgi Apparatus

The Golgi apparatus is responsible for sorting, modifying, and shipping off the products that come from the rough ER, much like a post-office. The Golgi apparatus looks like stacked flattened discs, almost like stacks of oddly shaped pancakes. Like the ER, these discs are membranous. The Golgi apparatus has two distinct sides, each with a different role. One side of the apparatus receives products in vesicles. These products are sorted through the apparatus, and then they are released from the opposite side after being repackaged into new vesicles. If the product is to be exported from the cell, the vesicle migrates to the cell surface and fuses to the cell membrane, and the cargo is secreted (Figure 3). [caption id="" align="alignright" width="550"]This figure shows the structure of the Golgi apparatus. The diagram in the left panel shows the location and structure of the Golgi apparatus. The right panel shows a micrograph showing the folds of the Golgi in detail. Figure 3. Golgi Apparatus. (a) The Golgi apparatus manipulates products from the rough ER, and also produces new organelles called lysosomes. Proteins and other products of the ER are sent to the Golgi apparatus, which organizes, modifies, packages, and tags them. Some of these products are transported to other areas of the cell and some are exported from the cell through exocytosis. Enzymatic proteins are packaged as new lysosomes (or packaged and sent for fusion with existing lysosomes). (b) An electron micrograph of the Golgi apparatus.[/caption]

Lysosomes

Some of the protein products packaged by the Golgi include digestive enzymes that are meant to remain inside the cell for use in breaking down certain materials. The enzyme-containing vesicles released by the Golgi may form new lysosomes, or fuse with existing, lysosomes. A lysosome is an organelle that contains enzymes that break down and digest unneeded cellular components, such as a damaged organelle. (A lysosome is similar to a wrecking crew that takes down old and unsound buildings in a neighborhood.) Autophagy (“self-eating”) is the process of a cell digesting its own structures. Lysosomes are also important for breaking down foreign material. For example, when certain immune defense cells (white blood cells) phagocytize bacteria, the bacterial cell is transported into a lysosome and digested by the enzymes inside. As one might imagine, such phagocytic defense cells contain large numbers of lysosomes.

Under certain circumstances, lysosomes perform a more grand and dire function. In the case of damaged or unhealthy cells, lysosomes can be triggered to open up and release their digestive enzymes into the cytoplasm of the cell, killing the cell. This “self-destruct” mechanism is called autolysis, and makes the process of cell death controlled (a mechanism called “apoptosis”).

Watch this Khan Academy video to learn more about the endomembrane system

Organelles for Energy Production and Detoxification

In addition to the jobs performed by the endomembrane system, the cell has many other important functions. Just as you must consume nutrients to provide yourself with energy, so must each of your cells take in nutrients, some of which convert to chemical energy that can be used to power biochemical reactions. Another important function of the cell is detoxification. Humans take in all sorts of toxins from the environment and also produce harmful chemicals as byproducts of cellular processes. Cells called hepatocytes in the liver detoxify many of these toxins.

Mitochondria

[caption id="" align="alignright" width="520"]This figure shows the structure of a mitochondrion. The inner and outer membrane, the cristae and the intermembrane space are labeled. The right panel shows a micrograph with the structure of a mitochondrion in detail. Figure 4. Mitochondrion. The mitochondria are the energy-conversion factories of the cell. (a) A mitochondrion is composed of two separate lipid bilayer membranes. Along the inner membrane are various molecules that work together to produce ATP, the cell’s major energy currency. (b) An electron micrograph of mitochondria. EM × 236,000. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

A mitochondrion (plural = mitochondria) is a membranous, bean-shaped organelle that is the “energy transformer” of the cell. Mitochondria consist of an outer lipid bilayer membrane as well as an additional inner lipid bilayer membrane (Figure 4). The inner membrane is highly folded into winding structures with a great deal of surface area, called cristae. It is along this inner membrane that a series of proteins, enzymes, and other molecules perform the biochemical reactions of cellular respiration. These reactions convert energy stored in nutrient molecules (such as glucose) into adenosine triphosphate (ATP), which provides usable cellular energy to the cell. Cells use ATP constantly, and so the mitochondria are constantly at work. Oxygen molecules are required during cellular respiration, which is why you must constantly breathe it in. One of the organ systems in the body that uses huge amounts of ATP is the muscular system because ATP is required to sustain muscle contraction. As a result, muscle cells are packed full of mitochondria. Nerve cells also need large quantities of ATP to run their sodium-potassium pumps. Therefore, an individual neuron will be loaded with over a thousand mitochondria. On the other hand, a bone cell, which is not nearly as metabolically-active, might only have a couple hundred mitochondria.

]]> 1955 1936 3 0 4.1 Types of Tissues https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1983 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1983

By the end of this section, you will be able to:
  • Identify the four main tissue types
  • Discuss the functions of each tissue type
  • Relate the structure of each tissue type to their function
  • Discuss the embryonic origin of tissue
  • Identify the three major germ layers
  • Identify the main types of tissue membranes

The term tissue is used to describe a group of cells found together in the body. The cells within a tissue share a common embryonic origin. Microscopic observation reveals that the cells in a tissue share morphological features and are arranged in an orderly pattern that achieves the tissue’s functions. From the evolutionary perspective, tissues appear in more complex organisms. For example, multicellular protists, ancient eukaryotes, do not have cells organized into tissues.

Although there are many types of cells in the human body, they are organized into four broad categories of tissues: epithelial, connective, muscle, and nervous. Each of these categories is characterized by specific functions that contribute to the overall health and maintenance of the body. A disruption of the structure is a sign of injury or disease. Such changes can be detected through histology, the microscopic study of tissue appearance, organization, and function.

The Four Types of Tissues

[caption id="" align="alignleft" width="500"]This diagram shows the silhouette of a female surrounded by four micrographs of tissue. Each micrograph has arrows pointing to the organs where that tissue is found. The upper left micrograph shows nervous tissue that is whitish with several large, purple, irregularly-shaped neurons embedded throughout. Nervous tissue is found in the brain, spinal cord and nerves. The upper right micrograph shows muscle tissue that is red with elongated cells and prominent, purple nuclei. Cardiac muscle is found in the heart. Smooth muscle is found in muscular internal organs, such as the stomach. Skeletal muscle is found in parts that are moved voluntarily, such as the arms. The lower left micrograph shows epithelial tissue. This tissue is purple with many round, purple cells with dark purple nuclei. Epithelial tissue is found in the lining of GI tract organs and other hollow organs such as the small intestine. Epithelial tissue also composes the outer layer of the skin, known as the epidermis. Finally, the lower right micrograph shows connective tissue, which is composed of very loosely packed purple cells and fibers. There are large open spaces between clumps of cells and fibers. Connective tissue is found in the leg within fat and other soft padding tissue as well as bones and tendons. Figure 1. Four Types of Tissue: Body. The four types of tissues are exemplified in nervous tissue, stratified squamous epithelial tissue, cardiac muscle tissue, and connective tissue in small intestine. Clockwise from nervous tissue, LM × 872, LM × 282, LM × 460, LM × 800. (Micrographs provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Epithelial tissue, also referred to as epithelium, refers to the sheets of cells that cover exterior surfaces of the body, lines internal cavities and passageways, and forms certain glands. Connective tissue, as its name implies, binds the cells and organs of the body together and functions in the protection, support, and integration of all parts of the body. Muscle tissue is excitable, responding to stimulation and contracting to provide movement, and occurs as three major types: skeletal (voluntary) muscle, smooth muscle, and cardiac muscle in the heart. Nervous tissue is also excitable, allowing the propagation of electrochemical signals in the form of nerve impulses that communicate between different regions of the body (Figure 1).

The next level of organization is the organ, where several types of tissues come together to form a working unit. Just as knowing the structure and function of cells helps you in your study of tissues, knowledge of tissues will help you understand how organs function. The epithelial and connective tissues are discussed in detail in this chapter. Muscle and nervous tissues will be discussed only briefly in this chapter.

Embryonic Origin of Tissues

[caption id="" align="alignright" width="520"]This is a two column-table containing both text and illustrations. The left column is titled germ layer while the right column is titled “Gives rise to.” The germ layer in the first row is ectoderm. Ectoderm gives rise to epidermis, glands on the skin, some cranial bones, the pituitary and adrenal medulla, the nervous system, the tissue between the cheeks and gums, and the anus. This row contains three pictures. The leftmost picture illustrates several layers of yellow, oval-shaped skin cells with purple nuclei. The middle diagram shows a neuron, which is a yellow, star shaped cell with finger like branches at its corners. The neuron also has a purple nucleus and a yellow tube that connects to the bottom of the cell. The right image in this row shows a brown pigment cell embedded at the bottom layer of several skin cells. It is secreting dark-colored pigment into the skin cells from tentacle-like projections. The germ layer in the second row is mesoderm. Mesoderm gives rise to connective tissues, bone, cartilage, blood, the endothelium of blood vessels, muscle, synovial membranes, serous membranes that line body cavities, the kidneys, and the lining of the gonads. Five images are given in this row to illustrate. The leftmost image is cardiac muscle, which is cylindrical and curved. There are many open spaces between neighboring cardiac muscles. The next image shows skeletal muscle, which is a series of closely stacked cylinders with well defined horizontal striping. The middle image shows three tubule cells of the kidney, which are square shaped and contain a brown nucleus. The fourth image shows a series of red blood cells, which are red and saucer shaped with a slight depression at the center. The fifth image shows smooth muscles which are tightly packed, diamond shaped cells with oval-shaped nuclei. Endoderm gives rise to the lining of the airways and digestive system (except the mouth and distal part of digestive system). Also, the rectum and anal canal, digestive glands, endocrine glands, and adrenal cortex all develop from endoderm. The leftmost image in this row shows a lung cell, which is a large, purple, trapezoid-shaped cell. The middle image shows a pair of thyroid cells, which are rectangle-shaped with the upper edge of each cell having a row of finger like projections, similar in appearance to carpet. The rightmost image in this row shows a pancreatic cell, which is large and wedge-shaped. The pancreatic cell has small indentations throughout its cell membrane. Figure 2. Embryonic Origin of Tissues and Major Organs[/caption]

The zygote, or fertilized egg, is a single cell formed by the fusion of an egg and sperm. After fertilization the zygote gives rise to rapid mitotic cycles, generating many cells to form the embryo. The first embryonic cells generated have the ability to differentiate into any type of cell in the body and, as such, are called totipotent, meaning each has the capacity to divide, differentiate, and develop into a new organism. As cell proliferation progresses, three major cell lineages are established within the embryo. Each of these lineages of embryonic cells forms the distinct germ layers from which all the tissues and organs of the human body eventually form. Each germ layer is identified by its relative position: ectoderm (ecto- = “outer”), mesoderm (meso- = “middle”), and endoderm (endo- = “inner”). Figure 2 shows the types of tissues and organs associated with the each of the three germ layers. Note that epithelial tissue originates in all three layers, whereas nervous tissue derives primarily from the ectoderm and muscle tissue from mesoderm.

Tissue Membranes

[caption id="" align="alignright" width="420"]This illustrations shows the silhouette of a human female from an anterior view. Several organs are showing in her neck, thorax, abdomen left arm and right leg. Text boxes point out and describe the mucous membranes in several different organs. The topmost box points to the mouth and trachea. It states that mucous membranes line the digestive, respiratory, urinary and reproductive tracts. They are coated with the secretions of mucous glands. The second box points to the outside edge of the lungs as well as the large intestine and states that serous membranes line body cavities that are closed to the exterior of the body, including the peritoneal, pleural and pericardial cavities. The third box points to the skin of the hand. It states that cutaneous membrane, also known as the skin, covers the body surface. The fourth box points to the right knee. It states that synovial membranes line joint cavities and produce the fluid within the joint. Figure 3. Tissue Membranes. The two broad categories of tissue membranes in the body are (1) connective tissue membranes, which include synovial membranes, and (2) epithelial membranes, which include mucous membranes, serous membranes, and the cutaneous membrane, in other words, the skin.[/caption]

A tissue membrane is a thin layer or sheet of cells that covers the outside of the body (for example, skin), the organs (for example, pericardium), internal passageways that lead to the exterior of the body (for example, abdominal mesenteries), and the lining of the moveable joint cavities. There are two basic types of tissue membranes: connective tissue and epithelial membranes (Figure 3).

Connective Tissue Membranes

The connective tissue membrane is formed solely from connective tissue. These membranes encapsulate organs, such as the kidneys, and line our movable joints. A synovial membrane is a type of connective tissue membrane that lines the cavity of a freely movable joint. For example, synovial membranes surround the joints of the shoulder, elbow, and knee. Fibroblasts in the inner layer of the synovial membrane release hyaluronan into the joint cavity. The hyaluronan effectively traps available water to form the synovial fluid, a natural lubricant that enables the bones of a joint to move freely against one another without much friction. This synovial fluid readily exchanges water and nutrients with blood, as do all body fluids.

Epithelial Membranes

The epithelial membrane is composed of epithelium attached to a layer of connective tissue, for example, your skin. The mucous membrane is also a composite of connective and epithelial tissues. Sometimes called mucosae, these epithelial membranes line the body cavities and hollow passageways that open to the external environment, and include the digestive, respiratory, excretory, and reproductive tracts. Mucous, produced by the epithelial exocrine glands, covers the epithelial layer. The underlying connective tissue, called the lamina propria (literally “own layer”), help support the fragile epithelial layer.

A serous membrane is an epithelial membrane composed of mesodermally derived epithelium called the mesothelium that is supported by connective tissue. These membranes line the coelomic cavities of the body, that is, those cavities that do not open to the outside, and they cover the organs located within those cavities. They are essentially membranous bags, with mesothelium lining the inside and connective tissue on the outside. Serous fluid secreted by the cells of the thin squamous mesothelium lubricates the membrane and reduces abrasion and friction between organs. Serous membranes are identified according locations. Three serous membranes line the thoracic cavity; the two pleura that cover the lungs and the pericardium that covers the heart. A fourth, the peritoneum, is the serous membrane in the abdominal cavity that covers abdominal organs and forms double sheets of mesenteries that suspend many of the digestive organs.

The skin is an epithelial membrane also called the cutaneous membrane. It is a stratified squamous epithelial membrane resting on top of connective tissue. The apical surface of this membrane is exposed to the external environment and is covered with dead, keratinized cells that help protect the body from desiccation and pathogens.

]]> 1983 1977 2 0 22.1 Organs and Structures of the Respiratory System https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-1-organs-and-structures-of-the-respiratory-system/ Thu, 13 Jul 2017 23:02:01 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2373

Learning Objectives

By the end of this section, you will be able to:
  • List the structures that make up the respiratory system
  • Describe how the respiratory system processes oxygen and CO2
  • Compare and contrast the functions of upper respiratory tract with the lower respiratory tract

The major organs of the respiratory system function primarily to provide oxygen to body tissues for cellular respiration, remove the waste product carbon dioxide, and help to maintain acid-base balance. Portions of the respiratory system are also used for non-vital functions, such as sensing odors, speech production, and for straining, such as during childbirth or coughing (Figure 1).

[caption id="" align="aligncenter" width="420"]This figure shows the upper half of the human body. The major organs in the respiratory system are labeled. Figure 1. Major Respiratory Structures. The major respiratory structures span the nasal cavity to the diaphragm.[/caption]

Functionally, the respiratory system can be divided into a conducting zone and a respiratory zone. The conducting zone of the respiratory system includes the organs and structures not directly involved in gas exchange. The gas exchange occurs in the respiratory zone.

Conducting Zone

The major functions of the conducting zone are to provide a route for incoming and outgoing air, remove debris and pathogens from the incoming air, and warm and humidify the incoming air. Several structures within the conducting zone perform other functions as well. The epithelium of the nasal passages, for example, is essential to sensing odors, and the bronchial epithelium that lines the lungs can metabolize some airborne carcinogens.

The Nose and its Adjacent Structures

The major entrance and exit for the respiratory system is through the nose. When discussing the nose, it is helpful to divide it into two major sections: the external nose, and the nasal cavity or internal nose.

The external nose consists of the surface and skeletal structures that result in the outward appearance of the nose and contribute to its numerous functions (Figure 2). The root is the region of the nose located between the eyebrows. The bridge is the part of the nose that connects the root to the rest of the nose. The dorsum nasi is the length of the nose. The apex is the tip of the nose. On either side of the apex, the nostrils are formed by the alae (singular = ala). An ala is a cartilaginous structure that forms the lateral side of each naris (plural = nares), or nostril opening. The philtrum is the concave surface that connects the apex of the nose to the upper lip.

[caption id="" align="aligncenter" width="450"]This figure shows the human nose. The top left panel shows the front view, and the top right panel shows the side view. The bottom panel shows the cartilaginous components of the nose. Figure 2. Nose. This illustration shows features of the external nose (top) and skeletal features of the nose (bottom).[/caption]

Underneath the thin skin of the nose are its skeletal features (see Figure 2, lower illustration). While the root and bridge of the nose consist of bone, the protruding portion of the nose is composed of cartilage. As a result, when looking at a skull, the nose is missing. The nasal bone is one of a pair of bones that lies under the root and bridge of the nose. The nasal bone articulates superiorly with the frontal bone and laterally with the maxillary bones. Septal cartilage is flexible hyaline cartilage connected to the nasal bone, forming the dorsum nasi. The alar cartilage consists of the apex of the nose; it surrounds the naris.

The nares open into the nasal cavity, which is separated into left and right sections by the nasal septum (Figure 3). The nasal septum is formed anteriorly by a portion of the septal cartilage (the flexible portion you can touch with your fingers) and posteriorly by the perpendicular plate of the ethmoid bone (a cranial bone located just posterior to the nasal bones) and the thin vomer bones (whose name refers to its plough shape). Each lateral wall of the nasal cavity has three bony projections, called the superior, middle, and inferior nasal conchae. The inferior conchae are separate bones, whereas the superior and middle conchae are portions of the ethmoid bone. Conchae serve to increase the surface area of the nasal cavity and to disrupt the flow of air as it enters the nose, causing air to bounce along the epithelium, where it is cleaned and warmed. The conchae and meatuses also conserve water and prevent dehydration of the nasal epithelium by trapping water during exhalation. The floor of the nasal cavity is composed of the palate. The hard palate at the anterior region of the nasal cavity is composed of bone. The soft palate at the posterior portion of the nasal cavity consists of muscle tissue. Air exits the nasal cavities via the internal nares and moves into the pharynx.

[caption id="" align="aligncenter" width="450"]This figure shows a cross section view of the nose and throat. The major parts are labeled. Figure 3. Upper Airway[/caption]

Several bones that help form the walls of the nasal cavity have air-containing spaces called the paranasal sinuses, which serve to warm and humidify incoming air. Sinuses are lined with a mucosa. Each paranasal sinus is named for its associated bone: frontal sinus, maxillary sinus, sphenoidal sinus, and ethmoidal sinus. The sinuses produce mucus and lighten the weight of the skull.

The nares and anterior portion of the nasal cavities are lined with mucous membranes, containing sebaceous glands and hair follicles that serve to prevent the passage of large debris, such as dirt, through the nasal cavity. An olfactory epithelium used to detect odors is found deeper in the nasal cavity.

The conchae, meatuses, and paranasal sinuses are lined by respiratory epithelium composed of pseudostratified ciliated columnar epithelium (Figure 4). The epithelium contains goblet cells, one of the specialized, columnar epithelial cells that produce mucus to trap debris. The cilia of the respiratory epithelium help remove the mucus and debris from the nasal cavity with a constant beating motion, sweeping materials towards the throat to be swallowed. Interestingly, cold air slows the movement of the cilia, resulting in accumulation of mucus that may in turn lead to a runny nose during cold weather. This moist epithelium functions to warm and humidify incoming air. Capillaries located just beneath the nasal epithelium warm the air by convection. Serous and mucus-producing cells also secrete the lysozyme enzyme and proteins called defensins, which have antibacterial properties. Immune cells that patrol the connective tissue deep to the respiratory epithelium provide additional protection.

[caption id="" align="aligncenter" width="500"]This figure shows a micrograph of pseudostratified epithelium. Figure 4. Pseudostratified Ciliated Columnar Epithelium. Respiratory epithelium is pseudostratified ciliated columnar epithelium. Seromucous glands provide lubricating mucus. LM × 680. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Pharynx

The pharynx is a tube formed by skeletal muscle and lined by mucous membrane that is continuous with that of the nasal cavities (see Figure 3). The pharynx is divided into three major regions: the nasopharynx, the oropharynx, and the laryngopharynx (Figure 5).

[caption id="" align="aligncenter" width="475"]This figure shows the side view of the face. The different parts of the pharynx are color-coded and labeled. Figure 5. Divisions of the Pharynx. The pharynx is divided into three regions: the nasopharynx, the oropharynx, and the laryngopharynx.[/caption]

The nasopharynx is flanked by the conchae of the nasal cavity, and it serves only as an airway. At the top of the nasopharynx are the pharyngeal tonsils. A pharyngeal tonsil, also called an adenoid, is an aggregate of lymphoid reticular tissue similar to a lymph node that lies at the superior portion of the nasopharynx. The function of the pharyngeal tonsil is not well understood, but it contains a rich supply of lymphocytes and is covered with ciliated epithelium that traps and destroys invading pathogens that enter during inhalation. The pharyngeal tonsils are large in children, but interestingly, tend to regress with age and may even disappear. The uvula is a small bulbous, teardrop-shaped structure located at the apex of the soft palate. Both the uvula and soft palate move like a pendulum during swallowing, swinging upward to close off the nasopharynx to prevent ingested materials from entering the nasal cavity. In addition, auditory (Eustachian) tubes that connect to each middle ear cavity open into the nasopharynx. This connection is why colds often lead to ear infections.

The oropharynx is a passageway for both air and food. The oropharynx is bordered superiorly by the nasopharynx and anteriorly by the oral cavity. The fauces is the opening at the connection between the oral cavity and the oropharynx. As the nasopharynx becomes the oropharynx, the epithelium changes from pseudostratified ciliated columnar epithelium to stratified squamous epithelium. The oropharynx contains two distinct sets of tonsils, the palatine and lingual tonsils. A palatine tonsil is one of a pair of structures located laterally in the oropharynx in the area of the fauces. The lingual tonsil is located at the base of the tongue. Similar to the pharyngeal tonsil, the palatine and lingual tonsils are composed of lymphoid tissue, and trap and destroy pathogens entering the body through the oral or nasal cavities.

The laryngopharynx is inferior to the oropharynx and posterior to the larynx. It continues the route for ingested material and air until its inferior end, where the digestive and respiratory systems diverge. The stratified squamous epithelium of the oropharynx is continuous with the laryngopharynx. Anteriorly, the laryngopharynx opens into the larynx, whereas posteriorly, it enters the esophagus.

Larynx

The larynx is a cartilaginous structure inferior to the laryngopharynx that connects the pharynx to the trachea and helps regulate the volume of air that enters and leaves the lungs (Figure 6). The structure of the larynx is formed by several pieces of cartilage. Three large cartilage pieces—the thyroid cartilage (anterior), epiglottis (superior), and cricoid cartilage (inferior)—form the major structure of the larynx. The thyroid cartilage is the largest piece of cartilage that makes up the larynx. The thyroid cartilage consists of the laryngeal prominence, or “Adam’s apple,” which tends to be more prominent in males. The thick cricoid cartilage forms a ring, with a wide posterior region and a thinner anterior region. Three smaller, paired cartilages—the arytenoids, corniculates, and cuneiforms—attach to the epiglottis and the vocal cords and muscle that help move the vocal cords to produce speech.

[caption id="" align="aligncenter" width="450"]The top panel of this figure shows the anterior view of the larynx, and the bottom panel shows the right lateral view of the larynx. Figure 6. Larynx. The larynx extends from the laryngopharynx and the hyoid bone to the trachea.[/caption]

The epiglottis, attached to the thyroid cartilage, is a very flexible piece of elastic cartilage that covers the opening of the trachea (see Figure 3). When in the “closed” position, the unattached end of the epiglottis rests on the glottis. The glottis is composed of the vestibular folds, the true vocal cords, and the space between these folds (Figure 7). A vestibular fold, or false vocal cord, is one of a pair of folded sections of mucous membrane. A true vocal cord is one of the white, membranous folds attached by muscle to the thyroid and arytenoid cartilages of the larynx on their outer edges. The inner edges of the true vocal cords are free, allowing oscillation to produce sound. The size of the membranous folds of the true vocal cords differs between individuals, producing voices with different pitch ranges. Folds in males tend to be larger than those in females, which create a deeper voice. The act of swallowing causes the pharynx and larynx to lift upward, allowing the pharynx to expand and the epiglottis of the larynx to swing downward, closing the opening to the trachea. These movements produce a larger area for food to pass through, while preventing food and beverages from entering the trachea.

[caption id="" align="aligncenter" width="450"]This diagram shows the cross section of the larynx. The different types of cartilages are labeled. Figure 7. Vocal Cords. The true vocal cords and vestibular folds of the larynx are viewed inferiorly from the laryngopharynx.[/caption]

Continuous with the laryngopharynx, the superior portion of the larynx is lined with stratified squamous epithelium, transitioning into pseudostratified ciliated columnar epithelium that contains goblet cells. Similar to the nasal cavity and nasopharynx, this specialized epithelium produces mucus to trap debris and pathogens as they enter the trachea. The cilia beat the mucus upward towards the laryngopharynx, where it can be swallowed down the esophagus.

Trachea

The trachea (windpipe) extends from the larynx toward the lungs (Figure 8a). The trachea is formed by 16 to 20 stacked, C-shaped pieces of hyaline cartilage that are connected by dense connective tissue. The trachealis muscle and elastic connective tissue together form the fibroelastic membrane, a flexible membrane that closes the posterior surface of the trachea, connecting the C-shaped cartilages. The fibroelastic membrane allows the trachea to stretch and expand slightly during inhalation and exhalation, whereas the rings of cartilage provide structural support and prevent the trachea from collapsing. In addition, the trachealis muscle can be contracted to force air through the trachea during exhalation. The trachea is lined with pseudostratified ciliated columnar epithelium, which is continuous with the larynx. The esophagus borders the trachea posteriorly.

[caption id="" align="aligncenter" width="560"]The top panel of this figure shows the trachea and its organs. The major parts including the larynx, trachea, bronchi, and lungs are labeled. Figure 8. Trachea. (a) The tracheal tube is formed by stacked, C-shaped pieces of hyaline cartilage. (b) The layer visible in this cross-section of tracheal wall tissue between the hyaline cartilage and the lumen of the trachea is the mucosa, which is composed of pseudostratified ciliated columnar epithelium that contains goblet cells. LM × 1220. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Bronchial Tree

The trachea branches into the right and left primary bronchi at the carina. These bronchi are also lined by pseudostratified ciliated columnar epithelium containing mucus-producing goblet cells (Figure 8b). The carina is a raised structure that contains specialized nervous tissue that induces violent coughing if a foreign body, such as food, is present. Rings of cartilage, similar to those of the trachea, support the structure of the bronchi and prevent their collapse. The primary bronchi enter the lungs at the hilum, a concave region where blood vessels, lymphatic vessels, and nerves also enter the lungs. The bronchi continue to branch into bronchial a tree. A bronchial tree (or respiratory tree) is the collective term used for these multiple-branched bronchi. The main function of the bronchi, like other conducting zone structures, is to provide a passageway for air to move into and out of each lung. In addition, the mucous membrane traps debris and pathogens.

A bronchiole branches from the tertiary bronchi. Bronchioles, which are about 1 mm in diameter, further branch until they become the tiny terminal bronchioles, which lead to the structures of gas exchange. There are more than 1000 terminal bronchioles in each lung. The muscular walls of the bronchioles do not contain cartilage like those of the bronchi. This muscular wall can change the size of the tubing to increase or decrease airflow through the tube.

Respiratory Zone

In contrast to the conducting zone, the respiratory zone includes structures that are directly involved in gas exchange. The respiratory zone begins where the terminal bronchioles join a respiratory bronchiole, the smallest type of bronchiole (Figure 9), which then leads to an alveolar duct, opening into a cluster of alveoli.

[caption id="" align="aligncenter" width="525"]This image shows the bronchioles and alveolar sacs in the lungs and depicts the exchange of oxygenated and deoxygenated blood in the pulmonary blood vessels. Figure 9. Respiratory Zone. Bronchioles lead to alveolar sacs in the respiratory zone, where gas exchange occurs.[/caption]

Alveoli

An alveolar duct is a tube composed of smooth muscle and connective tissue, which opens into a cluster of alveoli. An alveolus is one of the many small, grape-like sacs that are attached to the alveolar ducts.

An alveolar sac is a cluster of many individual alveoli that are responsible for gas exchange. An alveolus is approximately 200 μm in diameter with elastic walls that allow the alveolus to stretch during air intake, which greatly increases the surface area available for gas exchange. Alveoli are connected to their neighbors by alveolar pores, which help maintain equal air pressure throughout the alveoli and lung (Figure 10).

[caption id="" align="aligncenter" width="560"]This figure shows the detailed structure of the alveolus. The top panel shows the alveolar sacs and the bronchioles. The middle panel shows a magnified view of the alveolus, and the bottom panel shows a micrograph of the cross section of a bronchiole. Figure 10. Structures of the Respiratory Zone. (a) The alveolus is responsible for gas exchange. (b) A micrograph shows the alveolar structures within lung tissue. LM × 178. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]
The alveolar wall consists of three major cell types: type I alveolar cells, type II alveolar cells, and alveolar macrophages. A type I alveolar cell is a squamous epithelial cell of the alveoli, which constitute up to 97 percent of the alveolar surface area. These cells are about 25 nm thick and are highly permeable to gases. A type II alveolar cell is interspersed among the type I cells and secretes pulmonary surfactant, a substance composed of phospholipids and proteins that reduces the surface tension of the alveoli. Roaming around the alveolar wall is the alveolar macrophage, a phagocytic cell of the immune system that removes debris and pathogens that have reached the alveoli.

The simple squamous epithelium formed by type I alveolar cells is attached to a thin, elastic basement membrane. This epithelium is extremely thin and borders the endothelial membrane of capillaries. Taken together, the alveoli and capillary membranes form a respiratory membrane that is approximately 0.5 mm thick. The respiratory membrane allows gases to cross by simple diffusion, allowing oxygen to be picked up by the blood for transport and CO2 to be released into the air of the alveoli.

[caption id="attachment_2984" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of the respiratory system![/caption]
]]> 2373 2352 5 0 22.2 The Lungs https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-2-the-lungs/ Thu, 13 Jul 2017 23:02:21 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2376

Learning Objectives

By the end of this section, you will be able to:
  • Describe the overall function of the lung
  • Summarize the blood flow pattern associated with the lungs
  • Outline the anatomy of the blood supply to the lungs
  • Describe the pleura of the lungs and their function

A major organ of the respiratory system, each lung houses structures of both the conducting and respiratory zones. The main function of the lungs is to perform the exchange of oxygen and carbon dioxide with air from the atmosphere. To this end, the lungs exchange respiratory gases across a very large epithelial surface area—about 70 square meters—that is highly permeable to gases.

Gross Anatomy of the Lungs

The lungs are pyramid-shaped, paired organs that are connected to the trachea by the right and left bronchi; on the inferior surface, the lungs are bordered by the diaphragm. The diaphragm is the flat, dome-shaped muscle located at the base of the lungs and thoracic cavity. The lungs are enclosed by the pleurae, which are attached to the mediastinum. The right lung is shorter and wider than the left lung, and the left lung occupies a smaller volume than the right. The cardiac notch is an indentation on the surface of the left lung, and it allows space for the heart (Figure 1). The apex of the lung is the superior region, whereas the base is the opposite region near the diaphragm. The costal surface of the lung borders the ribs. The mediastinal surface faces the midline.

[caption id="" align="aligncenter" width="430"]This figure shows the structure of the lungs with the major parts labeled. Figure 1. Gross Anatomy of the Lungs.[/caption]

Each lung is composed of smaller units called lobes. Fissures separate these lobes from each other. The right lung consists of three lobes: the superior, middle, and inferior lobes. The left lung consists of two lobes: the superior and inferior lobes. A bronchopulmonary segment is a division of a lobe, and each lobe houses multiple bronchopulmonary segments. Each segment receives air from its own tertiary bronchus and is supplied with blood by its own artery. Some diseases of the lungs typically affect one or more bronchopulmonary segments, and in some cases, the diseased segments can be surgically removed with little influence on neighboring segments. A pulmonary lobule is a subdivision formed as the bronchi branch into bronchioles. Each lobule receives its own large bronchiole that has multiple branches. An interlobular septum is a wall, composed of connective tissue, which separates lobules from one another.

Blood Supply and Nervous Innervation of the Lungs

The blood supply of the lungs plays an important role in gas exchange and serves as a transport system for gases throughout the body. In addition, innervation by the both the parasympathetic and sympathetic nervous systems provides an important level of control through dilation and constriction of the airway.

Blood Supply

The major function of the lungs is to perform gas exchange, which requires blood from the pulmonary circulation. This blood supply contains deoxygenated blood and travels to the lungs where erythrocytes, also known as red blood cells, pick up oxygen to be transported to tissues throughout the body. The pulmonary artery is an artery that arises from the pulmonary trunk and carries deoxygenated, arterial blood to the alveoli. The pulmonary artery branches multiple times as it follows the bronchi, and each branch becomes progressively smaller in diameter. One arteriole and an accompanying venule supply and drain one pulmonary lobule. As they near the alveoli, the pulmonary arteries become the pulmonary capillary network. The pulmonary capillary network consists of tiny vessels with very thin walls that lack smooth muscle fibers. The capillaries branch and follow the bronchioles and structure of the alveoli. It is at this point that the capillary wall meets the alveolar wall, creating the respiratory membrane. Once the blood is oxygenated, it drains from the alveoli by way of multiple pulmonary veins, which exit the lungs through the hilum.

Nervous Innervation

Dilation and constriction of the airway are achieved through nervous control by the parasympathetic and sympathetic nervous systems. The parasympathetic system causes bronchoconstriction, whereas the sympathetic nervous system stimulates bronchodilation. Reflexes such as coughing, and the ability of the lungs to regulate oxygen and carbon dioxide levels, also result from this autonomic nervous system control. Sensory nerve fibers arise from the vagus nerve, and from the second to fifth thoracic ganglia. The pulmonary plexus is a region on the lung root formed by the entrance of the nerves at the hilum. The nerves then follow the bronchi in the lungs and branch to innervate muscle fibers, glands, and blood vessels.

Pleura of the Lungs

Each lung is enclosed within a cavity that is surrounded by the pleura. The pleura (plural = pleurae) is a serous membrane that surrounds the lung. The right and left pleurae, which enclose the right and left lungs, respectively, are separated by the mediastinum. The pleurae consist of two layers. The visceral pleura is the layer that is superficial to the lungs, and extends into and lines the lung fissures (Figure 2). In contrast, the parietal pleura is the outer layer that connects to the thoracic wall, the mediastinum, and the diaphragm. The visceral and parietal pleurae connect to each other at the hilum. The pleural cavity is the space between the visceral and parietal layers.
[caption id="" align="aligncenter" width="440"]This figure shows the lungs and the chest wall, which protects the lungs, in the left panel. In the right panel, a magnified image shows the pleural cavity and a pleural sac. Figure 2. Parietal and Visceral Pleurae of the Lungs.[/caption]

The pleurae perform two major functions: They produce pleural fluid and create cavities that separate the major organs. Pleural fluid is secreted by mesothelial cells from both pleural layers and acts to lubricate their surfaces. This lubrication reduces friction between the two layers to prevent trauma during breathing, and creates surface tension that helps maintain the position of the lungs against the thoracic wall. This adhesive characteristic of the pleural fluid causes the lungs to enlarge when the thoracic wall expands during ventilation, allowing the lungs to fill with air. The pleurae also create a division between major organs that prevents interference due to the movement of the organs, while preventing the spread of infection.

]]> 2376 2352 6 0 22.3 The Process of Breathing https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-3-the-process-of-breathing/ Thu, 13 Jul 2017 23:02:46 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2383

Learning Objectives

By the end of this section, you will be able to:
  • Describe the mechanisms that drive breathing
  • Discuss how pressure, volume, and resistance are related
  • List the steps involved in pulmonary ventilation
  • Discuss the physical factors related to breathing
  • Discuss the meaning of respiratory volume and capacities
  • Define respiratory rate
  • Outline the mechanisms behind the control of breathing
  • Describe the respiratory centers of the medulla oblongata
  • Describe the respiratory centers of the pons
  • Discuss factors that can influence the respiratory rate

Pulmonary ventilation is the act of breathing, which can be described as the movement of air into and out of the lungs. The major mechanisms that drive pulmonary ventilation are atmospheric pressure (Patm); the air pressure within the alveoli, called alveolar pressure (Palv); and the pressure within the pleural cavity, called intrapleural pressure (Pip).

Mechanisms of Breathing

The alveolar and intrapleural pressures are dependent on certain physical features of the lung. However, the ability to breathe—to have air enter the lungs during inspiration and air leave the lungs during expiration—is dependent on the air pressure of the atmosphere and the air pressure within the lungs.

Pressure Relationships

Inspiration (or inhalation) and expiration (or exhalation) are dependent on the differences in pressure between the atmosphere and the lungs. In a gas, pressure is a force created by the movement of gas molecules that are confined. For example, a certain number of gas molecules in a two-liter container has more room than the same number of gas molecules in a one-liter container (Figure 1). In this case, the force exerted by the movement of the gas molecules against the walls of the two-liter container is lower than the force exerted by the gas molecules in the one-liter container. Therefore, the pressure is lower in the two-liter container and higher in the one-liter container. At a constant temperature, changing the volume occupied by the gas changes the pressure, as does changing the number of gas molecules. Boyle’s law describes the relationship between volume and pressure in a gas at a constant temperature. Boyle discovered that the pressure of a gas is inversely proportional to its volume: If volume increases, pressure decreases. Likewise, if volume decreases, pressure increases. Pressure and volume are inversely related (P = k/V). Therefore, the pressure in the one-liter container (one-half the volume of the two-liter container) would be twice the pressure in the two-liter container. Boyle’s law is expressed by the following formula:
P1V1 = P2V2
In this formula, P1 represents the initial pressure and V1 represents the initial volume, whereas the final pressure and volume are represented by P2 and V2, respectively. If the two- and one-liter containers were connected by a tube and the volume of one of the containers were changed, then the gases would move from higher pressure (lower volume) to lower pressure (higher volume).
[caption id="" align="aligncenter" width="420"]This diagram shows two canisters containing a gas. The two canisters show how volume and pressure are inversely proportional, which illustrates Boyle’s law. Figure 1. Boyle's Law. In a gas, pressure increases as volume decreases.[/caption]

Pulmonary ventilation is dependent on three types of pressure: atmospheric, intra-alveolar, and interpleural. Atmospheric pressure is the amount of force that is exerted by gases in the air surrounding any given surface, such as the body. Atmospheric pressure can be expressed in terms of the unit atmosphere, abbreviated atm, or in millimeters of mercury (mm Hg). One atm is equal to 760 mm Hg, which is the atmospheric pressure at sea level. Typically, for respiration, other pressure values are discussed in relation to atmospheric pressure. Therefore, negative pressure is pressure lower than the atmospheric pressure, whereas positive pressure is pressure that it is greater than the atmospheric pressure. A pressure that is equal to the atmospheric pressure is expressed as zero.

Intra-alveolar pressure is the pressure of the air within the alveoli, which changes during the different phases of breathing (Figure 2). Because the alveoli are connected to the atmosphere via the tubing of the airways (similar to the two- and one-liter containers in the example above), the interpulmonary pressure of the alveoli always equalizes with the atmospheric pressure.
[caption id="" align="aligncenter" width="455"]This diagram shows the lungs and the air pressure in different regions. Figure 2. Intrapulmonary and Intrapleural Pressure Relationships. Alveolar pressure changes during the different phases of the cycle. It equalizes at 760 mm Hg but does not remain at 760 mm Hg.[/caption]

Intrapleural pressure is the pressure of the air within the pleural cavity, between the visceral and parietal pleurae. Similar to intra-alveolar pressure, intrapleural pressure also changes during the different phases of breathing. However, due to certain characteristics of the lungs, the intrapleural pressure is always lower than, or negative to, the intra-alveolar pressure (and therefore also to atmospheric pressure). Although it fluctuates during inspiration and expiration, intrapleural pressure remains approximately –4 mm Hg throughout the breathing cycle.

Competing forces within the thorax cause the formation of the negative intrapleural pressure. One of these forces relates to the elasticity of the lungs themselves—elastic tissue pulls the lungs inward, away from the thoracic wall. Surface tension of alveolar fluid, which is mostly water, also creates an inward pull of the lung tissue. This inward tension from the lungs is countered by opposing forces from the pleural fluid and thoracic wall. Surface tension within the pleural cavity pulls the lungs outward. Too much or too little pleural fluid would hinder the creation of the negative intrapleural pressure; therefore, the level must be closely monitored by the mesothelial cells and drained by the lymphatic system. Since the parietal pleura is attached to the thoracic wall, the natural elasticity of the chest wall opposes the inward pull of the lungs. Ultimately, the outward pull is slightly greater than the inward pull, creating the –4 mm Hg intrapleural pressure relative to the intra-alveolar pressure. Transpulmonary pressure is the difference between the intrapleural and intra-alveolar pressures, and it determines the size of the lungs. A higher transpulmonary pressure corresponds to a larger lung.

Physical Factors Affecting Ventilation

In addition to the differences in pressures, breathing is also dependent upon the contraction and relaxation of muscle fibers of both the diaphragm and thorax. The lungs themselves are passive during breathing, meaning they are not involved in creating the movement that helps inspiration and expiration. This is because of the adhesive nature of the pleural fluid, which allows the lungs to be pulled outward when the thoracic wall moves during inspiration. The recoil of the thoracic wall during expiration causes compression of the lungs. Contraction and relaxation of the diaphragm and intercostals muscles (found between the ribs) cause most of the pressure changes that result in inspiration and expiration. These muscle movements and subsequent pressure changes cause air to either rush in or be forced out of the lungs.

Other characteristics of the lungs influence the effort that must be expended to ventilate. Resistance is a force that slows motion, in this case, the flow of gases. The size of the airway is the primary factor affecting resistance. A small tubular diameter forces air through a smaller space, causing more collisions of air molecules with the walls of the airways. The following formula helps to describe the relationship between airway resistance and pressure changes:

F = ∆P / R

As noted earlier, there is surface tension within the alveoli caused by water present in the lining of the alveoli. This surface tension tends to inhibit expansion of the alveoli. However, pulmonary surfactant secreted by type II alveolar cells mixes with that water and helps reduce this surface tension. Without pulmonary surfactant, the alveoli would collapse during expiration.

Thoracic wall compliance is the ability of the thoracic wall to stretch while under pressure. This can also affect the effort expended in the process of breathing. In order for inspiration to occur, the thoracic cavity must expand. The expansion of the thoracic cavity directly influences the capacity of the lungs to expand. If the tissues of the thoracic wall are not very compliant, it will be difficult to expand the thorax to increase the size of the lungs.

Pulmonary Ventilation

The difference in pressures drives pulmonary ventilation because air flows down a pressure gradient, that is, air flows from an area of higher pressure to an area of lower pressure. Air flows into the lungs largely due to a difference in pressure; atmospheric pressure is greater than intra-alveolar pressure, and intra-alveolar pressure is greater than intrapleural pressure. Air flows out of the lungs during expiration based on the same principle; pressure within the lungs becomes greater than the atmospheric pressure.

Pulmonary ventilation comprises two major steps: inspiration and expiration. Inspiration is the process that causes air to enter the lungs, and expiration is the process that causes air to leave the lungs (Figure 3). A respiratory cycle is one sequence of inspiration and expiration. In general, two muscle groups are used during normal inspiration: the diaphragm and the external intercostal muscles. Additional muscles can be used if a bigger breath is required. When the diaphragm contracts, it moves inferiorly toward the abdominal cavity, creating a larger thoracic cavity and more space for the lungs. Contraction of the external intercostal muscles moves the ribs upward and outward, causing the rib cage to expand, which increases the volume of the thoracic cavity. Due to the adhesive force of the pleural fluid, the expansion of the thoracic cavity forces the lungs to stretch and expand as well. This increase in volume leads to a decrease in intra-alveolar pressure, creating a pressure lower than atmospheric pressure. As a result, a pressure gradient is created that drives air into the lungs.

[caption id="" align="aligncenter" width="450"]The left panel of this image shows a person inhaling air and the location of the chest muscles. The right panel shows the person exhaling air and the contraction of the thoracic cavity. Figure 3. Inspiration and Expiration. Inspiration and expiration occur due to the expansion and contraction of the thoracic cavity, respectively.[/caption]

The process of normal expiration is passive, meaning that energy is not required to push air out of the lungs. Instead, the elasticity of the lung tissue causes the lung to recoil, as the diaphragm and intercostal muscles relax following inspiration. In turn, the thoracic cavity and lungs decrease in volume, causing an increase in interpulmonary pressure. The interpulmonary pressure rises above atmospheric pressure, creating a pressure gradient that causes air to leave the lungs.

There are different types, or modes, of breathing that require a slightly different process to allow inspiration and expiration. Quiet breathing, also known as eupnea, is a mode of breathing that occurs at rest and does not require the cognitive thought of the individual. During quiet breathing, the diaphragm and external intercostals must contract.

A deep breath, called diaphragmatic breathing, requires the diaphragm to contract. As the diaphragm relaxes, air passively leaves the lungs. A shallow breath, called costal breathing, requires contraction of the intercostal muscles. As the intercostal muscles relax, air passively leaves the lungs.

In contrast, forced breathing, also known as hyperpnea, is a mode of breathing that can occur during exercise or actions that require the active manipulation of breathing, such as singing. During forced breathing, inspiration and expiration both occur due to muscle contractions. In addition to the contraction of the diaphragm and intercostal muscles, other accessory muscles must also contract. During forced inspiration, muscles of the neck, including the scalenes, contract and lift the thoracic wall, increasing lung volume. During forced expiration, accessory muscles of the abdomen, including the obliques, contract, forcing abdominal organs upward against the diaphragm. This helps to push the diaphragm further into the thorax, pushing more air out. In addition, accessory muscles (primarily the internal intercostals) help to compress the rib cage, which also reduces the volume of the thoracic cavity.

Respiratory Volumes and Capacities

Respiratory volume is the term used for various volumes of air moved by or associated with the lungs at a given point in the respiratory cycle. There are four major types of respiratory volumes: tidal, residual, inspiratory reserve, and expiratory reserve (Figure 4). Tidal volume (TV) is the amount of air that normally enters the lungs during quiet breathing, which is about 500 milliliters. Expiratory reserve volume (ERV) is the amount of air you can forcefully exhale past a normal tidal expiration, up to 1200 milliliters for men. Inspiratory reserve volume (IRV) is produced by a deep inhalation, past a tidal inspiration. This is the extra volume that can be brought into the lungs during a forced inspiration. Residual volume (RV) is the air left in the lungs if you exhale as much air as possible. The residual volume makes breathing easier by preventing the alveoli from collapsing. Respiratory volume is dependent on a variety of factors, and measuring the different types of respiratory volumes can provide important clues about a person’s respiratory health (Figure 5).

[caption id="" align="aligncenter" width="455"]The left panel shows a graph of different respiratory volumes. The right panel shows how the different respiratory volumes result in respiratory capacity. Figure 4. Respiratory Volumes and Capacities. These two graphs show (a) respiratory volumes and (b) the combination of volumes that results in respiratory capacity.[/caption]
[caption id="" align="aligncenter" width="500"]This tables describes methods of pulmonary function testing. Spirometry tests require a spirometer. These tests can measure forced vital capacity (FVC), the volume of air that is exhaled after maximum inhalation; foreced expiratory volume (FEV), the volume of air exhaled in one breath; forced expiratory flow, 25 to 75 percent, the air flow in the middle of exhalation; peak expiratory flow (PEF), the rate of exhalation; maximum voluntary ventilation (MVV), the volume of air that can be inspired and expired in 1 minute; slow vital capacity (SVC), the volume of air that can be slowly exhaled after inhaling past the tidal volume; total lung capacity (TLC), the volume of air in the lungs after maximum inhalation; functional residual capacity (FRC), the volume of air left in the lungs after normal expiration; residual volume (RV), the volume of air in the lungs after maximum exhalation; total lung capacity (TLC), the maximum volume of air that the lungs can hold; and expiratory reserve volume (ERV), the volume of air that can be exhaled beyond normal exhalation. Gas diffusion tests require a blood gas analyzer. These tests can measure arterial blood gases, the concentration of oxygen and carbon dioxide in the blood. Figure 5. Pulmonary Function Testing.[/caption]

Respiratory capacity is the combination of two or more selected volumes, which further describes the amount of air in the lungs during a given time. For example, total lung capacity (TLC) is the sum of all of the lung volumes (TV, ERV, IRV, and RV), which represents the total amount of air a person can hold in the lungs after a forceful inhalation. TLC is about 6000 mL air for men, and about 4200 mL for women. Vital capacity (VC) is the amount of air a person can move into or out of his or her lungs, and is the sum of all of the volumes except residual volume (TV, ERV, and IRV), which is between 4000 and 5000 milliliters. Inspiratory capacity (IC) is the maximum amount of air that can be inhaled past a normal tidal expiration, is the sum of the tidal volume and inspiratory reserve volume. On the other hand, the functional residual capacity (FRC) is the amount of air that remains in the lung after a normal tidal expiration; it is the sum of expiratory reserve volume and residual volume (see Figure 4).

In addition to the air that creates respiratory volumes, the respiratory system also contains anatomical dead space, which is air that is present in the airway that never reaches the alveoli and therefore never participates in gas exchange. Alveolar dead space involves air found within alveoli that are unable to function, such as those affected by disease or abnormal blood flow. Total dead space is the anatomical dead space and alveolar dead space together, and represents all of the air in the respiratory system that is not being used in the gas exchange process.

Respiratory Rate and Control of Ventilation

Breathing usually occurs without thought, although at times you can consciously control it, such as when you swim under water, sing a song, or blow bubbles. The respiratory rate is the total number of breaths, or respiratory cycles, that occur each minute. Respiratory rate can be an important indicator of disease, as the rate may increase or decrease during an illness or in a disease condition. The respiratory rate is controlled by the respiratory center located within the medulla oblongata in the brain, which responds primarily to changes in carbon dioxide, oxygen, and pH levels in the blood.

The normal respiratory rate of a child decreases from birth to adolescence. A child under 1 year of age has a normal respiratory rate between 30 and 60 breaths per minute, but by the time a child is about 10 years old, the normal rate is closer to 18 to 30. By adolescence, the normal respiratory rate is similar to that of adults, 12 to 18 breaths per minute.

Ventilation Control Centers

The control of ventilation is a complex interplay of multiple regions in the brain that signal the muscles used in pulmonary ventilation to contract (Table 1). The result is typically a rhythmic, consistent ventilation rate that provides the body with sufficient amounts of oxygen, while adequately removing carbon dioxide.

Summary of Ventilation Regulation (Table 1)
System component Function
Medullary respiratory renter Sets the basic rhythm of breathing
Ventral respiratory group (VRG) Generates the breathing rhythm and integrates data coming into the medulla
Dorsal respiratory group (DRG) Integrates input from the stretch receptors and the chemoreceptors in the periphery
Pontine respiratory group (PRG) Influences and modifies the medulla oblongata’s functions
Aortic body Monitors blood PCO2, PO2, and pH
Carotid body Monitors blood PCO2, PO2, and pH
Hypothalamus Monitors emotional state and body temperature
Cortical areas of the brain Control voluntary breathing
Proprioceptors Send impulses regarding joint and muscle movements
Pulmonary irritant reflexes Protect the respiratory zones of the system from foreign material
Inflation reflex Protects the lungs from over-inflating

Neurons that innervate the muscles of the respiratory system are responsible for controlling and regulating pulmonary ventilation. The major brain centers involved in pulmonary ventilation are the medulla oblongata and the pontine respiratory group (Figure 6).

[caption id="" align="aligncenter" width="465"]The top panel of this image shows the regions of the brain that control respiration. The middle panel shows a magnified view of these regions and links the regions of the brain to the specific organs that they control. Figure 6. Respiratory Centers of the Brain.[/caption]

The medulla oblongata contains the dorsal respiratory group (DRG) and the ventral respiratory group (VRG). The DRG is involved in maintaining a constant breathing rhythm by stimulating the diaphragm and intercostal muscles to contract, resulting in inspiration. When activity in the DRG ceases, it no longer stimulates the diaphragm and intercostals to contract, allowing them to relax, resulting in expiration. The VRG is involved in forced breathing, as the neurons in the VRG stimulate the accessory muscles involved in forced breathing to contract, resulting in forced inspiration. The VRG also stimulates the accessory muscles involved in forced expiration to contract.

The second respiratory center of the brain is located within the pons, called the pontine respiratory group, and consists of the apneustic and pneumotaxic centers. The apneustic center is a double cluster of neuronal cell bodies that stimulate neurons in the DRG, controlling the depth of inspiration, particularly for deep breathing. The pneumotaxic center is a network of neurons that inhibits the activity of neurons in the DRG, allowing relaxation after inspiration, and thus controlling the overall rate.

Factors That Affect the Rate and Depth of Respiration

The respiratory rate and the depth of inspiration are regulated by the medulla oblongata and pons; however, these regions of the brain do so in response to systemic stimuli. It is a dose-response, positive-feedback relationship in which the greater the stimulus, the greater the response. Thus, increasing stimuli results in forced breathing. Multiple systemic factors are involved in stimulating the brain to produce pulmonary ventilation.

The major factor that stimulates the medulla oblongata and pons to produce respiration is surprisingly not oxygen concentration, but rather the concentration of carbon dioxide in the blood. As you recall, carbon dioxide is a waste product of cellular respiration and can be toxic. Concentrations of chemicals are sensed by chemoreceptors. A central chemoreceptor is one of the specialized receptors that are located in the brain and brainstem, whereas a peripheral chemoreceptor is one of the specialized receptors located in the carotid arteries and aortic arch. Concentration changes in certain substances, such as carbon dioxide or hydrogen ions, stimulate these receptors, which in turn signal the respiration centers of the brain. In the case of carbon dioxide, as the concentration of CO2 in the blood increases, it readily diffuses across the blood-brain barrier, where it collects in the extracellular fluid. As will be explained in more detail later, increased carbon dioxide levels lead to increased levels of hydrogen ions, decreasing pH. The increase in hydrogen ions in the brain triggers the central chemoreceptors to stimulate the respiratory centers to initiate contraction of the diaphragm and intercostal muscles. As a result, the rate and depth of respiration increase, allowing more carbon dioxide to be expelled, which brings more air into and out of the lungs promoting a reduction in the blood levels of carbon dioxide, and therefore hydrogen ions, in the blood. In contrast, low levels of carbon dioxide in the blood cause low levels of hydrogen ions in the brain, leading to a decrease in the rate and depth of pulmonary ventilation, producing shallow, slow breathing.

Another factor involved in influencing the respiratory activity of the brain is systemic arterial concentrations of hydrogen ions. Increasing carbon dioxide levels can lead to increased H+ levels, as mentioned above, as well as other metabolic activities, such as lactic acid accumulation after strenuous exercise. Peripheral chemoreceptors of the aortic arch and carotid arteries sense arterial levels of hydrogen ions. When peripheral chemoreceptors sense decreasing, or more acidic, pH levels, they stimulate an increase in ventilation to remove carbon dioxide from the blood at a quicker rate. Removal of carbon dioxide from the blood helps to reduce hydrogen ions, thus increasing systemic pH.

Blood levels of oxygen are also important in influencing respiratory rate. The peripheral chemoreceptors are responsible for sensing large changes in blood oxygen levels. If blood oxygen levels become quite low—about 60 mm Hg or less—then peripheral chemoreceptors stimulate an increase in respiratory activity. The chemoreceptors are only able to sense dissolved oxygen molecules, not the oxygen that is bound to hemoglobin. As you recall, the majority of oxygen is bound by hemoglobin; when dissolved levels of oxygen drop, hemoglobin releases oxygen. Therefore, a large drop in oxygen levels is required to stimulate the chemoreceptors of the aortic arch and carotid arteries.

The hypothalamus and other brain regions associated with the limbic system also play roles in influencing the regulation of breathing by interacting with the respiratory centers. The hypothalamus and other regions associated with the limbic system are involved in regulating respiration in response to emotions, pain, and temperature. For example, an increase in body temperature causes an increase in respiratory rate. Feeling excited or the fight-or-flight response will also result in an increase in respiratory rate. Watch this CrashCourse video to earn more about the breathing process.
]]> 2383 2352 7 0 22.4 Gas Exchange https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-4-gas-exchange/ Thu, 13 Jul 2017 23:03:45 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2386

Learning Objectives

By the end of this section, you will be able to:
  • Compare the composition of atmospheric air and alveolar air
  • Describe the mechanisms that drive gas exchange
  • Discuss the importance of sufficient ventilation and perfusion, and how the body adapts when they are insufficient
  • Discuss the process of external respiration
  • Describe the process of internal respiration

The purpose of the respiratory system is to perform gas exchange. Pulmonary ventilation provides air to the alveoli for this gas exchange process. At the respiratory membrane, where the alveolar and capillary walls meet, gases move across the membranes, with oxygen entering the bloodstream and carbon dioxide exiting. It is through this mechanism that blood is oxygenated and carbon dioxide, the waste product of cellular respiration, is removed from the body.

Gas Exchange

In order to understand the mechanisms of gas exchange in the lung, it is important to understand the underlying principles of gases and their behavior. In addition to Boyle’s law, several other gas laws help to describe the behavior of gases.

Gas Laws and Air Composition

Gas molecules exert force on the surfaces with which they are in contact; this force is called pressure. In natural systems, gases are normally present as a mixture of different types of molecules. For example, the atmosphere consists of oxygen, nitrogen, carbon dioxide, and other gaseous molecules, and this gaseous mixture exerts a certain pressure referred to as atmospheric pressure (Table 2). Partial pressure (Px) is the pressure of a single type of gas in a mixture of gases. For example, in the atmosphere, oxygen exerts a partial pressure, and nitrogen exerts another partial pressure, independent of the partial pressure of oxygen (Figure 1). Total pressure is the sum of all the partial pressures of a gaseous mixture. Dalton’s law describes the behavior of nonreactive gases in a gaseous mixture and states that a specific gas type in a mixture exerts its own pressure; thus, the total pressure exerted by a mixture of gases is the sum of the partial pressures of the gases in the mixture.

Partial Pressures of Atmospheric Gases (Table 2)
Gas Percent of total composition Partial pressure
(mm Hg)
Nitrogen (N2) 78.6 597.4
Oxygen (O2) 20.9 158.8
Water (H2O) 0.04 3.0
Carbon dioxide (CO2) 0.004 0.3
Others 0.0006 0.5
Total composition/total atmospheric pressure 100% 760.0
Figure 1. Partial and Total Pressures of a Gas. Partial pressure is the force exerted by a gas. The sum of the partial pressures of all the gases in a mixture equals the total pressure.

Partial pressure is extremely important in predicting the movement of gases. Recall that gases tend to equalize their pressure in two regions that are connected. A gas will move from an area where its partial pressure is higher to an area where its partial pressure is lower. In addition, the greater the partial pressure difference between the two areas, the more rapid is the movement of gases.

Solubility of Gases in Liquids

Henry’s law describes the behavior of gases when they come into contact with a liquid, such as blood. Henry’s law states that the concentration of gas in a liquid is directly proportional to the solubility and partial pressure of that gas. The greater the partial pressure of the gas, the greater the number of gas molecules that will dissolve in the liquid. The concentration of the gas in a liquid is also dependent on the solubility of the gas in the liquid. For example, although nitrogen is present in the atmosphere, very little nitrogen dissolves into the blood, because the solubility of nitrogen in blood is very low. The exception to this occurs in scuba divers; the composition of the compressed air that divers breathe causes nitrogen to have a higher partial pressure than normal, causing it to dissolve in the blood in greater amounts than normal. Too much nitrogen in the bloodstream results in a serious condition that can be fatal if not corrected. Gas molecules establish an equilibrium between those molecules dissolved in liquid and those in air.

The composition of air in the atmosphere and in the alveoli differs. In both cases, the relative concentration of gases is nitrogen > oxygen > water vapor > carbon dioxide. The amount of water vapor present in alveolar air is greater than that in atmospheric air (Table 3). Recall that the respiratory system works to humidify incoming air, thereby causing the air present in the alveoli to have a greater amount of water vapor than atmospheric air. In addition, alveolar air contains a greater amount of carbon dioxide and less oxygen than atmospheric air. This is no surprise, as gas exchange removes oxygen from and adds carbon dioxide to alveolar air. Both deep and forced breathing cause the alveolar air composition to be changed more rapidly than during quiet breathing. As a result, the partial pressures of oxygen and carbon dioxide change, affecting the diffusion process that moves these materials across the membrane. This will cause oxygen to enter and carbon dioxide to leave the blood more quickly.

Composition and Partial Pressures of Alveolar Air (Table 3)
Gas Percent of total composition Partial pressure
(mm Hg)
Nitrogen (N2) 74.9 569
Oxygen (O2) 13.7 104
Water (H2O) 6.2 40
Carbon dioxide (CO2) 5.2 47
Total composition/total alveolar pressure 100% 760.0

Ventilation and Perfusion

Two important aspects of gas exchange in the lung are ventilation and perfusion. Ventilation is the movement of air into and out of the lungs, and perfusion is the flow of blood in the pulmonary capillaries. For gas exchange to be efficient, the volumes involved in ventilation and perfusion should be compatible. However, factors such as regional gravity effects on blood, blocked alveolar ducts, or disease can cause ventilation and perfusion to be imbalanced.

The partial pressure of oxygen in alveolar air is about 104 mm Hg, whereas the partial pressure of the oxygenated pulmonary venous blood is about 100 mm Hg. When ventilation is sufficient, oxygen enters the alveoli at a high rate, and the partial pressure of oxygen in the alveoli remains high. In contrast, when ventilation is insufficient, the partial pressure of oxygen in the alveoli drops. Without the large difference in partial pressure between the alveoli and the blood, oxygen does not diffuse efficiently across the respiratory membrane. The body has mechanisms that counteract this problem. In cases when ventilation is not sufficient for an alveolus, the body redirects blood flow to alveoli that are receiving sufficient ventilation. This is achieved by constricting the pulmonary arterioles that serves the dysfunctional alveolus, which redirects blood to other alveoli that have sufficient ventilation. At the same time, the pulmonary arterioles that serve alveoli receiving sufficient ventilation vasodilate, which brings in greater blood flow. Factors such as carbon dioxide, oxygen, and pH levels can all serve as stimuli for adjusting blood flow in the capillary networks associated with the alveoli.

Ventilation is regulated by the diameter of the airways, whereas perfusion is regulated by the diameter of the blood vessels. The diameter of the bronchioles is sensitive to the partial pressure of carbon dioxide in the alveoli. A greater partial pressure of carbon dioxide in the alveoli causes the bronchioles to increase their diameter as will a decreased level of oxygen in the blood supply, allowing carbon dioxide to be exhaled from the body at a greater rate. As mentioned above, a greater partial pressure of oxygen in the alveoli causes the pulmonary arterioles to dilate, increasing blood flow.

Gas Exchange

Gas exchange occurs at two sites in the body: in the lungs, where oxygen is picked up and carbon dioxide is released at the respiratory membrane, and at the tissues, where oxygen is released and carbon dioxide is picked up. External respiration is the exchange of gases with the external environment, and occurs in the alveoli of the lungs. Internal respiration is the exchange of gases with the internal environment, and occurs in the tissues. The actual exchange of gases occurs due to simple diffusion. Energy is not required to move oxygen or carbon dioxide across membranes. Instead, these gases follow pressure gradients that allow them to diffuse. The anatomy of the lung maximizes the diffusion of gases: The respiratory membrane is highly permeable to gases; the respiratory and blood capillary membranes are very thin; and there is a large surface area throughout the lungs.

External Respiration

The pulmonary artery carries deoxygenated blood into the lungs from the heart, where it branches and eventually becomes the capillary network composed of pulmonary capillaries. These pulmonary capillaries create the respiratory membrane with the alveoli (Figure 2). As the blood is pumped through this capillary network, gas exchange occurs. Although a small amount of the oxygen is able to dissolve directly into plasma from the alveoli, most of the oxygen is picked up by erythrocytes (red blood cells) and binds to a protein called hemoglobin, a process described later in this chapter. Oxygenated hemoglobin is red, causing the overall appearance of bright red oxygenated blood, which returns to the heart through the pulmonary veins. Carbon dioxide is released in the opposite direction of oxygen, from the blood to the alveoli. Some of the carbon dioxide is returned on hemoglobin, but can also be dissolved in plasma or is present as a converted form, also explained in greater detail later in this chapter.

External respiration occurs as a function of partial pressure differences in oxygen and carbon dioxide between the alveoli and the blood in the pulmonary capillaries.

[caption id="" align="aligncenter" width="425"]This figure shows the pathway in which external respiration takes place. The exchange of oxygen and carbon dioxide between the alveolus and blood plasma is detailed. Figure 3. External Respiration. In external respiration, oxygen diffuses across the respiratory membrane from the alveolus to the capillary, whereas carbon dioxide diffuses out of the capillary into the alveolus.[/caption]

Although the solubility of oxygen in blood is not high, there is a drastic difference in the partial pressure of oxygen in the alveoli versus in the blood of the pulmonary capillaries. This difference is about 64 mm Hg: The partial pressure of oxygen in the alveoli is about 104 mm Hg, whereas its partial pressure in the blood of the capillary is about 40 mm Hg. This large difference in partial pressure creates a very strong pressure gradient that causes oxygen to rapidly cross the respiratory membrane from the alveoli into the blood.

The partial pressure of carbon dioxide is also different between the alveolar air and the blood of the capillary. However, the partial pressure difference is less than that of oxygen, about 5 mm Hg. The partial pressure of carbon dioxide in the blood of the capillary is about 45 mm Hg, whereas its partial pressure in the alveoli is about 40 mm Hg. However, the solubility of carbon dioxide is much greater than that of oxygen—by a factor of about 20—in both blood and alveolar fluids. As a result, the relative concentrations of oxygen and carbon dioxide that diffuse across the respiratory membrane are similar.

Internal Respiration

Internal respiration is gas exchange that occurs at the level of body tissues (Figure 3). Similar to external respiration, internal respiration also occurs as simple diffusion due to a partial pressure gradient. However, the partial pressure gradients are opposite of those present at the respiratory membrane. The partial pressure of oxygen in tissues is low, about 40 mm Hg, because oxygen is continuously used for cellular respiration. In contrast, the partial pressure of oxygen in the blood is about 100 mm Hg. This creates a pressure gradient that causes oxygen to dissociate from hemoglobin, diffuse out of the blood, cross the interstitial space, and enter the tissue. Hemoglobin that has little oxygen bound to it loses much of its brightness, so that blood returning to the heart is more burgundy in color.

Considering that cellular respiration continuously produces carbon dioxide, the partial pressure of carbon dioxide is lower in the blood than it is in the tissue, causing carbon dioxide to diffuse out of the tissue, cross the interstitial fluid, and enter the blood. It is then carried back to the lungs either bound to hemoglobin, dissolved in plasma, or in a converted form. By the time blood returns to the heart, the partial pressure of oxygen has returned to about 40 mm Hg, and the partial pressure of carbon dioxide has returned to about 45 mm Hg. The blood is then pumped back to the lungs to be oxygenated once again during external respiration.

[caption id="" align="aligncenter" width="425"]This diagram details the pathway of internal respiration. The exchange of oxygen and carbon dioxide between a red blood cell and a tissue cell is shown. Figure 3. Internal Respiration. Oxygen diffuses out of the capillary and into cells, whereas carbon dioxide diffuses out of cells and into the capillary.[/caption]
]]> 2386 2352 8 0 22.5 Transport of Gases https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-5-transport-of-gases/ Thu, 13 Jul 2017 23:04:01 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2393

Learning Objectives

By the end of this section, you will be able to:
  • Describe the principles of oxygen transport
  • Describe the structure of hemoglobin
  • Compare and contrast fetal and adult hemoglobin
  • Describe the principles of carbon dioxide transport

The other major activity in the lungs is the process of respiration, the process of gas exchange. The function of respiration is to provide oxygen for use by body cells during cellular respiration and to eliminate carbon dioxide, a waste product of cellular respiration, from the body. In order for the exchange of oxygen and carbon dioxide to occur, both gases must be transported between the external and internal respiration sites. Although carbon dioxide is more soluble than oxygen in blood, both gases require a specialized transport system for the majority of the gas molecules to be moved between the lungs and other tissues.

Oxygen Transport in the Blood

Even though oxygen is transported via the blood, you may recall that oxygen is not very soluble in liquids. A small amount of oxygen does dissolve in the blood and is transported in the bloodstream, but it is only about 1.5% of the total amount. The majority of oxygen molecules are carried from the lungs to the body’s tissues by a specialized transport system, which relies on the erythrocyte—the red blood cell. Erythrocytes contain a metalloprotein, hemoglobin, which serves to bind oxygen molecules to the erythrocyte (Figure 1). Heme is the portion of hemoglobin that contains iron, and it is heme that binds oxygen. One hemoglobin molecule contains iron-containing Heme molecules, and because of this, each hemoglobin molecule is capable of carrying up to four molecules of oxygen. As oxygen diffuses across the respiratory membrane from the alveolus to the capillary, it also diffuses into the red blood cell and is bound by hemoglobin. The following reversible chemical reaction describes the production of the final product, oxyhemoglobin (Hb–O2), which is formed when oxygen binds to hemoglobin. Oxyhemoglobin is a bright red-colored molecule that contributes to the bright red color of oxygenated blood.

Hb + O2 ↔ Hb − O2

In this formula, Hb represents reduced hemoglobin, that is, hemoglobin that does not have oxygen bound to it. There are multiple factors involved in how readily heme binds to and dissociates from oxygen, which will be discussed in the subsequent sections.

[caption id="" align="aligncenter" width="300"]This diagram shows a red blood cell and the structure of a hemoglobin molecule. Figure 1. Erythrocyte and Hemoglobin. Hemoglobin consists of four subunits, each of which contains one molecule of iron.[/caption]

Function of Hemoglobin

Hemoglobin is composed of subunits, a protein structure that is referred to as a quaternary structure. Each of the four subunits that make up hemoglobin is arranged in a ring-like fashion, with an iron atom covalently bound to the heme in the center of each subunit. Binding of the first oxygen molecule causes a conformational change in hemoglobin that allows the second molecule of oxygen to bind more readily. As each molecule of oxygen is bound, it further facilitates the binding of the next molecule, until all four heme sites are occupied by oxygen. The opposite occurs as well: After the first oxygen molecule dissociates and is “dropped off” at the tissues, the next oxygen molecule dissociates more readily. When all four heme sites are occupied, the hemoglobin is said to be saturated. When one to three heme sites are occupied, the hemoglobin is said to be partially saturated. Therefore, when considering the blood as a whole, the percent of the available heme units that are bound to oxygen at a given time is called hemoglobin saturation. Hemoglobin saturation of 100 percent means that every heme unit in all of the erythrocytes of the body is bound to oxygen. In a healthy individual with normal hemoglobin levels, hemoglobin saturation generally ranges from 95 percent to 99 percent.

Oxygen Dissociation from Hemoglobin

Partial pressure is an important aspect of the binding of oxygen to and disassociation from heme. An oxygen–hemoglobin dissociation curve is a graph that describes the relationship of partial pressure to the binding of oxygen to heme and its subsequent dissociation from heme (Figure 2). Remember that gases travel from an area of higher partial pressure to an area of lower partial pressure. In addition, the affinity of an oxygen molecule for heme increases as more oxygen molecules are bound. Therefore, in the oxygen–hemoglobin saturation curve, as the partial pressure of oxygen increases, a proportionately greater number of oxygen molecules are bound by heme. Not surprisingly, the oxygen–hemoglobin saturation/dissociation curve also shows that the lower the partial pressure of oxygen, the fewer oxygen molecules are bound to heme. As a result, the partial pressure of oxygen plays a major role in determining the degree of binding of oxygen to heme at the site of the respiratory membrane, as well as the degree of dissociation of oxygen from heme at the site of body tissues.

The top panel of this figure shows a graph with oxygen saturation of the y-axis and partial pressure of oxygen on the x-axis.The middle panel shows oxygen saturation versus partial pressure of oxygen as a function of pH. [caption id="" align="aligncenter" width="400"]The bottom panel shows the same relationship as a function of temperature. Figure 2. Oxygen-Hemoglobin Dissociation and Effects of pH and Temperature. These three graphs show (a) the relationship between the partial pressure of oxygen and hemoglobin saturation, (b) the effect of pH on the oxygen–hemoglobin dissociation curve, and (c) the effect of temperature on the oxygen–hemoglobin dissociation curve.[/caption]

The mechanisms behind the oxygen–hemoglobin saturation/dissociation curve also serve as automatic control mechanisms that regulate how much oxygen is delivered to different tissues throughout the body. This is important because some tissues have a higher metabolic rate than others. Highly active tissues, such as muscle, rapidly use oxygen to produce ATP, lowering the partial pressure of oxygen in the tissue to about 20 mm Hg. The partial pressure of oxygen inside capillaries is about 100 mm Hg, so the difference between the two becomes quite high, about 80 mm Hg. As a result, a greater number of oxygen molecules dissociate from hemoglobin and enter the tissues. The reverse is true of tissues, such as adipose (body fat), which have lower metabolic rates. Because less oxygen is used by these cells, the partial pressure of oxygen within such tissues remains relatively high, resulting in fewer oxygen molecules dissociating from hemoglobin and entering the tissue interstitial fluid. Although venous blood is said to be deoxygenated, some oxygen is still bound to hemoglobin in its red blood cells. This provides an oxygen reserve that can be used when tissues suddenly demand more oxygen.

Factors other than partial pressure also affect the oxygen–hemoglobin saturation/dissociation curve. For example, a higher temperature promotes hemoglobin and oxygen to dissociate faster, whereas a lower temperature inhibits dissociation (see Figure 2, middle). However, the human body tightly regulates temperature, so this factor may not affect gas exchange throughout the body. The exception to this is in highly active tissues, which may release a larger amount of energy than is given off as heat. As a result, oxygen readily dissociates from hemoglobin, which is a mechanism that helps to provide active tissues with more oxygen.

Certain hormones, such as androgens, epinephrine, thyroid hormones, and growth hormone, can affect the oxygen–hemoglobin saturation/disassociation curve by stimulating the production of a compound called 2,3-bisphosphoglycerate (BPG) by erythrocytes. BPG is a byproduct of glycolysis. Because erythrocytes do not contain mitochondria, glycolysis is the sole method by which these cells produce ATP. BPG promotes the disassociation of oxygen from hemoglobin. Therefore, the greater the concentration of BPG, the more readily oxygen dissociates from hemoglobin, despite its partial pressure.

The pH of the blood is another factor that influences the oxygen–hemoglobin saturation/dissociation curve (see Figure 2). The Bohr effect is a phenomenon that arises from the relationship between pH and oxygen’s affinity for hemoglobin: A lower, more acidic pH promotes oxygen dissociation from hemoglobin. In contrast, a higher, or more basic, pH inhibits oxygen dissociation from hemoglobin. The greater the amount of carbon dioxide in the blood, the more molecules that must be converted, which in turn generates hydrogen ions and thus lowers blood pH. Furthermore, blood pH may become more acidic when certain byproducts of cell metabolism, such as lactic acid, carbonic acid, and carbon dioxide, are released into the bloodstream.

Hemoglobin of the Fetus

The fetus has its own circulation with its own erythrocytes; however, it is dependent on the mother for oxygen. Blood is supplied to the fetus by way of the umbilical cord, which is connected to the placenta and separated from maternal blood by the chorion. The mechanism of gas exchange at the chorion is similar to gas exchange at the respiratory membrane. However, the partial pressure of oxygen is lower in the maternal blood in the placenta, at about 35 to 50 mm Hg, than it is in maternal arterial blood. The difference in partial pressures between maternal and fetal blood is not large, as the partial pressure of oxygen in fetal blood at the placenta is about 20 mm Hg. Therefore, there is not as much diffusion of oxygen into the fetal blood supply. The fetus’ hemoglobin overcomes this problem by having a greater affinity for oxygen than maternal hemoglobin (Figure 3). Both fetal and adult hemoglobin have four subunits, but two of the subunits of fetal hemoglobin have a different structure that causes fetal hemoglobin to have a greater affinity for oxygen than does adult hemoglobin.

[caption id="" align="aligncenter" width="430"]This graph shows the oxygen saturation versus the partial pressure of oxygen in fetal hemoglobin and adult hemoglobin. Figure 3. Oxygen-Hemoglobin Dissociation Curves in Fetus and Adult. Fetal hemoglobin has a greater affinity for oxygen than does adult hemoglobin.[/caption]

Carbon Dioxide Transport in the Blood

Carbon dioxide is transported by three major mechanisms. The first mechanism of carbon dioxide transport is by blood plasma, as some carbon dioxide molecules dissolve in the blood. The second mechanism is transport in the form of bicarbonate (HCO3–), which also dissolves in plasma. The third mechanism of carbon dioxide transport is similar to the transport of oxygen by erythrocytes (Figure 4).

[caption id="" align="aligncenter" width="435"]This figure shows how carbon dioxide is transported from the tissue to the red blood cell. Carbon Dioxide Transport Carbon dioxide is transported by three different methods: (a) in erythrocytes; (b) after forming carbonic acid (H2CO3 ), which is dissolved in plasma; (c) and in plasma.[/caption]

Dissolved Carbon Dioxide

Although carbon dioxide is not considered to be highly soluble in blood, a small fraction—about 7 to 10 percent—of the carbon dioxide that diffuses into the blood from the tissues dissolves in plasma. The dissolved carbon dioxide then travels in the bloodstream and when the blood reaches the pulmonary capillaries, the dissolved carbon dioxide diffuses across the respiratory membrane into the alveoli, where it is then exhaled during pulmonary ventilation.

Bicarbonate Buffer

A large fraction—about 70 percent—of the carbon dioxide molecules that diffuse into the blood is transported to the lungs as bicarbonate. Most bicarbonate is produced in erythrocytes after carbon dioxide diffuses into the capillaries, and subsequently into red blood cells. Carbonic anhydrase (CA) causes carbon dioxide and water to form carbonic acid (H2CO3), which dissociates into two ions: bicarbonate (HCO3–) and hydrogen (H+). The following formula depicts this reaction:

CO2 + H2O CA ↔ H2CO3↔H+ + HCO3−

Bicarbonate tends to build up in the erythrocytes, so that there is a greater concentration of bicarbonate in the erythrocytes than in the surrounding blood plasma. As a result, some of the bicarbonate will leave the erythrocytes and move down its concentration gradient into the plasma in exchange for chloride (Cl–) ions. This phenomenon is referred to as the chloride shift and occurs because by exchanging one negative ion for another negative ion, neither the electrical charge of the erythrocytes nor that of the blood is altered.

At the pulmonary capillaries, the chemical reaction that produced bicarbonate (shown above) is reversed, and carbon dioxide and water are the products. Much of the bicarbonate in the plasma re-enters the erythrocytes in exchange for chloride ions. Hydrogen ions and bicarbonate ions join to form carbonic acid, which is converted into carbon dioxide and water by carbonic anhydrase. Carbon dioxide diffuses out of the erythrocytes and into the plasma, where it can further diffuse across the respiratory membrane into the alveoli to be exhaled during pulmonary ventilation.

Carbaminohemoglobin

About 20 percent of carbon dioxide is bound by hemoglobin and is transported to the lungs. Carbon dioxide does not bind to iron as oxygen does; instead, carbon dioxide binds amino acid moieties on the globin portions of hemoglobin to form carbaminohemoglobin, which forms when hemoglobin and carbon dioxide bind. When hemoglobin is not transporting oxygen, it tends to have a bluish-purple tone to it, creating the darker maroon color typical of deoxygenated blood. The following formula depicts this reversible reaction:

CO2 + Hb ↔ HbCO2

Similar to the transport of oxygen by heme, the binding and dissociation of carbon dioxide to and from hemoglobin is dependent on the partial pressure of carbon dioxide. Because carbon dioxide is released from the lungs, blood that leaves the lungs and reaches body tissues has a lower partial pressure of carbon dioxide than is found in the tissues. As a result, carbon dioxide leaves the tissues because of its higher partial pressure, enters the blood, and then moves into red blood cells, binding to hemoglobin. In contrast, in the pulmonary capillaries, the partial pressure of carbon dioxide is high compared to within the alveoli. As a result, carbon dioxide dissociates readily from hemoglobin and diffuses across the respiratory membrane into the air.

In addition to the partial pressure of carbon dioxide, the oxygen saturation of hemoglobin and the partial pressure of oxygen in the blood also influence the affinity of hemoglobin for carbon dioxide. The Haldane effect is a phenomenon that arises from the relationship between the partial pressure of oxygen and the affinity of hemoglobin for carbon dioxide. Hemoglobin that is saturated with oxygen does not readily bind carbon dioxide. However, when oxygen is not bound to heme and the partial pressure of oxygen is low, hemoglobin readily binds to carbon dioxide. [caption id="attachment_2986" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of how oxygen is exchanged![/caption]  
]]> 2393 2352 9 0 22.6 Modifications in Respiratory Functions https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-6-modifications-in-respiratory-functions/ Thu, 13 Jul 2017 23:04:16 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2394

Learning Objectives

By the end of this section, you will be able to:
  • Define the terms hyperpnea and hyperventilation
  • Describe the effect of exercise on the respiratory system
  • Describe the effect of high altitude on the respiratory system
  • Discuss the process of acclimatization

At rest, the respiratory system performs its functions at a constant, rhythmic pace, as regulated by the respiratory centers of the brain. At this pace, ventilation provides sufficient oxygen to all the tissues of the body. However, there are times that the respiratory system must alter the pace of its functions in order to accommodate the oxygen demands of the body.

Hyperpnea

Hyperpnea is an increased depth and rate of ventilation to meet an increase in oxygen demand as might be seen in exercise or disease, particularly diseases that target the respiratory or digestive tracts. This does not significantly alter blood oxygen or carbon dioxide levels, but merely increases the depth and rate of ventilation to meet the demand of the cells. In contrast, hyperventilation is an increased ventilation rate that is independent of the cellular oxygen needs and leads to abnormally low blood carbon dioxide levels and high (alkaline) blood pH.

Interestingly, exercise does not cause hyperpnea as one might think. Muscles that perform work during exercise do increase their demand for oxygen, stimulating an increase in ventilation. However, hyperpnea during exercise appears to occur before a drop in oxygen levels within the muscles can occur. Therefore, hyperpnea must be driven by other mechanisms, either instead of or in addition to a drop in oxygen levels. The exact mechanisms behind exercise hyperpnea are not well understood, and some hypotheses are somewhat controversial. However, in addition to low oxygen, high carbon dioxide, and low pH levels, there appears to be a complex interplay of factors related to the nervous system and the respiratory centers of the brain.

First, a conscious decision to partake in exercise, or another form of physical exertion, results in a psychological stimulus that may trigger the respiratory centers of the brain to increase ventilation. In addition, the respiratory centers of the brain may be stimulated through the activation of motor neurons that innervate muscle groups that are involved in the physical activity. Finally, physical exertion stimulates proprioceptors, which are receptors located within the muscles, joints, and tendons, which sense movement and stretching; proprioceptors thus create a stimulus that may also trigger the respiratory centers of the brain. These neural factors are consistent with the sudden increase in ventilation that is observed immediately as exercise begins. Because the respiratory centers are stimulated by psychological, motor neuron, and proprioceptor inputs throughout exercise, the fact that there is also a sudden decrease in ventilation immediately after the exercise ends when these neural stimuli cease, further supports the idea that they are involved in triggering the changes of ventilation.

High Altitude Effects

An increase in altitude results in a decrease in atmospheric pressure. Although the proportion of oxygen relative to gases in the atmosphere remains at 21 percent, its partial pressure decreases (Table 4). As a result, it is more difficult for a body to achieve the same level of oxygen saturation at high altitude than at low altitude, due to lower atmospheric pressure. In fact, hemoglobin saturation is lower at high altitudes compared to hemoglobin saturation at sea level. For example, hemoglobin saturation is about 67 percent at 19,000 feet above sea level, whereas it reaches about 98 percent at sea level.

Partial Pressure of Oxygen at Different Altitudes (Table 4)
Example location Altitude (feet above sea level) Atmospheric pressure (mm Hg) Partial pressure of oxygen (mm Hg)
New York City, New York 0 760 159
Boulder, Colorado 5000 632 133
Aspen, Colorado 8000 565 118
Pike’s Peak, Colorado 14,000 447 94
Denali (Mt. McKinley), Alaska 20,000 350 73
Mt. Everest, Tibet 29,000 260 54

As you recall, partial pressure is extremely important in determining how much gas can cross the respiratory membrane and enter the blood of the pulmonary capillaries. A lower partial pressure of oxygen means that there is a smaller difference in partial pressures between the alveoli and the blood, so less oxygen crosses the respiratory membrane. As a result, fewer oxygen molecules are bound by hemoglobin. Despite this, the tissues of the body still receive a sufficient amount of oxygen during rest at high altitudes. This is due to two major mechanisms. First, the number of oxygen molecules that enter the tissue from the blood is nearly equal between sea level and high altitudes. At sea level, hemoglobin saturation is higher, but only a quarter of the oxygen molecules are actually released into the tissue. At high altitudes, a greater proportion of molecules of oxygen are released into the tissues. Secondly, at high altitudes, a greater amount of BPG is produced by erythrocytes, which enhances the dissociation of oxygen from hemoglobin. Physical exertion, such as skiing or hiking, can lead to altitude sickness due to the low amount of oxygen reserves in the blood at high altitudes. At sea level, there is a large amount of oxygen reserve in venous blood (even though venous blood is thought of as “deoxygenated”) from which the muscles can draw during physical exertion. Because the oxygen saturation is much lower at higher altitudes, this venous reserve is small, resulting in pathological symptoms of low blood oxygen levels. You may have heard that it is important to drink more water when traveling at higher altitudes than you are accustomed to. This is because your body will increase micturition (urination) at high altitudes to counteract the effects of lower oxygen levels. By removing fluids, blood plasma levels drop but not the total number of erythrocytes. In this way, the overall concentration of erythrocytes in the blood increases, which helps tissues obtain the oxygen they need.

Acute mountain sickness (AMS), or altitude sickness, is a condition that results from acute exposure to high altitudes due to a low partial pressure of oxygen at high altitudes. AMS typically can occur at 2400 meters (8000 feet) above sea level. AMS is a result of low blood oxygen levels, as the body has acute difficulty adjusting to the low partial pressure of oxygen. In serious cases, AMS can cause pulmonary or cerebral edema. Symptoms of AMS include nausea, vomiting, fatigue, lightheadedness, drowsiness, feeling disoriented, increased pulse, and nosebleeds. The only treatment for AMS is descending to a lower altitude; however, pharmacologic treatments and supplemental oxygen can improve symptoms. AMS can be prevented by slowly ascending to the desired altitude, allowing the body to acclimate, as well as maintaining proper hydration.

Acclimatization

Especially in situations where the ascent occurs too quickly, traveling to areas of high altitude can cause AMS. Acclimatization is the process of adjustment that the respiratory system makes due to chronic exposure to a high altitude. Over a period of time, the body adjusts to accommodate the lower partial pressure of oxygen. The low partial pressure of oxygen at high altitudes results in a lower oxygen saturation level of hemoglobin in the blood. In turn, the tissue levels of oxygen are also lower. As a result, the kidneys are stimulated to produce the hormone erythropoietin (EPO), which stimulates the production of erythrocytes, resulting in a greater number of circulating erythrocytes in an individual at a high altitude over a long period. With more red blood cells, there is more hemoglobin to help transport the available oxygen. Even though there is low saturation of each hemoglobin molecule, there will be more hemoglobin present, and therefore more oxygen in the blood. Over time, this allows the person to partake in physical exertion without developing AMS.

]]> 2394 2352 10 0 25.2 Gross Anatomy of Urine Transport https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2718 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2718

Learning Objectives

By the end of this section, you will be able to:
  • Identify the ureters, urinary bladder, and urethra, as well as their location, structure, histology, and function
  • Compare and contrast male and female urethras
  • Describe the micturition reflex
  • Describe voluntary and involuntary neural control of micturition

Rather than start with urine formation, this section will start with urine excretion. Urine is a fluid of variable composition that requires specialized structures to remove it from the body safely and efficiently. Blood is filtered, and the filtrate is transformed into urine at a relatively constant rate throughout the day. This processed liquid is stored until a convenient time for excretion. All structures involved in the transport and storage of the urine are large enough to be visible to the naked eye. This transport and storage system not only stores the waste, but it protects the tissues from damage due to the wide range of pH and osmolarity of the urine, prevents infection by foreign organisms, and for the male, provides reproductive functions.

Urethra

The urethra transports urine from the bladder to the outside of the body for disposal. The urethra is the only urologic organ that shows any significant anatomic difference between males and females; all other urine transport structures are identical (Figure 1).

[caption id="" align="aligncenter" width="560"]The top panel of this figure shows the organs in the female urinary system. Figure 1. Female and Male Urethras. The urethra transports urine from the bladder to the outside of the body. This image shows (a) a female urethra and (b) a male urethra.[/caption]

The urethra in both males and females begins inferior and central to the two ureteral openings forming the three points of a triangular-shaped area at the base of the bladder called the trigone (Greek tri- = “triangle” and the root of the word “trigonometry”). The urethra tracks posterior and inferior to the pubic symphysis (see Figure 1a). In both males and females, the proximal urethra is lined by transitional epithelium, whereas the terminal portion is a nonkeratinized, stratified squamous epithelium. In the male, pseudostratified columnar epithelium lines the urethra between these two cell types. Voiding is regulated by an involuntary autonomic nervous system-controlled internal urinary sphincter, consisting of smooth muscle and voluntary skeletal muscle that forms the external urinary sphincter below it.

Female Urethra

The external urethral orifice is embedded in the anterior vaginal wall inferior to the clitoris, superior to the vaginal opening (introitus), and medial to the labia minora. Its short length, about 4 cm, is less of a barrier to fecal bacteria than the longer male urethra and the best explanation for the greater incidence of UTI in women. Voluntary control of the external urethral sphincter is a function of the pudendal nerve. It arises in the sacral region of the spinal cord, traveling via the S2–S4 nerves of the sacral plexus.

Male Urethra

The male urethra passes through the prostate gland immediately inferior to the bladder before passing below the pubic symphysis (see Figure 1b). The length of the male urethra varies between men but averages 20 cm in length. It is divided into four regions: the preprostatic urethra, the prostatic urethra, the membranous urethra, and the spongy or penile urethra. The preprostatic urethra is very short and incorporated into the bladder wall. The prostatic urethra passes through the prostate gland. During sexual intercourse, it receives sperm via the ejaculatory ducts and secretions from the seminal vesicles. Paired Cowper’s glands (bulbourethral glands) produce and secrete mucus into the urethra to buffer urethral pH during sexual stimulation. The mucus neutralizes the usually acidic environment and lubricates the urethra, decreasing the resistance to ejaculation. The membranous urethra passes through the deep muscles of the perineum, where it is invested by the overlying urethral sphincters. The spongy urethra exits at the tip (external urethral orifice) of the penis after passing through the corpus spongiosum. Mucous glands are found along much of the length of the urethra and protect the urethra from extremes of urine pH. Innervation is the same in both males and females.

Bladder

The urinary bladder collects urine from both ureters (Figure 2). The bladder lies anterior to the uterus in females, posterior to the pubic bone and anterior to the rectum. During late pregnancy, its capacity is reduced due to compression by the enlarging uterus, resulting in increased frequency of urination. In males, the anatomy is similar, minus the uterus, and with the addition of the prostate inferior to the bladder. The bladder is partially retroperitoneal (outside the peritoneal cavity) with its peritoneal-covered “dome” projecting into the abdomen when the bladder is distended with urine.

[caption id="" align="aligncenter" width="500"]The left panel of this figure shows the cross section of the bladder and the major parts are labeled. The right panel shows a micrograph of the bladder. Figure 2. Bladder. (a) Anterior cross section of the bladder. (b) The detrusor muscle of the bladder (source: monkey tissue) LM × 448. (Micrograph provided by the Regents of the University of Michigan Medical School © 2012)[/caption]

The bladder is a highly distensible organ comprised of irregular crisscrossing bands of smooth muscle collectively called the detrusor muscle. The interior surface is made of transitional cellular epithelium that is structurally suited for the large volume fluctuations of the bladder. When empty, it resembles columnar epithelia, but when stretched, it “transitions” (hence the name) to a squamous appearance (see Figure 2). Volumes in adults can range from nearly zero to 500–600 mL.

The detrusor muscle contracts with significant force in the young. The bladder’s strength diminishes with age, but voluntary contractions of abdominal skeletal muscles can increase intra-abdominal pressure to promote more forceful bladder emptying. Such voluntary contraction is also used in forceful defecation and childbirth.

Micturition Reflex

Micturition is a less-often used, but proper term for urination or voiding. It results from an interplay of involuntary and voluntary actions by the internal and external urethral sphincters. When bladder volume reaches about 150 mL, an urge to void is sensed but is easily overridden. Voluntary control of urination relies on consciously preventing relaxation of the external urethral sphincter to maintain urinary continence. As the bladder fills, subsequent urges become harder to ignore. Ultimately, voluntary constraint fails with resulting incontinence, which will occur as bladder volume approaches 300 to 400 mL.

Normal micturition is a result of stretch receptors in the bladder wall that transmit nerve impulses to the sacral region of the spinal cord to generate a spinal reflex. The resulting parasympathetic neural outflow causes contraction of the detrusor muscle and relaxation of the involuntary internal urethral sphincter. At the same time, the spinal cord inhibits somatic motor neurons, resulting in the relaxation of the skeletal muscle of the external urethral sphincter. The micturition reflex is active in infants but with maturity, children learn to override the reflex by asserting external sphincter control, thereby delaying voiding (potty training). This reflex may be preserved even in the face of spinal cord injury that results in paraplegia or quadriplegia. However, relaxation of the external sphincter may not be possible in all cases, and therefore, periodic catheterization may be necessary for bladder emptying.

Nerves involved in the control of urination include the hypogastric, pelvic, and pudendal (Figure 3). Voluntary micturition requires an intact spinal cord and functional pudendal nerve arising from the sacral micturition center. Since the external urinary sphincter is voluntary skeletal muscle, actions by cholinergic neurons maintain contraction (and thereby continence) during filling of the bladder. At the same time, sympathetic nervous activity via the hypogastric nerves suppresses contraction of the detrusor muscle. With further bladder stretch, afferent signals traveling over sacral pelvic nerves activate parasympathetic neurons. This activates efferent neurons to release acetylcholine at the neuromuscular junctions, producing detrusor contraction and bladder emptying.

[caption id="" align="aligncenter" width="425"]This image shows the female urinary system and identifies the nerves that are important in this system. Figure 3. Nerves Innervating the Urinary System.[/caption]

Ureters

The kidneys and ureters are completely retroperitoneal, and the bladder has a peritoneal covering only over the dome. As urine is formed, it drains into the calyces of the kidney, which merge to form the funnel-shaped renal pelvis in the hilum of each kidney. The hilum narrows to become the ureter of each kidney. As urine passes through the ureter, it does not passively drain into the bladder but rather is propelled by waves of peristalsis. As the ureters enter the pelvis, they sweep laterally, hugging the pelvic walls. As they approach the bladder, they turn medially and pierce the bladder wall obliquely. This is important because it creates an one-way valve (a physiological sphincter rather than an anatomical sphincter) that allows urine into the bladder but prevents reflux of urine from the bladder back into the ureter. Children born lacking this oblique course of the ureter through the bladder wall are susceptible to “vesicoureteral reflux,” which dramatically increases their risk of serious UTI. Pregnancy also increases the likelihood of reflux and UTI.

The ureters are approximately 30 cm long. The inner mucosa is lined with transitional epithelium (Figure 4) and scattered goblet cells that secrete protective mucus. The muscular layer of the ureter consists of longitudinal and circular smooth muscles that create the peristaltic contractions to move the urine into the bladder without the aid of gravity. Finally, a loose adventitial layer composed of collagen and fat anchors the ureters between the parietal peritoneum and the posterior abdominal wall.

[caption id="" align="aligncenter" width="425"]A micrograph shows the lumen of the ureter. Figure 4. Ureter. Peristaltic contractions help to move urine through the lumen with contributions from fluid pressure and gravity. LM × 128. (Micrograph provided by the Regents of the University of Michigan Medical School © 2012)[/caption]

]]> 2718 2711 2 0 25.3 Gross Anatomy of the Kidney - 1203 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2724 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2724

Learning Objectives

By the end of this section, you will be able to:
  • Describe the external structure of the kidney, including its location, support structures, and covering
  • Identify the major internal divisions and structures of the kidney
  • Identify the major blood vessels associated with the kidney and trace the path of blood through the kidney
  • Compare and contrast the cortical and juxtamedullary nephrons
  • Name structures found in the cortex and medulla
  • Describe the physiological characteristics of the cortex and medulla
The kidneys lie on either side of the spine in the retroperitoneal space between the parietal peritoneum and the posterior abdominal wall, well protected by muscle, fat, and ribs. They are roughly the size of your fist, and the male kidney is typically a bit larger than the female kidney. The kidneys are well vascularized, receiving about 25 percent of the cardiac output at rest. [caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this video to learn about the TED (Technology, Entertainment, Design) Conference held in March 2011.[/caption]

External Anatomy

The left kidney is located at about the T12 to L3 vertebrae, whereas the right is lower due to slight displacement by the liver. Upper portions of the kidneys are somewhat protected by the eleventh and twelfth ribs (Figure 1). Each kidney weighs about 125–175 g in males and 115–155 g in females. They are about 11–14 cm in length, 6 cm wide, and 4 cm thick, and are directly covered by a fibrous capsule composed of dense, irregular connective tissue that helps to hold their shape and protect them. This capsule is covered by a shock-absorbing layer of adipose tissue called the renal fat pad, which in turn is encompassed by a tough renal fascia. The fascia and, to a lesser extent, the overlying peritoneum serve to firmly anchor the kidneys to the posterior abdominal wall in a retroperitoneal position.

[caption id="" align="aligncenter" width="425"]This image shows a human torso and shows the location of the kidneys within the torso. Figure 1. Kidneys. The kidneys are slightly protected by the ribs and are surrounded by fat for protection (not shown).[/caption]

On the superior aspect of each kidney is the adrenal gland. The adrenal cortex directly influences renal function through the production of the hormone aldosterone to stimulate sodium reabsorption.

Internal Anatomy

A frontal section through the kidney reveals an outer region called the renal cortex and an inner region called the medulla (Figure 2). The renal columns are connective tissue extensions that radiate downward from the cortex through the medulla to separate the most characteristic features of the medulla, the renal pyramids and renal papillae. The papillae are bundles of collecting ducts that transport urine made by nephrons to the calyces of the kidney for excretion. The renal columns also serve to divide the kidney into 6–8 lobes and provide a supportive framework for vessels that enter and exit the cortex. The pyramids and renal columns taken together constitute the kidney lobes.

[caption id="" align="aligncenter" width="475"]The left panel of this figure shows the location of the kidneys in the abdomen. The right panel shows the cross section of the kidney. Figure 2. Left Kidney.[/caption]

Renal Hilum

The renal hilum is the entry and exit site for structures servicing the kidneys: vessels, nerves, lymphatics, and ureters. The medial-facing hila are tucked into the sweeping convex outline of the cortex. Emerging from the hilum is the renal pelvis, which is formed from the major and minor calyxes in the kidney. The smooth muscle in the renal pelvis funnels urine via peristalsis into the ureter. The renal arteries form directly from the descending aorta, whereas the renal veins return cleansed blood directly to the inferior vena cava. The artery, vein, and renal pelvis are arranged in an anterior-to-posterior order.

Nephrons and Vessels

The renal artery first divides into segmental arteries, followed by further branching to form interlobar arteries that pass through the renal columns to reach the cortex (Figure 3). The interlobar arteries, in turn, branch into arcuate arteries, cortical radiate arteries, and then into afferent arterioles. The afferent arterioles service about 1.3 million nephrons in each kidney.

[caption id="" align="aligncenter" width="475"]This figure shows the network of blood vessels and the blood flow in the kidneys. Figure 3. Blood Flow in the Kidney.[/caption]

Nephrons are the “functional units” of the kidney; they cleanse the blood and balance the constituents of the circulation. The afferent arterioles form a tuft of high-pressure capillaries about 200 µm in diameter, the glomerulus. The rest of the nephron consists of a continuous sophisticated tubule whose proximal end surrounds the glomerulus in an intimate embrace—this is Bowman’s capsule. The glomerulus and Bowman’s capsule together form the renal corpuscle. As mentioned earlier, these glomerular capillaries filter the blood based on particle size. After passing through the renal corpuscle, the capillaries form a second arteriole, the efferent arteriole (Figure 4). These will next form a capillary network around the more distal portions of the nephron tubule, the peritubular capillaries and vasa recta, before returning to the venous system. As the glomerular filtrate progresses through the nephron, these capillary networks recover most of the solutes and water, and return them to the circulation. Since a capillary bed (the glomerulus) drains into a vessel that in turn forms a second capillary bed, the definition of a portal system is met. This is the only portal system in which an arteriole is found between the first and second capillary beds. (Portal systems also link the hypothalamus to the anterior pituitary, and the blood vessels of the digestive viscera to the liver.)

[caption id="" align="aligncenter" width="380"]This image shows the blood vessels and the direction of blood flow in the nephron. Figure 4. Blood Flow in the Nephron. The two capillary beds are clearly shown in this figure. The efferent arteriole is the connecting vessel between the glomerulus and the peritubular capillaries and vasa recta.[/caption]

Cortex

In a dissected kidney, it is easy to identify the cortex; it appears lighter in color compared to the rest of the kidney. All of the renal corpuscles as well as both the proximal convoluted tubules (PCTs) and distal convoluted tubules are found here. Some nephrons have a short loop of Henle that does not dip beyond the cortex. These nephrons are called cortical nephrons. About 15 percent of nephrons have long loops of Henle that extend deep into the medulla and are called juxtamedullary nephrons.

]]> 2724 2711 3 0 25.4 Microscopic Anatomy of the Kidney https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2730 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2730

Learning Objectives

By the end of this section, you will be able to:
  • Distinguish the histological differences between the renal cortex and medulla
  • Describe the structure of the filtration membrane
  • Identify the major structures and subdivisions of the renal corpuscles, renal tubules, and renal capillaries
  • Discuss the function of the peritubular capillaries and vasa recta
  • Identify the location of the juxtaglomerular apparatus and describe the cells that line it
  • Describe the histology of the proximal convoluted tubule, loop of Henle, distal convoluted tubule, and collecting ducts

The renal structures that conduct the essential work of the kidney cannot be seen by the naked eye. Only a light or electron microscope can reveal these structures. Even then, serial sections and computer reconstruction are necessary to give us a comprehensive view of the functional anatomy of the nephron and its associated blood vessels.

Nephrons: The Functional Unit

Nephrons take a simple filtrate of the blood and modify it into urine. Many changes take place in the different parts of the nephron before urine is created for disposal. The term forming urine will be used hereafter to describe the filtrate as it is modified into true urine. The principle task of the nephron population is to balance the plasma to homeostatic set points and excrete potential toxins in the urine. They do this by accomplishing three principle functions—filtration, reabsorption, and secretion. They also have additional secondary functions that exert control in three areas: blood pressure (via production of renin), red blood cell production (via the hormone EPO), and calcium absorption (via conversion of calcidiol into calcitriol, the active form of vitamin D).

Renal Corpuscle

As discussed earlier, the renal corpuscle consists of a tuft of capillaries called the glomerulus that is largely surrounded by Bowman’s (glomerular) capsule. The glomerulus is a high-pressure capillary bed between afferent and efferent arterioles. Bowman’s capsule surrounds the glomerulus to form a lumen, and captures and directs this filtrate to the PCT. The outermost part of Bowman’s capsule, the parietal layer, is a simple squamous epithelium. It transitions onto the glomerular capillaries in an intimate embrace to form the visceral layer of the capsule. Here, the cells are not squamous, but uniquely shaped cells (podocytes) extending finger-like arms (pedicels) to cover the glomerular capillaries (Figure 1). These projections interdigitate to form filtration slits, leaving small gaps between the digits to form a sieve. As blood passes through the glomerulus, 10 to 20 percent of the plasma filters between these sieve-like fingers to be captured by Bowman’s capsule and funneled to the PCT. Where the fenestrae (windows) in the glomerular capillaries match the spaces between the podocyte “fingers,” the only thing separating the capillary lumen and the lumen of Bowman’s capsule is their shared basement membrane (Figure 2). These three features comprise what is known as the filtration membrane. This membrane permits very rapid movement of filtrate from capillary to capsule though pores that are only 70 nm in diameter.

[caption id="" align="aligncenter" width="480"]The left panel of this figure shows an image of a podocyte. The right panel shows a tube-like structure that illustrates the filtration slits and the cell bodies. Figure 1. Podocytes. Podocytes interdigitate with structures called pedicels and filter substances in a way similar to fenestrations. In (a), the large cell body can be seen at the top right corner, with branches extending from the cell body. The smallest finger-like extensions are the pedicels. Pedicels on one podocyte always interdigitate with the pedicels of another podocyte. (b) This capillary has three podocytes wrapped around it.[/caption]
[caption id="" align="aligncenter" width="345"]The top panel of this figure shows a tube-like structure with the basement membrane and other parts labeled. Figure 2. Fenestrated Capillary. Fenestrations allow many substances to diffuse from the blood based primarily on size.[/caption]

The fenestrations prevent filtration of blood cells or large proteins, but allow most other constituents through. These substances cross readily if they are less than 4 nm in size and most pass freely up to 8 nm in size. An additional factor affecting the ability of substances to cross this barrier is their electric charge. The proteins associated with these pores are negatively charged, so they tend to repel negatively charged substances and allow positively charged substances to pass more readily. The basement membrane prevents filtration of medium-to-large proteins such as globulins. There are also mesangial cells in the filtration membrane that can contract to help regulate the rate of filtration of the glomerulus. Overall, filtration is regulated by fenestrations in capillary endothelial cells, podocytes with filtration slits, membrane charge, and the basement membrane between capillary cells. The result is the creation of a filtrate that does not contain cells or large proteins, and has a slight predominance of positively charged substances.

Lying just outside Bowman’s capsule and the glomerulus is the juxtaglomerular apparatus (JGA) (Figure 3). At the juncture where the afferent and efferent arterioles enter and leave Bowman’s capsule, the initial part of the distal convoluted tubule (DCT) comes into direct contact with the arterioles. The wall of the DCT at that point forms a part of the JGA known as the macula densa. This cluster of cuboidal epithelial cells monitors the fluid composition of fluid flowing through the DCT. In response to the concentration of Na+ in the fluid flowing past them, these cells release paracrine signals. They also have a single, nonmotile cilium that responds to the rate of fluid movement in the tubule. The paracrine signals released in response to changes in flow rate and Na+ concentration are adenosine triphosphate (ATP) and adenosine.

[caption id="" align="aligncenter" width="550"]The top panel of this image shows the cross section of the juxtaglomerular apparatus. The major parts are labeled. Figure 3. Juxtaglomerular Apparatus and Glomerulus. (a) The JGA allows specialized cells to monitor the composition of the fluid in the DCT and adjust the glomerular filtration rate. (b) This micrograph shows the glomerulus and surrounding structures. LM × 1540. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

A second cell type in this apparatus is the juxtaglomerular cell. This is a modified, smooth muscle cell lining the afferent arteriole that can contract or relax in response to ATP or adenosine released by the macula densa. Such contraction and relaxation regulate blood flow to the glomerulus. If the osmolarity of the filtrate is too high (hyperosmotic), the juxtaglomerular cells will contract, decreasing the glomerular filtration rate (GFR) so less plasma is filtered, leading to less urine formation and greater retention of fluid. This will ultimately decrease blood osmolarity toward the physiologic norm. If the osmolarity of the filtrate is too low, the juxtaglomerular cells will relax, increasing the GFR and enhancing the loss of water to the urine, causing blood osmolarity to rise. In other words, when osmolarity goes up, filtration and urine formation decrease and water is retained. When osmolarity goes down, filtration and urine formation increase and water is lost by way of the urine. The net result of these opposing actions is to keep the rate of filtration relatively constant. A second function of the macula densa cells is to regulate renin release from the juxtaglomerular cells of the afferent arteriole (Figure 4). Active renin is a protein comprised of 304 amino acids that cleaves several amino acids from angiotensinogen to produce angiotensin I. Angiotensin I is not biologically active until converted to angiotensin II by angiotensin-converting enzyme (ACE) from the lungs. Angiotensin II is a systemic vasoconstrictor that helps to regulate blood pressure by increasing it. Angiotensin II also stimulates the release of the steroid hormone aldosterone from the adrenal cortex. Aldosterone stimulates Na+ reabsorption by the kidney, which also results in water retention and increased blood pressure.

[caption id="" align="aligncenter" width="600"]This diagram shows the pathway of action of the renin-aldosterone-angiotensin system. An arrow in the center of the image shows the sequence of events that take place, and branching off from this arrow are indications of where in the body these events take place. Figure 4. Conversion of Angiotensin I to Angiotensin II. The enzyme renin converts the pro-enzyme angiotensin I; the lung-derived enzyme ACE converts angiotensin I into active angiotensin II.[/caption]

Proximal Convoluted Tubule (PCT)

Filtered fluid collected by Bowman’s capsule enters into the PCT. It is called convoluted due to its tortuous path. Simple cuboidal cells form this tubule with prominent microvilli on the luminal surface, forming a brush border. These microvilli create a large surface area to maximize the absorption and secretion of solutes (Na+, Cl–, glucose, etc.), the most essential function of this portion of the nephron. These cells actively transport ions across their membranes, so they possess a high concentration of mitochondria in order to produce sufficient ATP.

Loop of Henle

The descending and ascending portions of the loop of Henle (sometimes referred to as the nephron loop) are, of course, just continuations of the same tubule. They run adjacent and parallel to each other after having made a hairpin turn at the deepest point of their descent. The descending loop of Henle consists of an initial short, thick portion and long, thin portion, whereas the ascending loop consists of an initial short, thin portion followed by a long, thick portion. The descending thick portion consists of simple cuboidal epithelium similar to that of the PCT. The descending and ascending thin portions consists of simple squamous epithelium. As you will see later, these are important differences, since different portions of the loop have different permeabilities for solutes and water. The ascending thick portion consists of simple cuboidal epithelium similar to the DCT.

Distal Convoluted Tubule (DCT)

The DCT, like the PCT, is very tortuous and formed by simple cuboidal epithelium, but it is shorter than the PCT. These cells are not as active as those in the PCT; thus, there are fewer microvilli on the apical surface. However, these cells must also pump ions against their concentration gradient, so you will find of large numbers of mitochondria, although fewer than in the PCT.

Collecting Ducts

The collecting ducts are continuous with the nephron but not technically part of it. In fact, each duct collects filtrate from several nephrons for final modification. Collecting ducts merge as they descend deeper in the medulla to form about 30 terminal ducts, which empty at a papilla. They are lined with simple squamous epithelium with receptors for ADH. When stimulated by ADH, these cells will insert aquaporin channel proteins into their membranes, which as their name suggests, allow water to pass from the duct lumen through the cells and into the interstitial spaces to be recovered by the vasa recta. This process allows for the recovery of large amounts of water from the filtrate back into the blood. In the absence of ADH, these channels are not inserted, resulting in the excretion of water in the form of dilute urine. Most, if not all, cells of the body contain aquaporin molecules, whose channels are so small that only water can pass. At least 10 types of aquaporins are known in humans, and six of those are found in the kidney. The function of all aquaporins is to allow the movement of water across the lipid-rich, hydrophobic cell membrane (Figure 5).

[caption id="attachment_1836" align="aligncenter" width="500"]This figure shows an aquaporin water channel in the bilayer membrane with water molecules passing through. Figure 5. Aquaporin Water Channel. Positive charges inside the channel prevent the leakage of electrolytes across the cell membrane, while allowing water to move due to osmosis.[/caption]
]]> 2730 2711 4 0 25.6 Tubular Reabsorption https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2737 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2737

Learning Objectives

By the end of this section, you will be able to:
  • List specific transport mechanisms occurring in different parts of the nephron, including active transport, osmosis, facilitated diffusion, and passive electrochemical gradients
  • List the different membrane proteins of the nephron, including channels, transporters, and ATPase pumps
  • Compare and contrast passive and active tubular reabsorption
  • Explain why the differential permeability or impermeability of specific sections of the nephron tubules is necessary for urine formation
  • Describe how and where water, organic compounds, and ions are reabsorbed in the nephron
  • Explain the role of the loop of Henle, the vasa recta, and the countercurrent multiplication mechanisms in the concentration of urine
  • List the locations in the nephron where tubular secretion occurs

With up to 180 liters per day passing through the nephrons of the kidney, it is quite obvious that most of that fluid and its contents must be reabsorbed. That recovery occurs in the PCT, loop of Henle, DCT, and the collecting ducts (Table 5 and Figure 1). Various portions of the nephron differ in their capacity to reabsorb water and specific solutes. While much of the reabsorption and secretion occur passively based on concentration gradients, the amount of water that is reabsorbed or lost is tightly regulated. This control is exerted directly by ADH and aldosterone, and indirectly by renin. Most water is recovered in the PCT, loop of Henle, and DCT. About 10 percent (about 18 L) reaches the collecting ducts. The collecting ducts, under the influence of ADH, can recover almost all of the water passing through them, in cases of dehydration, or almost none of the water, in cases of over-hydration.

[caption id="" align="aligncenter" width="280"]This diagram shows the different ions and chemicals that are secreted and reabsorbed along the nephron. Arrows show the direction of the movement of the substance. Figure 1. Locations of Secretion and Reabsorption in the Nephron.[/caption]
Substances Secreted or Reabsorbed in the Nephron and Their Locations (Table 5)
Substance PCT Loop of Henle DCT Collecting ducts
Glucose Almost 100 percent reabsorbed; secondary active transport with Na+
Oligopeptides, proteins, amino acids Almost 100 percent reabsorbed; symport with Na+
Vitamins Reabsorbed
Lactate Reabsorbed
Creatinine Secreted
Urea 50 percent reabsorbed by diffusion; also secreted Secretion, diffusion in descending limb Reabsorption in medullary collecting ducts; diffusion
Sodium 65 percent actively reabsorbed 25 percent reabsorbed in thick ascending limb; active transport 5 percent reabsorbed; active 5 percent reabsorbed, stimulated by aldosterone; active
Chloride Reabsorbed, symport with Na+, diffusion Reabsorbed in thin and thick ascending limb; diffusion in ascending limb Reabsorbed; diffusion Reabsorbed; symport
Water 67 percent reabsorbed osmotically with solutes 15 percent reabsorbed in descending limb; osmosis 8 percent reabsorbed if ADH; osmosis Variable amounts reabsorbed, controlled by ADH, osmosis
Bicarbonate 80–90 percent symport reabsorption with Na+ Reabsorbed, symport with Na+ and antiport with Cl–; in ascending limb Reabsorbed antiport with Cl–
H+ Secreted; diffusion Secreted; active Secreted; active
NH4+ Secreted; diffusion Secreted; diffusion Secreted; diffusion
HCO3– Reabsorbed; diffusion Reabsorbed; diffusion in ascending limb Reabsorbed; diffusion Reabsorbed; antiport with Na+
Some drugs Secreted Secreted; active Secreted; active
Potassium 65 percent reabsorbed; diffusion 20 percent reabsorbed in thick ascending limb; symport Secreted; active Secretion controlled by aldosterone; active
Calcium Reabsorbed; diffusion Reabsorbed in thick ascending limb; diffusion Reabsorbed if parathyroid hormone present; active
Magnesium Reabsorbed; diffusion Reabsorbed in thick ascending limb; diffusion Reabsorbed
Phosphate 85 percent reabsorbed, inhibited by parathyroid hormone, diffusion Reabsorbed; diffusion

Mechanisms of Recovery

Mechanisms by which substances move across membranes for reabsorption or secretion include active transport, diffusion, facilitated diffusion, secondary active transport, and osmosis. These were discussed in an earlier chapter, and you may wish to review them.

Active transport utilizes energy, usually the energy found in a phosphate bond of ATP, to move a substance across a membrane from a low to a high concentration. It is very specific and must have an appropriately shaped receptor for the substance to be transported. An example would be the active transport of Na+ out of a cell and K+ into a cell by the Na+/K+ pump. Both ions are moved in opposite directions from a lower to a higher concentration.

Simple diffusion moves a substance from a higher to a lower concentration down its concentration gradient. It requires no energy and only needs to be soluble.

Facilitated diffusion is similar to diffusion in that it moves a substance down its concentration gradient. The difference is that it requires specific membrane receptors or channel proteins for movement. The movement of glucose and, in certain situations, Na+ ions, is an example of facilitated diffusion. In some cases of facilitated diffusion, two different substances share the same channel protein port; these mechanisms are described by the terms symport and antiport.

Symport mechanisms move two or more substances in the same direction at the same time, whereas antiport mechanisms move two or more substances in opposite directions across the cell membrane. Both mechanisms may utilize concentration gradients maintained by ATP pumps. This is a mechanism described by the term “secondary active transport.” For example, a Na+ ATPase pump on the basilar membrane of a cell may constantly pump Na+ out of a cell, maintaining a strong electrochemical gradient. On the opposite (apical) surface, a Na+/glucose symport protein channel assists both Na+ and glucose into the cell as Na+ moves down the concentration gradient created by the basilar Na+ ATPase pumps. The glucose molecule then diffuses across the basal membrane by facilitated diffusion into the interstitial space and from there into peritubular capillaries.

Most of the Ca++, Na+, glucose, and amino acids must be reabsorbed by the nephron to maintain homeostatic plasma concentrations. Other substances, such as urea, K+, ammonia (NH3), creatinine, and some drugs are secreted into the filtrate as waste products. Acid–base balance is maintained through actions of the lungs and kidneys: The lungs rid the body of H+, whereas the kidneys secrete or reabsorb H+ and HCO3– (Table 6). In the case of urea, about 50 percent is passively reabsorbed by the PCT. More is recovered by in the collecting ducts as needed. ADH induces the insertion of urea transporters and aquaporin channel proteins.

Substances Filtered and Reabsorbed by the Kidney per 24 Hours (Table 6)
Substance Amount filtered (grams) Amount reabsorbed (grams) Amount in urine (grams)
Water 180 L 179 L 1 L
Proteins 10–20 10–20 0
Chlorine 630 625 5
Sodium 540 537 3
Bicarbonate 300 299.7 0.3
Glucose 180 180 0
Urea 53 28 25
Potassium 28 24 4
Uric acid 8.5 7.7 0.8
Creatinine 1.4 0 1.4

Reabsorption and Secretion in the PCT

The renal corpuscle filters the blood to create a filtrate that differs from blood mainly in the absence of cells and large proteins. From this point to the ends of the collecting ducts, the filtrate or forming urine is undergoing modification through secretion and reabsorption before true urine is produced. The first point at which the forming urine is modified is in the PCT. Here, some substances are reabsorbed, whereas others are secreted. Note the use of the term “reabsorbed.” All of these substances were “absorbed” in the digestive tract—99 percent of the water and most of the solutes filtered by the nephron must be reabsorbed. Water and substances that are reabsorbed are returned to the circulation by the peritubular and vasa recta capillaries. It is important to understand the difference between the glomerulus and the peritubular and vasa recta capillaries. The glomerulus has a relatively high pressure inside its capillaries and can sustain this by dilating the afferent arteriole while constricting the efferent arteriole. This assures adequate filtration pressure even as the systemic blood pressure varies. Movement of water into the peritubular capillaries and vasa recta will be influenced primarily by osmolarity and concentration gradients. Sodium is actively pumped out of the PCT into the interstitial spaces between cells and diffuses down its concentration gradient into the peritubular capillary. As it does so, water will follow passively to maintain an isotonic fluid environment inside the capillary. This is called obligatory water reabsorption, because water is “obliged” to follow the Na+ (Figure 2).

[caption id="" align="aligncenter" width="535"]This diagram shows the different substances that are secreted and reabsorbed by the proximal collecting tubule. Arrows show the direction of the movement of the substance. Figure 2. Substances Reabsorbed and Secreted by the PCT.[/caption]

More substances move across the membranes of the PCT than any other portion of the nephron. Many of these substances (amino acids and glucose) use symport mechanisms for transport along with Na+. Antiport, active transport, diffusion, and facilitated diffusion are additional mechanisms by which substances are moved from one side of a membrane to the other. Recall that cells have two surfaces: apical and basal. The apical surface is the one facing the lumen or open space of a cavity or tube, in this case, the inside of the PCT. The basal surface of the cell faces the connective tissue base to which the cell attaches (basement membrane) or the cell membrane closer to the basement membrane if there is a stratified layer of cells. In the PCT, there is a single layer of simple cuboidal endothelial cells against the basement membrane. The numbers and particular types of pumps and channels vary between the apical and basilar surfaces (Table 7). A few of the substances that are transported with Na+ (symport mechanism) on the apical membrane include Cl–, Ca++, amino acids, glucose, and PO43−. Sodium is actively exchanged for K+ using ATP on the basal membrane. Most of the substances transported by a symport mechanism on the apical membrane are transported by facilitated diffusion on the basal membrane. At least three ions, K+, Ca++, and Mg++, diffuse laterally between adjacent cell membranes (transcellular).

Reabsorption of Major Solutes by the PCT (Table 7)
Basal membrane Apical membrane
Active transport Symport with Na+
Na+ (exchange for K+) K+
Facilitated diffusion Cl–
K+ Ca++
Cl– Mg++
Ca++ HCO3–
HCO3– PO43−
PO43− Amino acids
Amino acids Glucose
Glucose Fructose
Fructose Galactose
Galactose Lactate
Lactate Succinate
Succinate Citrate
Citrate Diffusion between nephron cells
K+
Ca++
Mg++

About 67 percent of the water, Na+, and K+ entering the nephron is reabsorbed in the PCT and returned to the circulation. Almost 100 percent of glucose, amino acids, and other organic substances such as vitamins are normally recovered here. Some glucose may appear in the urine if circulating glucose levels are high enough that all the glucose transporters in the PCT are saturated, so that their capacity to move glucose is exceeded (transport maximum, or Tm). In men, the maximum amount of glucose that can be recovered is about 375 mg/min, whereas in women, it is about 300 mg/min. This recovery rate translates to an arterial concentration of about 200 mg/dL. Though an exceptionally high sugar intake might cause sugar to appear briefly in the urine, the appearance of glycosuria usually points to type I or II diabetes mellitus. The transport of glucose from the lumen of the PCT to the interstitial space is similar to the way it is absorbed by the small intestine. Both glucose and Na+ bind simultaneously to the same symport proteins on the apical surface of the cell to be transported in the same direction, toward the interstitial space. Sodium moves down its electrochemical and concentration gradient into the cell and takes glucose with it. Na+ is then actively pumped out of the cell at the basal surface of the cell into the interstitial space. Glucose leaves the cell to enter the interstitial space by facilitated diffusion. The energy to move glucose comes from the Na+/K+ ATPase that pumps Na+ out of the cell on the basal surface. Fifty percent of Cl– and variable quantities of Ca++, Mg++, and HPO42− are also recovered in the PCT.

Recovery of bicarbonate (HCO3–) is vital to the maintenance of acid–base balance, since it is a very powerful and fast-acting buffer. An important enzyme is used to catalyze this mechanism: carbonic anhydrase (CA). This same enzyme and reaction is used in red blood cells in the transportation of CO2, in the stomach to produce hydrochloric acid, and in the pancreas to produce HCO3– to buffer acidic chyme from the stomach. In the kidney, most of the CA is located within the cell, but a small amount is bound to the brush border of the membrane on the apical surface of the cell. In the lumen of the PCT, HCO3– combines with hydrogen ions to form carbonic acid (H2CO3). This is enzymatically catalyzed into CO2 and water, which diffuse across the apical membrane into the cell. Water can move osmotically across the lipid bilayer membrane due to the presence of aquaporin water channels. Inside the cell, the reverse reaction occurs to produce bicarbonate ions (HCO3–). These bicarbonate ions are cotransported with Na+ across the basal membrane to the interstitial space around the PCT (Figure 3). At the same time this is occurring, a Na+/H+ antiporter excretes H+ into the lumen, while it recovers Na+. Note how the hydrogen ion is recycled so that bicarbonate can be recovered. Also, note that a Na+ gradient is created by the Na+/K+ pump.

HCO3−+ H+ ↔ H2CO3 ↔ CO2 + H2O

The significant recovery of solutes from the PCT lumen to the interstitial space creates an osmotic gradient that promotes water recovery. As noted before, water moves through channels created by the aquaporin proteins. These proteins are found in all cells in varying amounts and help regulate water movement across membranes and through cells by creating a passageway across the hydrophobic lipid bilayer membrane. Changing the number of aquaporin proteins in membranes of the collecting ducts also helps to regulate the osmolarity of the blood. The movement of many positively charged ions also creates an electrochemical gradient. This charge promotes the movement of negative ions toward the interstitial spaces and the movement of positive ions toward the lumen.

[caption id="" align="aligncenter" width="400"]This diagram shows the process of reabsorption of bicarbonate by the proximal collecting tubule. Figure 3. Reabsorption of Bicarbonate from the PCT.[/caption]

Reabsorption and Secretion in the Loop of Henle

The loop of Henle consists of two sections: thick and thin descending and thin and thick ascending sections. The loops of cortical nephrons do not extend into the renal medulla very far, if at all. Juxtamedullary nephrons have loops that extend variable distances, some very deep into the medulla. The descending and ascending portions of the loop are highly specialized to enable recovery of much of the Na+ and water that were filtered by the glomerulus. As the forming urine moves through the loop, the osmolarity will change from isosmotic with blood (about 278–300 mOsmol/kg) to both a very hypertonic solution of about 1200 mOsmol/kg and a very hypotonic solution of about 100 mOsmol/kg. These changes are accomplished by osmosis in the descending limb and active transport in the ascending limb. Solutes and water recovered from these loops are returned to the circulation by way of the vasa recta.

Descending Loop

The majority of the descending loop is comprised of simple squamous epithelial cells; to simplify the function of the loop, this discussion focuses on these cells. These membranes have permanent aquaporin channel proteins that allow unrestricted movement of water from the descending loop into the surrounding interstitium as osmolarity increases from about 300 mOsmol/kg to about 1200 mOsmol/kg. This increase results in reabsorption of up to 15 percent of the water entering the nephron. Modest amounts of urea, Na+, and other ions are also recovered here.

Most of the solutes that were filtered in the glomerulus have now been recovered along with a majority of water, about 82 percent. As the forming urine enters the ascending loop, major adjustments will be made to the concentration of solutes to create what you perceive as urine.

Ascending Loop

The ascending loop is made of very short thin and longer thick portions. Once again, to simplify the function, this section only considers the thick portion. The thick portion is lined with simple cuboidal epithelium without a brush border. It is completely impermeable to water due to the absence of aquaporin proteins, but ions, mainly Na+, are actively pumped out of the loop by large quantities of the Na+/K+ ATPase pump. This has two significant effects: Removal of Na+ while retaining water leads to a hypotonic filtrate by the time it reaches the DCT; pumping Na+ into the interstitial space contributes to the hyperosmotic environment in the kidney medulla.

The Na+/K+ ATPase pumps in the basal membrane create an electrochemical gradient, allowing reabsorption of Cl– by Na+/Cl– symporters in the apical membrane. At the same time that Na+ is actively pumped from the basal side of the cell into the interstitial fluid, Cl– follows the Na+ from the lumen into the interstitial fluid by a paracellular route between cells through leaky tight junctions. These are found between cells of the ascending loop, where they allow certain solutes to move according to their concentration gradient. Most of the K+ that enters the cell via symporters returns to the lumen (down its concentration gradient) through leaky channels in the apical membrane. Note the environment now created in the interstitial space: With the “back door exiting” K+, there is one Na+ and two Cl– ions left in the interstitium surrounding the ascending loop. Therefore, in comparison to the lumen of the loop, the interstitial space is now a negatively charged environment. This negative charge attracts cations (Na+, K+, Ca++, and Mg++) from the lumen via a paracellular route to the interstitial space and vasa recta.

Countercurrent Multiplier System

The structure of the loop of Henle and associated vasa recta create a countercurrent multiplier system (Figure 4). The countercurrent term comes from the fact that the descending and ascending loops are next to each other and their fluid flows in opposite directions (countercurrent). The multiplier term is due to the action of solute pumps that increase (multiply) the concentrations of urea and Na+ deep in the medulla.

[caption id="" align="aligncenter" width="450"]The left panel of this image shows the location of the loop of Henle. The right panel shows the interstitial osmolality and the exchange of sodium and chloride ions, as well as water and urea. Figure 4. Countercurrent Multiplier System.[/caption]

As discussed above, the ascending loop has many Na+ pumps that actively pump Na+ out of the forming urine into the interstitial spaces. In addition, collecting ducts have urea pumps that actively pump urea into the interstitial spaces. This results in the recovery of Na+ to the circulation via the vasa recta and creates a high osmolar environment in the depths of the medulla.

Ammonia (NH3) is a toxic byproduct of protein metabolism. It is formed as amino acids are deaminated by liver hepatocytes. That means that the amine group, NH2, is removed from amino acids as they are broken down. Most of the resulting ammonia is converted into urea by liver hepatocytes. Urea is not only less toxic but is utilized to aid in the recovery of water by the loop of Henle and collecting ducts. At the same time that water is freely diffusing out of the descending loop through aquaporin channels into the interstitial spaces of the medulla, urea freely diffuses into the lumen of the descending loop as it descends deeper into the medulla, much of it to be reabsorbed from the forming urine when it reaches the collecting duct. Thus, the movement of Na+ and urea into the interstitial spaces by these mechanisms creates the hyperosmotic environment of the medulla. The net result of this countercurrent multiplier system is to recover both water and Na+ in the circulation.

The amino acid glutamine can be deaminated by the kidney. As NH2 from the amino acid is converted into NH3 and pumped into the lumen of the PCT, Na+ and HCO3– are excreted into the interstitial fluid of the renal pyramid via a symport mechanism. When this process occurs in the cells of the PCT, the added benefit is a net loss of a hydrogen ion (complexed to ammonia to form the weak acid NH4+) in the urine and a gain of a bicarbonate ion (HCO3–) in the blood. Ammonia and bicarbonate are exchanged in a one-to-one ratio. This exchange is yet another means by which the body can buffer and excrete acid. The presence of aquaporin channels in the descending loop allows prodigious quantities of water to leave the loop and enter the hyperosmolar interstitium of the pyramid, where it is returned to the circulation by the vasa recta. As the loop turns to become the ascending loop, there is an absence of aquaporin channels, so water cannot leave the loop. However, in the basal membrane of cells of the thick ascending loop, ATPase pumps actively remove Na+ from the cell. A Na+/K+/2Cl– symporter in the apical membrane passively allows these ions to enter the cell cytoplasm from the lumen of the loop down a concentration gradient created by the pump. This mechanism works to dilute the fluid of the ascending loop ultimately to approximately 50–100 mOsmol/L.

At the transition from the DCT to the collecting duct, about 20 percent of the original water is still present and about 10 percent of the sodium. If no other mechanism for water reabsorption existed, about 20–25 liters of urine would be produced. Now consider what is happening in the adjacent capillaries, the vasa recta. They are recovering both solutes and water at a rate that preserves the countercurrent multiplier system. In general, blood flows slowly in capillaries to allow time for exchange of nutrients and wastes. In the vasa recta particularly, this rate of flow is important for two additional reasons. The flow must be slow to allow blood cells to lose and regain water without either crenating or bursting. Second, a rapid flow would remove too much Na+ and urea, destroying the osmolar gradient that is necessary for the recovery of solutes and water. Thus, by flowing slowly to preserve the countercurrent mechanism, as the vasa recta descend, Na+ and urea are freely able to enter the capillary, while water freely leaves; as they ascend, Na+ and urea are secreted into the surrounding medulla, while water reenters and is removed. Watch this CrashCourse video to learn more about reabsorption!

Reabsorption and Secretion in the Distal Convoluted Tubule

Approximately 80 percent of filtered water has been recovered by the time the dilute forming urine enters the DCT. The DCT will recover another 10–15 percent before the forming urine enters the collecting ducts. Aldosterone increases the amount of Na+/K+ ATPase in the basal membrane of the DCT and collecting duct. The movement of Na+ out of the lumen of the collecting duct creates a negative charge that promotes the movement of Cl– out of the lumen into the interstitial space by a paracellular route across tight junctions. Peritubular capillaries receive the solutes and water, returning them to the circulation.

Cells of the DCT also recover Ca++ from the filtrate. Receptors for parathyroid hormone (PTH) are found in DCT cells and when bound to PTH, induce the insertion of calcium channels on their luminal surface. The channels enhance Ca++ recovery from the forming urine. In addition, as Na+ is pumped out of the cell, the resulting electrochemical gradient attracts Ca++ into the cell. Finally, calcitriol (1,25 dihydroxyvitamin D, the active form of vitamin D) is very important for calcium recovery. It induces the production of calcium-binding proteins that transport Ca++ into the cell. These binding proteins are also important for the movement of calcium inside the cell and aid in exocytosis of calcium across the basolateral membrane. Any Ca++ not reabsorbed at this point is lost in the urine.

Collecting Ducts and Recovery of Water

Solutes move across the membranes of the collecting ducts, which contain two distinct cell types, principal cells and intercalated cells. A principal cell possesses channels for the recovery or loss of sodium and potassium. An intercalated cell secretes or absorbs acid or bicarbonate. As in other portions of the nephron, there is an array of micromachines (pumps and channels) on display in the membranes of these cells.

Regulation of urine volume and osmolarity are major functions of the collecting ducts. By varying the amount of water that is recovered, the collecting ducts play a major role in maintaining the body’s normal osmolarity. If the blood becomes hyperosmotic, the collecting ducts recover more water to dilute the blood; if the blood becomes hyposmotic, the collecting ducts recover less of the water, leading to concentration of the blood. Another way of saying this is: If plasma osmolarity rises, more water is recovered and urine volume decreases; if plasma osmolarity decreases, less water is recovered and urine volume increases. This function is regulated by the posterior pituitary hormone ADH (vasopressin). With mild dehydration, plasma osmolarity rises slightly. This increase is detected by osmoreceptors in the hypothalamus, which stimulates the release of ADH from the posterior pituitary. If plasma osmolarity decreases slightly, the opposite occurs.

When stimulated by ADH, aquaporin channels are inserted into the apical membrane of principal cells, which line the collecting ducts. As the ducts descend through the medulla, the osmolarity surrounding them increases (due to the countercurrent mechanisms described above). If aquaporin water channels are present, water will be osmotically pulled from the collecting duct into the surrounding interstitial space and into the peritubular capillaries. Therefore, the final urine will be more concentrated. If less ADH is secreted, fewer aquaporin channels are inserted and less water is recovered, resulting in dilute urine. By altering the number of aquaporin channels, the volume of water recovered or lost is altered. This, in turn, regulates the blood osmolarity, blood pressure, and osmolarity of the urine.

As Na+ is pumped from the forming urine, water is passively recaptured for the circulation; this preservation of vascular volume is critically important for the maintenance of a normal blood pressure. Aldosterone is secreted by the adrenal cortex in response to angiotensin II stimulation. As an extremely potent vasoconstrictor, angiotensin II functions immediately to increase blood pressure. By also stimulating aldosterone production, it provides a longer-lasting mechanism to support blood pressure by maintaining vascular volume (water recovery).

In addition to receptors for ADH, principal cells have receptors for the steroid hormone aldosterone. While ADH is primarily involved in the regulation of water recovery, aldosterone regulates Na+ recovery. Aldosterone stimulates principal cells to manufacture luminal Na+ and K+ channels as well as Na+/K+ ATPase pumps on the basal membrane of the cells. When aldosterone output increases, more Na+ is recovered from the forming urine and water follows the Na+ passively. As the pump recovers Na+ for the body, it is also pumping K+ into the forming urine, since the pump moves K+ in the opposite direction. When aldosterone decreases, more Na+ remains in the forming urine and more K+ is recovered in the circulation. Symport channels move Na+ and Cl– together. Still other channels in the principal cells secrete K+ into the collecting duct in direct proportion to the recovery of Na+.

Intercalated cells play significant roles in regulating blood pH. Intercalated cells reabsorb K+ and HCO3– while secreting H+. This function lowers the acidity of the plasma while increasing the acidity of the urine.

]]> 2737 2711 6 0 26.1 Body Fluids and Fluid Compartments https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2748 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2748

Learning Objectives

By the end of this section, you will be able to:
  • Explain the importance of water in the body
  • Contrast the composition of the intracellular fluid with that of the extracellular fluid
  • Explain the importance of protein channels in the movement of solutes
  • Identify the causes and symptoms of edema

The chemical reactions of life take place in aqueous solutions. The dissolved substances in a solution are called solutes. In the human body, solutes vary in different parts of the body, but may include proteins—including those that transport lipids, carbohydrates, and, very importantly, electrolytes. Often in medicine, a mineral dissociated from a salt that carries an electrical charge (an ion) is called and electrolyte. For instance, sodium ions (Na+) and chloride ions (Cl-) are often referred to as electrolytes.

In the body, water moves through semi-permeable membranes of cells and from one compartment of the body to another by a process called osmosis. Osmosis is basically the diffusion of water from regions of higher concentration to regions of lower concentration, along an osmotic gradient across a semi-permeable membrane. As a result, water will move into and out of cells and tissues, depending on the relative concentrations of the water and solutes found there. An appropriate balance of solutes inside and outside of cells must be maintained to ensure normal function.

Body Water Content

Human beings are mostly water, ranging from about 75 percent of body mass in infants to about 50–60 percent in adult men and women, to as low as 45 percent in old age. The percent of body water changes with development, because the proportions of the body given over to each organ and to muscles, fat, bone, and other tissues change from infancy to adulthood (Figure 1). Your brain and kidneys have the highest proportions of water, which composes 80–85 percent of their masses. In contrast, teeth have the lowest proportion of water, at 8–10 percent.

[caption id="" align="aligncenter" width="450"]This illustration shows a silhouette of a human body with various organs highlighted. The percent of water contained in each organ is given. The brain typically contains 80% to 85% water, teeth contain 8% to 10% water, a single lung contains 75% to 80% water, the heart contains 75% to 80% water, the bones contain 20% to 25% water, the liver contains 70% to 75% water, the kidneys contain 80% to 85% water, the skin contains 70% to 75% water and the muscles also contain 70% to 75% water. Figure 1. Water Content of the Body’s Organs and Tissues. Water content varies in different body organs and tissues, from as little as 8 percent in the teeth to as much as 85 percent in the brain.[/caption]

Fluid Compartments

Body fluids can be discussed in terms of their specific fluid compartment, a location that is largely separate from another compartment by some form of a physical barrier. The intracellular fluid (ICF) compartment is the system that includes all fluid enclosed in cells by their plasma membranes. Extracellular fluid (ECF) surrounds all cells in the body. Extracellular fluid has two primary constituents: the fluid component of the blood (called plasma) and the interstitial fluid (IF) that surrounds all cells not in the blood (Figure 2).

[caption id="" align="aligncenter" width="380"]This diagram shows a small blood vessel surrounded by several body cells. The fluid between the body cells is the interstitial fluid (IF), which is a type of extracellular fluid (ECF). The fluid in the blood vessel is also an example of extracellular fluid. The fluid in the cytoplasm of each body cell is intracellular fluid, or ICF. Figure 2. Fluid Compartments in the Human Body. The intracellular fluid (ICF) is the fluid within cells. The interstitial fluid (IF) is part of the extracellular fluid (ECF) between the cells. Blood plasma is the second part of the ECF. Materials travel between cells and the plasma in capillaries through the IF.[/caption]

Intracellular Fluid

The ICF lies within cells and is the principal component of the cytosol/cytoplasm. The ICF makes up about 60 percent of the total water in the human body, and in an average-size adult male, the ICF accounts for about 25 liters (seven gallons) of fluid (Figure 3). This fluid volume tends to be very stable, because the amount of water in living cells is closely regulated. If the amount of water inside a cell falls to a value that is too low, the cytosol becomes too concentrated with solutes to carry on normal cellular activities; if too much water enters a cell, the cell may burst and be destroyed.

[caption id="" align="aligncenter" width="380"]This pie chart shows that about 55% of water in the human body is intracellular fluid. About 30% of the water in the human body is interstitial fluid. Most of the remaining 15% of water is plasma, along with a small percentage labeled “other fluid”. Figure 3. A Pie Graph Showing the Proportion of Total Body Fluid in Each of the Body’s Fluid Compartments. Most of the water in the body is intracellular fluid. The second largest volume is the interstitial fluid, which surrounds cells that are not blood cells.[/caption]

Extracellular Fluid

The ECF accounts for the other one-third of the body’s water content. Approximately 20 percent of the ECF is found in plasma. Plasma travels through the body in blood vessels and transports a range of materials, including blood cells, proteins (including clotting factors and antibodies), electrolytes, nutrients, gases, and wastes. Gases, nutrients, and waste materials travel between capillaries and cells through the IF. Cells are separated from the IF by a selectively permeable cell membrane that helps regulate the passage of materials between the IF and the interior of the cell.

The body has other water-based ECF. These include the cerebrospinal fluid that bathes the brain and spinal cord, lymph, the synovial fluid in joints, the pleural fluid in the pleural cavities, the pericardial fluid in the cardiac sac, the peritoneal fluid in the peritoneal cavity, and the aqueous humor of the eye. Because these fluids are outside of cells, these fluids are also considered components of the ECF compartment.

Composition of Body Fluids

The compositions of the two components of the ECF—plasma and IF—are more similar to each other than either is to the ICF (Figure 4). Blood plasma has high concentrations of sodium, chloride, bicarbonate, and protein. The IF has high concentrations of sodium, chloride, and bicarbonate, but a relatively lower concentration of protein. In contrast, the ICF has elevated amounts of potassium, phosphate, magnesium, and protein. Overall, the ICF contains high concentrations of potassium and phosphate (HPO42−HPO42−), whereas both plasma and the ECF contain high concentrations of sodium and chloride.

[caption id="" align="aligncenter" width="380"]This bar graph shows the concentration of several ions and proteins in intracellular fluid, interstitial fluid and plasma. The ions and proteins are categories on the X axis . The Y axis shows concentration, in milliequivalents per liter, ranging from zero to 160. Three different colored bars are shown above each compound on the X axis. One bar represents intracellular fluid (ICF), a second bar represents interstitial fluid (IF, which is part of ECF) and the third bar represents plasma (ECF). Intracellular fluid contains high concentrations of K plus and HPO four two minus. It has lower concentrations of MG two plus and protein, and negligible amounts of the other compounds. Interstitial fluid contains high concentrations of NA plus and CL minus, along with a smaller amount of HCO 3 minus, and negligible amounts of the other compounds. Plasma contains large concentrations of NA plus and CL minus, with smaller concentrations of HCO 3 minus and protein, and negligible amounts of the other compounds. Figure 4. The Concentrations of Different Elements in Key Bodily Fluids. The graph shows the composition of the ICF, IF, and plasma. The compositions of plasma and IF are similar to one another but are quite different from the composition of the ICF.[/caption]
[caption id="" align="aligncenter" width="120"]QR Codes representing a URL Watch this video to learn more about body fluids, fluid compartments, and electrolytes.[/caption]

Most body fluids are neutral in charge. Thus, cations, or positively charged ions, and anions, or negatively charged ions, are balanced in fluids. As seen in the previous graph, sodium (Na+) ions and chloride (Cl-) ions are concentrated in the ECF of the body, whereas potassium (K+) ions are concentrated inside cells. Although sodium and potassium can “leak” through “pores” into and out of cells, respectively, the high levels of potassium and low levels of sodium in the ICF are maintained by sodium-potassium pumps in the cell membranes. These pumps use the energy supplied by ATP to pump sodium out of the cell and potassium into the cell (Figure 5).

[caption id="" align="aligncenter" width="520"]This diagram shows a sodium potassium pump embedded in the cell membrane. In the first step, the pump is opened to the cytosol and closed to the extracellular fluid. First, three sodium ions move into the pump from the cytosol. An ATP molecule binds to the cytosol side of the pump, causing the pump to change shape and open to the extracellular fluid. The pump is now closed to the cytosol. The sodium ions are then released into the extracellular fluid, after which two potassium ions enter the pump. Also at this point, the used ADP detaches from the cytosol side of the pump, leaving a single phosphate attached. The pump then changes shape again so that it closes to the extracellular fluid and again opens to the cytosol. This releases the two potassium ions into the cytosol. The single phosphate also detaches from the pump at this point so that the cycle can start anew. Two bars along the right hand side of the figure indicate that sodium normally diffuses into the cell down its concentration gradient while potassium usually diffuses out of the cell down its concentration gradient. Therefore, the sodium potassium pump is working against these natural concentration gradients. Figure 5. The Sodium-Potassium Pump. The sodium-potassium pump is powered by ATP to transfer sodium out of the cytoplasm and into the ECF. The pump also transfers potassium out of the ECF and into the cytoplasm. (credit: modification of work by Mariana Ruiz Villarreal)[/caption]

Fluid Movement between Compartments

Hydrostatic pressure, the force exerted by a fluid against a wall, causes movement of fluid between compartments. The hydrostatic pressure of blood is the pressure exerted by blood against the walls of the blood vessels by the pumping action of the heart. In capillaries, hydrostatic pressure (also known as capillary blood pressure) is higher than the opposing “colloid osmotic pressure” in blood—a “constant” pressure primarily produced by circulating albumin—at the arteriolar end of the capillary (Figure 6). This pressure forces plasma and nutrients out of the capillaries and into surrounding tissues. Fluid and the cellular wastes in the tissues enter the capillaries at the venule end, where the hydrostatic pressure is less than the osmotic pressure in the vessel. Filtration pressure squeezes fluid from the plasma in the blood to the IF surrounding the tissue cells. The surplus fluid in the interstitial space that is not returned directly back to the capillaries is drained from tissues by the lymphatic system, and then re-enters the vascular system at the subclavian veins.
[caption id="" align="aligncenter" width="480"]Alt text to come. Figure 6. Capillary Exchange. Net filtration occurs near the arterial end of the capillary since capillary hydrostatic pressure (CHP) is greater than blood colloidal osmotic pressure (BCOP). There is no net movement of fluid near the midpoint of the capillary since CHP = BCOP. Net reabsorption occurs near the venous end of the capillary since BCOP is greater than CHP.[/caption]
[caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this video to see an explanation of the dynamics of fluid in the body’s compartments.[/caption]

Hydrostatic pressure is especially important in governing the movement of water in the nephrons of the kidneys to ensure proper filtering of the blood to form urine. As hydrostatic pressure in the kidneys increases, the amount of water leaving the capillaries also increases, and more urine filtrate is formed. If hydrostatic pressure in the kidneys drops too low, as can happen in dehydration, the functions of the kidneys will be impaired, and less nitrogenous wastes will be removed from the bloodstream. Extreme dehydration can result in kidney failure.

Fluid also moves between compartments along an osmotic gradient. Recall that an osmotic gradient is produced by the difference in concentration of all solutes on either side of a semi-permeable membrane. The magnitude of the osmotic gradient is proportional to the difference in the concentration of solutes on one side of the cell membrane to that on the other side. Water will move by osmosis from the side where its concentration is high (and the concentration of solute is low) to the side of the membrane where its concentration is low (and the concentration of solute is high). In the body, water moves by osmosis from plasma to the IF (and the reverse) and from the IF to the ICF (and the reverse). In the body, water moves constantly into and out of fluid compartments as conditions change in different parts of the body.

For example, if you are sweating, you will lose water through your skin. Sweating depletes your tissues of water and increases the solute concentration in those tissues. As this happens, water diffuses from your blood into sweat glands and surrounding skin tissues that have become dehydrated because of the osmotic gradient. Additionally, as water leaves the blood, it is replaced by the water in other tissues throughout your body that are not dehydrated. If this continues, dehydration spreads throughout the body. When a dehydrated person drinks water and rehydrates, the water is redistributed by the same gradient, but in the opposite direction, replenishing water in all of the tissues.

Solute Movement between Compartments

The movement of some solutes between compartments is active, which consumes energy and is an active transport process, whereas the movement of other solutes is passive, which does not require energy. Active transport allows cells to move a specific substance against its concentration gradient through a membrane protein, requiring energy in the form of ATP. For example, the sodium-potassium pump employs active transport to pump sodium out of cells and potassium into cells, with both substances moving against their concentration gradients.

Passive transport of a molecule or ion depends on its ability to pass through the membrane, as well as the existence of a concentration gradient that allows the molecules to diffuse from an area of higher concentration to an area of lower concentration. Some molecules, like gases, lipids, and water itself (which also utilizes water channels in the membrane called aquaporins), slip fairly easily through the cell membrane; others, including polar molecules like glucose, amino acids, and ions do not. Some of these molecules enter and leave cells using facilitated transport, whereby the molecules move down a concentration gradient through specific protein channels in the membrane. This process does not require energy. For example, glucose is transferred into cells by glucose transporters that use facilitated transport (Figure 7).

[caption id="" align="aligncenter" width="480"]This diagram shows a carrier protein embedded in the plasma membrane between the cytoplasm and the extracellular fluid. There are several glucose molecules in the extracellular fluid. In the first step, the carrier protein is open to the extracellular fluid and closed to the cytosol. One of the glucose molecules travels from the extracellular fluid into the carrier protein. The protein then changes shape, closing at both ends. This pushes the glucose down into the carrier protein, closer to the cytosol end. The protein then opens on the cytosol side and closes on the extracellular fluid side, allowing the glucose to enter the cytosol. Figure 7. Facilitated Diffusion. Glucose molecules use facilitated diffusion to move down a concentration gradient through the carrier protein channels in the membrane. (credit: modification of work by Mariana Ruiz Villarreal)[/caption]
]]> 2748 2738 1 0 26.2 Water Balance https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2752 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2752

Learning Objectives

By the end of this section, you will be able to:
  • Explain how water levels in the body influence the thirst cycle
  • Identify the main route by which water leaves the body
  • Describe the role of ADH and its effect on body water levels
  • Define dehydration and identify common causes of dehydration

On a typical day, the average adult will take in about 2500 mL (almost 3 quarts) of aqueous fluids. Although most of the intake comes through the digestive tract, about 230 mL (8 ounces) per day is generated metabolically, in the last steps of aerobic respiration. Additionally, each day about the same volume (2500 mL) of water leaves the body by different routes; most of this lost water is removed as urine. The kidneys also can adjust blood volume though mechanisms that draw water out of the filtrate and urine. The kidneys can regulate water levels in the body; they conserve water if you are dehydrated, and they can make urine more dilute to expel excess water if necessary. Water is lost through the skin through evaporation from the skin surface without overt sweating and from air expelled from the lungs. This type of water loss is called insensible water loss because a person is usually unaware of it.

Regulation of Water Intake

Osmolality is the ratio of solutes in a solution to a volume of solvent in a solution. Plasma osmolality is thus the ratio of solutes to water in blood plasma. A person’s plasma osmolality value reflects his or her state of hydration. A healthy body maintains plasma osmolality within a narrow range, by employing several mechanisms that regulate both water intake and output.

Drinking water is considered voluntary. So how is water intake regulated by the body? Consider someone who is experiencing dehydration, a net loss of water that results in insufficient water in blood and other tissues. The water that leaves the body, as exhaled air, sweat, or urine, is ultimately extracted from blood plasma. As the blood becomes more concentrated, the thirst response—a sequence of physiological processes—is triggered (Figure 1). Osmoreceptors are sensory receptors in the thirst center in the hypothalamus that monitor the concentration of solutes (osmolality) of the blood. If blood osmolality increases above its ideal value, the hypothalamus transmits signals that result in a conscious awareness of thirst. The person should (and normally does) respond by drinking water. The hypothalamus of a dehydrated person also releases antidiuretic hormone (ADH) through the posterior pituitary gland. ADH signals the kidneys to recover water from urine, effectively diluting the blood plasma. To conserve water, the hypothalamus of a dehydrated person also sends signals via the sympathetic nervous system to the salivary glands in the mouth. The signals result in a decrease in watery, serous output (and an increase in stickier, thicker mucus output). These changes in secretions result in a “dry mouth” and the sensation of thirst.

[caption id="" align="aligncenter" width="380"]This figure is a top-to bottom flowchart describing the thirst response. The topmost box of the chart states that there is insufficient water in the body, which has two effects. The left branch of the chart leads to decreased blood volume, which leads to decreased blood pressure. This triggers an increase in angiotensin two. Angiotensin two stimulates the thirst center in the hypothalamus. On the right branch, insufficient water in the body leads to increased blood osmolality, which causes dry mouth. Increased blood osmolality and dry mouth is sensed by osmoreceptors in the hypothalamus. This stimulates the thirst center in the hypothalamus to increase thirst, giving a person the urge to drink. Drinking decreases blood osmolality back to homeostatic levels. Figure 1. A Flowchart Showing the Thirst Response. The thirst response begins when osmoreceptors detect a decrease in water levels in the blood.[/caption]

Decreased blood volume resulting from water loss has two additional effects. First, baroreceptors, blood-pressure receptors in the arch of the aorta and the carotid arteries in the neck, detect a decrease in blood pressure that results from decreased blood volume. The heart is ultimately signaled to increase its rate and/or strength of contractions to compensate for the lowered blood pressure.

Second, the kidneys have a renin-angiotensin hormonal system that increases the production of the active form of the hormone angiotensin II, which helps stimulate thirst, but also stimulates the release of the hormone aldosterone from the adrenal glands. Aldosterone increases the reabsorption of sodium in the distal tubules of the nephrons in the kidneys, and water follows this reabsorbed sodium back into the blood.

If adequate fluids are not consumed, dehydration results and a person’s body contains too little water to function correctly. A person who repeatedly vomits or who has diarrhea may become dehydrated, and infants, because their body mass is so low, can become dangerously dehydrated very quickly. Endurance athletes such as distance runners often become dehydrated during long races. Dehydration can be a medical emergency, and a dehydrated person may lose consciousness, become comatose, or die, if his or her body is not rehydrated quickly.

Regulation of Water Output

Water loss from the body occurs predominantly through the renal system. A person produces an average of 1.5 liters (1.6 quarts) of urine per day. Although the volume of urine varies in response to hydration levels, there is a minimum volume of urine production required for proper bodily functions. The kidney excretes 100 to 1200 milliosmoles of solutes per day to rid the body of a variety of excess salts and other water-soluble chemical wastes, most notably creatinine, urea, and uric acid. Failure to produce the minimum volume of urine means that metabolic wastes cannot be effectively removed from the body, a situation that can impair organ function. The minimum level of urine production necessary to maintain normal function is about 0.47 liters (0.5 quarts) per day.

The kidneys also must make adjustments in the event of ingestion of too much fluid. Diuresis, which is the production of urine in excess of normal levels, begins about 30 minutes after drinking a large quantity of fluid. Diuresis reaches a peak after about 1 hour, and normal urine production is reestablished after about 3 hours.

Role of ADH

Antidiuretic hormone (ADH), also known as vasopressin, controls the amount of water reabsorbed from the collecting ducts and tubules in the kidney. This hormone is produced in the hypothalamus and is delivered to the posterior pituitary for storage and release (Figure 2). When the osmoreceptors in the hypothalamus detect an increase in the concentration of blood plasma, the hypothalamus signals the release of ADH from the posterior pituitary into the blood.

[caption id="" align="aligncenter" width="450"]This set of diagrams shows the effects of ADH on various structures within the body. In the brain, ADH affects the cerebrum by influencing social behavior in some mammals. ADH is also produced in the brain by the hypothalamus and released in the posterior pituitary. ADH also constricts arterioles in the body, which are the small arteries that enter into capillary beds. Finally, a kidney is shown because ADH increases the reabsorption of water in the kidneys. Figure 2. Antidiuretic Hormone (ADH). ADH is produced in the hypothalamus and released by the posterior pituitary gland. It causes the kidneys to retain water, constricts arterioles in the peripheral circulation, and affects some social behaviors in mammals.[/caption]
ADH has two major effects. It constricts the arterioles in the peripheral circulation, which reduces the flow of blood to the extremities and thereby increases the blood supply to the core of the body. ADH also causes the epithelial cells that line the renal collecting tubules to move water channel proteins, called aquaporins, from the interior of the cells to the apical surface, where these proteins are inserted into the cell membrane (Figure 3). The result is an increase in the water permeability of these cells and, thus, a large increase in water passage from the urine through the walls of the collecting tubules, leading to more reabsorption of water into the bloodstream. When the blood plasma becomes less concentrated and the level of ADH decreases, aquaporins are removed from collecting tubule cell membranes, and the passage of water out of urine and into the blood decreases.
[caption id="" align="aligncenter" width="480"]This diagram depicts a cross section of the right wall of a kidney collecting tubule. The wall is composed of three block-shaped cells arranged vertically one on top of each other. The lumen of the collecting tubule is to the left of the three cells. Yellow-colored urine is flowing through the lumen. There is a small strip of blue interstitial fluid to the right of the three cells. To the right of the interstitial fluid is a cross section of a blood vessel. Arrows show that water in the urine is entering the left side of the wall cells through aquaporins. The water travels through the cells and then leaves the kidney tubule through additional aquaporins in the right side of the wall cells. The water travels through the interstitial space and enters into the blood in the blood vessel. The aquaporins in the wall cells are being released from aquaporin storage vesicles within their cytoplasm. Figure 3. Aquaporins. The binding of ADH to receptors on the cells of the collecting tubule results in aquaporins being inserted into the plasma membrane, shown in the lower cell. This dramatically increases the flow of water out of the tubule and into the bloodstream.[/caption]
A diuretic is a compound that increases urine output and therefore decreases water conservation by the body. Diuretics are used to treat hypertension, congestive heart failure, and fluid retention associated with menstruation. Alcohol acts as a diuretic by inhibiting the release of ADH. Additionally, caffeine, when consumed in high concentrations, acts as a diuretic.
]]> 2752 2738 2 0 26.3 Electrolyte Balance https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2756 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2756

Learning Objectives

By the end of this section, you will be able to:
  • List the role of the six most important electrolytes in the body
  • Name the disorders associated with abnormally high and low levels of the six electrolytes
  • Identify the predominant extracellular anion
  • Describe the role of aldosterone on the level of water in the body

The body contains a large variety of ions, or electrolytes, which perform a variety of functions. Some ions assist in the transmission of electrical impulses along cell membranes in neurons and muscles. Other ions help to stabilize protein structures in enzymes. Still others aid in releasing hormones from endocrine glands. All of the ions in plasma contribute to the osmotic balance that controls the movement of water between cells and their environment.

Electrolytes in living systems include sodium, potassium, chloride, bicarbonate, calcium, phosphate, magnesium, copper, zinc, iron, manganese, molybdenum, copper, and chromium. In terms of body functioning, six electrolytes are most important: sodium, potassium, chloride, bicarbonate, calcium, and phosphate.

Roles of Electrolytes

These six ions aid in nerve excitability, endocrine secretion, membrane permeability, buffering body fluids, and controlling the movement of fluids between compartments. These ions enter the body through the digestive tract. More than 90 percent of the calcium and phosphate that enters the body is incorporated into bones and teeth, with bone serving as a mineral reserve for these ions. In the event that calcium and phosphate are needed for other functions, bone tissue can be broken down to supply the blood and other tissues with these minerals. Phosphate is a normal constituent of nucleic acids; hence, blood levels of phosphate will increase whenever nucleic acids are broken down.

Excretion of ions occurs mainly through the kidneys, with lesser amounts lost in sweat and in feces. Excessive sweating may cause a significant loss, especially of sodium and chloride. Severe vomiting or diarrhea will cause a loss of chloride and bicarbonate ions. Adjustments in respiratory and renal functions allow the body to regulate the levels of these ions in the ECF.

Table 1 lists the reference values for blood plasma, cerebrospinal fluid (CSF), and urine for the six ions addressed in this section. In a clinical setting, sodium, potassium, and chloride are typically analyzed in a routine urine sample. In contrast, calcium and phosphate analysis requires a collection of urine across a 24-hour period, because the output of these ions can vary considerably over the course of a day. Urine values reflect the rates of excretion of these ions. Bicarbonate is the one ion that is not normally excreted in urine; instead, it is conserved by the kidneys for use in the body’s buffering systems.

Electrolyte and Ion Reference Values (Table 1)
Name Chemical symbol Plasma CSF Urine
Sodium Na+ 136.00–146.00 (mM) 138.00–150.00 (mM) 40.00–220.00 (mM)
Potassium K+ 3.50–5.00 (mM) 0.35–3.5 (mM) 25.00–125.00 (mM)
Chloride Cl- 98.00–107.00 (mM) 118.00–132.00 (mM) 110.00–250.00 (mM)
Bicarbonate HCO3- 22.00–29.00 (mM) ------ ------
Calcium Ca++ 2.15–2.55 (mmol/day) ------ Up to 7.49 (mmol/day)
Phosphate HPO42−HPO42− 0.81–1.45 (mmol/day) ------ 12.90–42.00 (mmol/day)

Sodium

Sodium is the major cation of the extracellular fluid. It is responsible for one-half of the osmotic pressure gradient that exists between the interior of cells and their surrounding environment. People eating a typical Western diet, which is very high in NaCl, routinely take in 130 to 160 mmol/day of sodium, but humans require only 1 to 2 mmol/day. This excess sodium appears to be a major factor in hypertension (high blood pressure) in some people. Excretion of sodium is accomplished primarily by the kidneys. Sodium is freely filtered through the glomerular capillaries of the kidneys, and although much of the filtered sodium is reabsorbed in the proximal convoluted tubule, some remains in the filtrate and urine, and is normally excreted.

Hyponatremia is a lower-than-normal concentration of sodium, usually associated with excess water accumulation in the body, which dilutes the sodium. An absolute loss of sodium may be due to a decreased intake of the ion coupled with its continual excretion in the urine. An abnormal loss of sodium from the body can result from several conditions, including excessive sweating, vomiting, or diarrhea; the use of diuretics; excessive production of urine, which can occur in diabetes; and acidosis, either metabolic acidosis or diabetic ketoacidosis.

A relative decrease in blood sodium can occur because of an imbalance of sodium in one of the body’s other fluid compartments, like IF, or from a dilution of sodium due to water retention related to edema or congestive heart failure. At the cellular level, hyponatremia results in increased entry of water into cells by osmosis, because the concentration of solutes within the cell exceeds the concentration of solutes in the now-diluted ECF. The excess water causes swelling of the cells; the swelling of red blood cells—decreasing their oxygen-carrying efficiency and making them potentially too large to fit through capillaries—along with the swelling of neurons in the brain can result in brain damage or even death.

Hypernatremia is an abnormal increase of blood sodium. It can result from water loss from the blood, resulting in the hemoconcentration of all blood constituents. Hormonal imbalances involving ADH and aldosterone may also result in higher-than-normal sodium values.

Potassium

Potassium is the major intracellular cation. It helps establish the resting membrane potential in neurons and muscle fibers after membrane depolarization and action potentials. In contrast to sodium, potassium has very little effect on osmotic pressure. The low levels of potassium in blood and CSF are due to the sodium-potassium pumps in cell membranes, which maintain the normal potassium concentration gradients between the ICF and ECF. The recommendation for daily intake/consumption of potassium is 4700 mg. Potassium is excreted, both actively and passively, through the renal tubules, especially the distal convoluted tubule and collecting ducts. Potassium participates in the exchange with sodium in the renal tubules under the influence of aldosterone, which also relies on basolateral sodium-potassium pumps.

Hypokalemia is an abnormally low potassium blood level. Similar to the situation with hyponatremia, hypokalemia can occur because of either an absolute reduction of potassium in the body or a relative reduction of potassium in the blood due to the redistribution of potassium. An absolute loss of potassium can arise from decreased intake, frequently related to starvation. It can also come about from vomiting, diarrhea, or alkalosis.

Some insulin-dependent diabetic patients experience a relative reduction of potassium in the blood from the redistribution of potassium. When insulin is administered and glucose is taken up by cells, potassium passes through the cell membrane along with glucose, decreasing the amount of potassium in the blood and IF, which can cause hyperpolarization of the cell membranes of neurons, reducing their responses to stimuli. Hyperkalemia, an elevated potassium blood level, also can impair the function of skeletal muscles, the nervous system, and the heart. Hyperkalemia can result from increased dietary intake of potassium. In such a situation, potassium from the blood ends up in the ECF in abnormally high concentrations. This can result in a partial depolarization (excitation) of the plasma membrane of skeletal muscle fibers, neurons, and cardiac cells of the heart, and can also lead to an inability of cells to repolarize. For the heart, this means that it won’t relax after a contraction, and will effectively “seize” and stop pumping blood, which is fatal within minutes. Because of such effects on the nervous system, a person with hyperkalemia may also exhibit mental confusion, numbness, and weakened respiratory muscles.

Chloride

Chloride is the predominant extracellular anion. Chloride is a major contributor to the osmotic pressure gradient between the ICF and ECF, and plays an important role in maintaining proper hydration. Chloride functions to balance cations in the ECF, maintaining the electrical neutrality of this fluid. The paths of secretion and reabsorption of chloride ions in the renal system follow the paths of sodium ions.

Hypochloremia, or lower-than-normal blood chloride levels, can occur because of defective renal tubular absorption. Vomiting, diarrhea, and metabolic acidosis can also lead to hypochloremia. Hyperchloremia, or higher-than-normal blood chloride levels, can occur due to dehydration, excessive intake of dietary salt (NaCl) or swallowing of sea water, aspirin intoxication, congestive heart failure, and the hereditary, chronic lung disease, cystic fibrosis. In people who have cystic fibrosis, chloride levels in sweat are two to five times those of normal levels, and analysis of sweat is often used in the diagnosis of the disease.
[caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this video to see an explanation of the effect of seawater on humans. What effect does drinking seawater have on the body?[/caption]

Bicarbonate

Bicarbonate is the second most abundant anion in the blood. Its principal function is to maintain your body’s acid-base balance by being part of buffer systems. This role will be discussed in a different section.

Bicarbonate ions result from a chemical reaction that starts with carbon dioxide (CO2) and water, two molecules that are produced at the end of aerobic metabolism. Only a small amount of CO2 can be dissolved in body fluids. Thus, over 90 percent of the CO2 is converted into bicarbonate ions, HCO3–, through the following reactions:

CO2 + H2O ↔ H2CO3 ↔ H2CO3- + H+

The bidirectional arrows indicate that the reactions can go in either direction, depending on the concentrations of the reactants and products. Carbon dioxide is produced in large amounts in tissues that have a high metabolic rate. Carbon dioxide is converted into bicarbonate in the cytoplasm of red blood cells through the action of an enzyme called carbonic anhydrase. Bicarbonate is transported in the blood. Once in the lungs, the reactions reverse direction, and CO2 is regenerated from bicarbonate to be exhaled as metabolic waste.

Calcium

About two pounds of calcium in your body are bound up in bone, which provides hardness to the bone and serves as a mineral reserve for calcium and its salts for the rest of the tissues. Teeth also have a high concentration of calcium within them. A little more than one-half of blood calcium is bound to proteins, leaving the rest in its ionized form. Calcium ions, Ca2+, are necessary for muscle contraction, enzyme activity, and blood coagulation. In addition, calcium helps to stabilize cell membranes and is essential for the release of neurotransmitters from neurons and of hormones from endocrine glands.

Calcium is absorbed through the intestines under the influence of activated vitamin D. A deficiency of vitamin D leads to a decrease in absorbed calcium and, eventually, a depletion of calcium stores from the skeletal system, potentially leading to rickets in children and osteomalacia in adults, contributing to osteoporosis.

Hypocalcemia, or abnormally low calcium blood levels, is seen in hypoparathyroidism, which may follow the removal of the thyroid gland, because the four nodules of the parathyroid gland are embedded in it. Hypercalcemia, or abnormally high calcium blood levels, is seen in primary hyperparathyroidism. Some malignancies may also result in hypercalcemia.

Phosphate

Phosphate is present in the body in three ionic forms: H2PO4−, HPO42, and PO43−. The most common form is HPO42−HPO42−. Bone and teeth bind up 85 percent of the body’s phosphate as part of calcium-phosphate salts. Phosphate is found in phospholipids, such as those that make up the cell membrane, and in ATP, nucleotides, and buffers.

Hypophosphatemia, or abnormally low phosphate blood levels, occurs with heavy use of antacids, during alcohol withdrawal, and during malnourishment. In the face of phosphate depletion, the kidneys usually conserve phosphate, but during starvation, this conservation is impaired greatly. Hyperphosphatemia, or abnormally increased levels of phosphates in the blood, occurs if there is decreased renal function or in cases of acute lymphocytic leukemia. Additionally, because phosphate is a major constituent of the ICF, any significant destruction of cells can result in dumping of phosphate into the ECF.

Regulation of Sodium and Potassium

Sodium is reabsorbed from the renal filtrate, and potassium is excreted into the filtrate in the renal collecting tubule. The control of this exchange is governed principally by two hormones—aldosterone and angiotensin II.

Aldosterone

Recall that aldosterone increases the excretion of potassium and the reabsorption of sodium in the distal tubule. Aldosterone is released if blood levels of potassium increase, if blood levels of sodium severely decrease, or if blood pressure decreases. Its net effect is to conserve and increase water levels in the plasma by reducing the excretion of sodium, and thus water, from the kidneys. In a negative feedback loop, increased osmolality of the ECF (which follows aldosterone-stimulated sodium absorption) inhibits the release of the hormone (Figure 1).
[caption id="" align="aligncenter" width="250"]This flow chart shows how potassium and sodium ion concentrations in the blood are regulated by aldosterone. Rising K plus and falling NA plus levels in the blood trigger aldosterone release from the adrenal cortex. Aldosterone targets the kidneys, causing a decrease in K plus release from the kidneys, which reduces the amount of K plus in the blood back to homeostatic levels. Aldosterone also increases sodium reabsorption by the kidneys, which increases the amount of NA plus in the blood back to homeostatic levels. Figure 1. The Aldosterone Feedback Loop. Aldosterone, which is released by the adrenal gland, facilitates reabsorption of Na+ and thus the reabsorption of water.[/caption]

Angiotensin II

Angiotensin II causes vasoconstriction and an increase in systemic blood pressure. This action increases the glomerular filtration rate, resulting in more material filtered out of the glomerular capillaries and into Bowman’s capsule. Angiotensin II also signals an increase in the release of aldosterone from the adrenal cortex.

In the distal convoluted tubules and collecting ducts of the kidneys, aldosterone stimulates the synthesis and activation of the sodium-potassium pump (Figure 2). Sodium passes from the filtrate, into and through the cells of the tubules and ducts, into the ECF and then into capillaries. Water follows the sodium due to osmosis. Thus, aldosterone causes an increase in blood sodium levels and blood volume. Aldosterone’s effect on potassium is the reverse of that of sodium; under its influence, excess potassium is pumped into the renal filtrate for excretion from the body.

[caption id="" align="aligncenter" width="550"]This figure shows the hormone cascade that that increases kidney reabsorption of NA plus and water. In the first step, the kidneys release renin into the blood stream. The blood stream is depicted with a red arrow pointing from left to right. At the same time, the liver releases angiotensinogen into the blood, which combines with the renin, yielding angiotensin one. The blood flow then leads to the lungs. Within the pulmonary blood, angiotensin-converting enzyme (ACE) converts angiotensin one to angiotensin two. The blood then flows to the adrenal cortex, where angiotensin two stimulates the adrenal cortex to secrete aldosterone. Aldosterone causes the kidney tubules to increase reabsorption of NA plus and water into the blood. Figure 2. The Renin-Angiotensin System. Angiotensin II stimulates the release of aldosterone from the adrenal cortex.[/caption]

Regulation of Calcium and Phosphate

Calcium and phosphate are both regulated through the actions of three hormones: parathyroid hormone (PTH), dihydroxyvitamin D (calcitriol), and calcitonin. All three are released or synthesized in response to the blood levels of calcium.

PTH is released from the parathyroid gland in response to a decrease in the concentration of blood calcium. The hormone activates osteoclasts to break down bone matrix and release inorganic calcium-phosphate salts. PTH also increases the gastrointestinal absorption of dietary calcium by converting vitamin D into dihydroxyvitamin D (calcitriol), an active form of vitamin D that intestinal epithelial cells require to absorb calcium.

PTH raises blood calcium levels by inhibiting the loss of calcium through the kidneys. PTH also increases the loss of phosphate through the kidneys.

Calcitonin is released from the thyroid gland in response to elevated blood levels of calcium. The hormone increases the activity of osteoblasts, which remove calcium from the blood and incorporate calcium into the bony matrix.

]]> 2756 2738 3 0 26.4 Acid-Base Balance https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2760 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2760

Learning Objectives

By the end of this section, you will be able to:
  • Identify the most powerful buffer system in the body
  • Explain the way in which the respiratory system affects blood pH

Proper physiological functioning depends on a very tight balance between the concentrations of acids and bases in the blood. Acid-balance balance is measured using the pH scale, as shown in Figure 1. A variety of buffering systems permits blood and other bodily fluids to maintain a narrow pH range, even in the face of perturbations. A buffer is a chemical system that prevents a radical change in fluid pH by dampening the change in hydrogen ion concentrations in the case of excess acid or base. Most commonly, the substance that absorbs the ions is either a weak acid, which takes up hydroxyl ions, or a weak base, which takes up hydrogen ions.

[caption id="" align="aligncenter" width="320"]This table gives examples of solutions from PH of zero to 14. Examples of solutions with a PH of zero include battery acid and strong hydrofluoric acid. An example of a solution with a pH of one is the hydrochloric acid secreted by the stomach lining. Examples of solutions with a PH of two include lemon juice and vinegar. Examples of solutions with a PH of three include grapefruit juice, orange juice and soda. Examples of solutions with a PH of four include tomato juice and acid rain. Examples of solutions with a PH of five include soft drinking water and black coffee. Examples of solutions with a PH of six include urine and saliva. An example of a solution with a PH of seven is pure water. An example of a solution with a PH of eight is sea water. An example of a solution with a PH of nine is baking soda. Examples of solutions with a PH of ten include saline lake water and milk of magnesia. An example of a solution with a PH of eleven is an ammonia solution. An example of a solution with a PH of twelve is soapy water. Examples of solutions with a PH of thirteen include bleach and oven cleaner. An example of a solution with a PH of fourteen is liquid drain cleaner. Figure 1. The pH Scale. This chart shows where many common substances fall on the pH scale.[/caption]

Buffer Systems in the Body

The buffer systems in the human body are extremely efficient, and different systems work at different rates. It takes only seconds for the chemical buffers in the blood to make adjustments to pH. The respiratory tract can adjust the blood pH upward in minutes by exhaling CO2 from the body. The renal system can also adjust blood pH through the excretion of hydrogen ions (H+) and the conservation of bicarbonate, but this process takes hours to days to have an effect.

The buffer systems functioning in blood plasma include plasma proteins, phosphate, and bicarbonate and carbonic acid buffers. The kidneys help control acid-base balance by excreting hydrogen ions and generating bicarbonate that helps maintain blood plasma pH within a normal range. Protein buffer systems work predominantly inside cells.

Protein Buffers in Blood Plasma and Cells

Nearly all proteins can function as buffers. Proteins are made up of amino acids, which contain positively charged amino groups and negatively charged carboxyl groups. The charged regions of these molecules can bind hydrogen and hydroxyl ions, and thus function as buffers. Buffering by proteins accounts for two-thirds of the buffering power of the blood and most of the buffering within cells.

Hemoglobin as a Buffer

Hemoglobin is the principal protein inside of red blood cells and accounts for one-third of the mass of the cell. During the conversion of CO2 into bicarbonate, hydrogen ions liberated in the reaction are buffered by hemoglobin, which is reduced by the dissociation of oxygen. This buffering helps maintain normal pH. The process is reversed in the pulmonary capillaries to re-form CO2, which then can diffuse into the air sacs to be exhaled into the atmosphere. This process is discussed in detail in the chapter on the respiratory system.

Phosphate Buffer

Phosphates are found in the blood in two forms: sodium dihydrogen phosphate (Na2H2PO4−), which is a weak acid, and sodium monohydrogen phosphate (Na2HPO42-), which is a weak base. When Na2HPO42- comes into contact with a strong acid, such as HCl, the base picks up a second hydrogen ion to form the weak acid Na2H2PO4− and sodium chloride, NaCl. When Na2HPO42− (the weak acid) comes into contact with a strong base, such as sodium hydroxide (NaOH), the weak acid reverts back to the weak base and produces water. Acids and bases are still present, but they hold onto the ions.

HCl + Na2HPO4→NaH2PO4 + NaCl
(strong acid) + (weak base) → (weak acid) + (salt)
NaOH + NaH2PO4→Na2HPO4 + H2O
(strong base) + (weak acid) → (weak base) + (water)

Bicarbonate-Carbonic Acid Buffer

The bicarbonate-carbonic acid buffer works in a fashion similar to phosphate buffers. The bicarbonate is regulated in the blood by sodium, as are the phosphate ions. When sodium bicarbonate (NaHCO3), comes into contact with a strong acid, such as HCl, carbonic acid (H2CO3), which is a weak acid, and NaCl are formed. When carbonic acid comes into contact with a strong base, such as NaOH, bicarbonate and water are formed.

NaHCO3 + HCl →  H2CO3+NaCl
(sodium bicarbonate) + (strong acid) → (weak acid) + (salt)
H2CO3 + NaOH→HCO3- + H2O
(weak acid) + (strong base)→(bicarbonate) + (water)

As with the phosphate buffer, a weak acid or weak base captures the free ions, and a significant change in pH is prevented. Bicarbonate ions and carbonic acid are present in the blood in a 20:1 ratio if the blood pH is within the normal range. With 20 times more bicarbonate than carbonic acid, this capture system is most efficient at buffering changes that would make the blood more acidic. This is useful because most of the body’s metabolic wastes, such as lactic acid and ketones, are acids. Carbonic acid levels in the blood are controlled by the expiration of CO2 through the lungs. In red blood cells, carbonic anhydrase forces the dissociation of the acid, rendering the blood less acidic. Because of this acid dissociation, CO2 is exhaled (see equations above). The level of bicarbonate in the blood is controlled through the renal system, where bicarbonate ions in the renal filtrate are conserved and passed back into the blood. However, the bicarbonate buffer is the primary buffering system of the IF surrounding the cells in tissues throughout the body.

Respiratory Regulation of Acid-Base Balance

The respiratory system contributes to the balance of acids and bases in the body by regulating the blood levels of carbonic acid (Figure 2). CO2 in the blood readily reacts with water to form carbonic acid, and the levels of CO2 and carbonic acid in the blood are in equilibrium. When the CO2 level in the blood rises (as it does when you hold your breath), the excess CO2 reacts with water to form additional carbonic acid, lowering blood pH. Increasing the rate and/or depth of respiration (which you might feel the “urge” to do after holding your breath) allows you to exhale more CO2. The loss of CO2 from the body reduces blood levels of carbonic acid and thereby adjusts the pH upward, toward normal levels. As you might have surmised, this process also works in the opposite direction. Excessive deep and rapid breathing (as in hyperventilation) rids the blood of CO2 and reduces the level of carbonic acid, making the blood too alkaline. This brief alkalosis can be remedied by rebreathing air that has been exhaled into a paper bag. Rebreathing exhaled air will rapidly bring blood pH down toward normal.

[caption id="" align="aligncenter" width="280"]This top to bottom flowchart describes the regulation of PH in the blood. The left branch shows acidosis, which is when the PH of the blood drops. Acidosis stimulates brain and arterial receptors, triggering an increase in respiratory rate. This causes a drop in blood CO two and H two CO three. A drop in these two acidic compounds causes the blood PH to rise back to homeostatic levels. The right branch shows alkalosis which is when the PH of the blood rises. Alkalosis also stimulates brain and arterial receptors, but these now trigger a decrease in respiratory rate. This causes an increase in blood CO two and H two CO three, which lowers the PH of the blood back to homeostatic levels. Figure 2. Respiratory Regulation of Blood pH. The respiratory system can reduce blood pH by removing CO2 from the blood.[/caption]

The chemical reactions that regulate the levels of CO2 and carbonic acid occur in the lungs when blood travels through the lung’s pulmonary capillaries. Minor adjustments in breathing are usually sufficient to adjust the pH of the blood by changing how much CO2 is exhaled. In fact, doubling the respiratory rate for less than 1 minute, removing “extra” CO2, would increase the blood pH by 0.2. This situation is common if you are exercising strenuously over a period of time. To keep up the necessary energy production, you would produce excess CO2 (and lactic acid if exercising beyond your aerobic threshold). In order to balance the increased acid production, the respiration rate goes up to remove the CO2. This helps to keep you from developing acidosis.

The body regulates the respiratory rate by the use of chemoreceptors, which primarily use CO2 as a signal. Peripheral blood sensors are found in the walls of the aorta and carotid arteries. These sensors signal the brain to provide immediate adjustments to the respiratory rate if CO2 levels rise or fall. Yet other sensors are found in the brain itself. Changes in the pH of CSF affect the respiratory center in the medulla oblongata, which can directly modulate breathing rate to bring the pH back into the normal range.

Hypercapnia, or abnormally elevated blood levels of CO2, occurs in any situation that impairs respiratory functions, including pneumonia and congestive heart failure. Reduced breathing (hypoventilation) due to drugs such as morphine, barbiturates, or ethanol (or even just holding one’s breath) can also result in hypercapnia. Hypocapnia, or abnormally low blood levels of CO2, occurs with any cause of hyperventilation that drives off the CO2, such as salicylate toxicity, elevated room temperatures, fever, or hysteria.

Renal Regulation of Acid-Base Balance

The renal regulation of the body’s acid-base balance addresses the metabolic component of the buffering system. Whereas the respiratory system (together with breathing centers in the brain) controls the blood levels of carbonic acid by controlling the exhalation of CO2, the renal system controls the blood levels of bicarbonate. A decrease of blood bicarbonate can result from the inhibition of carbonic anhydrase by certain diuretics or from excessive bicarbonate loss due to diarrhea. Blood bicarbonate levels are also typically lower in people who have Addison’s disease (chronic adrenal insufficiency), in which aldosterone levels are reduced, and in people who have renal damage, such as chronic nephritis. Finally, low bicarbonate blood levels can result from elevated levels of ketones (common in unmanaged diabetes mellitus), which bind bicarbonate in the filtrate and prevent its conservation.

Bicarbonate ions, HCO3-, found in the filtrate, are essential to the bicarbonate buffer system, yet the cells of the tubule are not permeable to bicarbonate ions. The steps involved in supplying bicarbonate ions to the system are seen in Figure 3 and are summarized below:

  • Step 1: Sodium ions are reabsorbed from the filtrate in exchange for H+ by an antiport mechanism in the apical membranes of cells lining the renal tubule.
  • Step 2: The cells produce bicarbonate ions that can be shunted to peritubular capillaries.
  • Step 3: When CO2 is available, the reaction is driven to the formation of carbonic acid, which dissociates to form a bicarbonate ion and a hydrogen ion.
  • Step 4: The bicarbonate ion passes into the peritubular capillaries and returns to the blood. The hydrogen ion is secreted into the filtrate, where it can become part of new water molecules and be reabsorbed as such, or removed in the urine.
[caption id="" align="aligncenter" width="550"]This diagram depicts a cross section of the left wall of a kidney proximal tubule. The wall is composed of two block-shaped cells arranged vertically one on top of each other. The lumen of the proximal tubule is to the left of the two cells. Yellow-colored urine is flowing through the lumen. There is a small strip of blue interstitial fluid to the right of the two cells. To the right of the interstitial fluid is a cross section of a blood vessel. A loop of chemical reactions is occurring in the diagram. Within the lumen of the proximal tubule, HCO three minus is combining with an H plus ion that enters the lumen from a proximal tubule cell. This reaction forms H two CO three. H two CO three then breaks into H two O and CO two, a reaction catalyzed by the enzyme carbonic anhydrase. The CO two then moves from the lumen of the proximal tubule into one of the proximal tubule cells. There, the reaction runs in reverse, with CO two combining with H two O to form H two CO three. The H two CO three then splits into H plus and HCO three minus. The H plus moves into the lumen, reinitiating the first step of the loop. The HCO three minus leaves the proximal tubule cell and enters the blood stream. Figure 3. Conservation of Bicarbonate in the Kidney. Tubular cells are not permeable to bicarbonate; thus, bicarbonate is conserved rather than reabsorbed. Steps 1 and 2 of bicarbonate conservation are indicated.[/caption]

It is also possible that salts in the filtrate, such as sulfates, phosphates, or ammonia, will capture hydrogen ions. If this occurs, the hydrogen ions will not be available to combine with bicarbonate ions and produce CO2. In such cases, bicarbonate ions are not conserved from the filtrate to the blood, which will also contribute to a pH imbalance and acidosis.

The hydrogen ions also compete with potassium to exchange with sodium in the renal tubules. If more potassium is present than normal, potassium, rather than the hydrogen ions, will be exchanged, and increased potassium enters the filtrate. When this occurs, fewer hydrogen ions in the filtrate participate in the conversion of bicarbonate into CO2 and less bicarbonate is conserved. If there is less potassium, more hydrogen ions enter the filtrate to be exchanged with sodium and more bicarbonate is conserved.

Chloride ions are important in neutralizing positive ion charges in the body. If chloride is lost, the body uses bicarbonate ions in place of the lost chloride ions. Thus, lost chloride results in an increased reabsorption of bicarbonate by the renal system.

]]> 2760 2738 4 0 26.5 Disorders of Acid-Base Balance https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2762 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2762

Learning Objectives

By the end of this section, you will be able to:
  • Identify the three blood variables considered when making a diagnosis of acidosis or alkalosis
  • Identify the source of compensation for blood pH problems of a respiratory origin
  • Identify the source of compensation for blood pH problems of a metabolic/renal origin
Normal arterial blood pH is restricted to a very narrow range of 7.35 to 7.45. A person who has a blood pH below 7.35 is considered to be in acidosis (actually, “physiological acidosis,” because blood is not truly acidic until its pH drops below 7), and a continuous blood pH below 7.0 can be fatal. Acidosis has several symptoms, including headache and confusion, and the individual can become lethargic and easily fatigued (Figure 1). A person who has a blood pH above 7.45 is considered to be in alkalosis, and a pH above 7.8 is fatal. Some symptoms of alkalosis include cognitive impairment (which can progress to unconsciousness), tingling or numbness in the extremities, muscle twitching and spasm, and nausea and vomiting. Both acidosis and alkalosis can be caused by either metabolic or respiratory disorders.

As discussed earlier in this chapter, the concentration of carbonic acid in the blood is dependent on the level of CO2 in the body and the amount of CO2 gas exhaled through the lungs. Thus, the respiratory contribution to acid-base balance is usually discussed in terms of CO2 (rather than of carbonic acid). Remember that a molecule of carbonic acid is lost for every molecule of CO2 exhaled, and a molecule of carbonic acid is formed for every molecule of CO2 retained.

[caption id="" align="aligncenter" width="480"]This figure points out the symptoms of acidosis and alkalosis on a silhouette of a human torso. The effects of acidosis on the central nervous system include headache, sleepiness, confusion, loss of consciousness and coma. The effects of acidosis are given on the left side of the diagram. The effects of acidosis on the respiratory system include shortness of breath and coughing. The effects of acidosis on the heart include arrhythmia and increased heart rate. The effects of acidosis on the muscular system include seizures and weakness. The effects of acidosis on the digestive system include nausea, vomiting and diarrhea. The right side of the diagram describes the symptoms of alkalosis. The effects of alkalosis on the central nervous system include confusion, light-headedness, stupor, and coma. The effects of alkalosis on the peripheral nervous system include hand tremor and numbness or tingling in the face, hands, and feet. The effects of alkalosis on the muscular system include twitching and prolonged spasms. The effects of alkalosis on the digestive system include nausea and vomiting. Figure 1. Symptoms of Acidosis and Alkalosis. Symptoms of acidosis affect several organ systems. Both acidosis and alkalosis can be diagnosed using a blood test.[/caption]

Metabolic Acidosis: Primary Bicarbonate Deficiency

Metabolic acidosis occurs when the blood is too acidic (pH below 7.35) due to too little bicarbonate, a condition called primary bicarbonate deficiency. At the normal pH of 7.40, the ratio of bicarbonate to carbonic acid buffer is 20:1. If a person’s blood pH drops below 7.35, then he or she is in metabolic acidosis. The most common cause of metabolic acidosis is the presence of organic acids or excessive ketones in the blood. Table 2 lists some other causes of metabolic acidosis.

*Acid metabolites from ingested chemical.
Common Causes of Metabolic Acidosis and Blood Metabolites (Table 2)
Cause Metabolite
Diarrhea Bicarbonate
Uremia Phosphoric, sulfuric, and lactic acids
Diabetic ketoacidosis Increased ketones
Strenuous exercise Lactic acid
Methanol Formic acid*
Paraldehyde β-Hydroxybutyric acid*
Isopropanol Propionic acid*
Ethylene glycol Glycolic acid, and some oxalic and formic acids*
Salicylate/aspirin Sulfasalicylic acid (SSA)*

The first three of the eight causes of metabolic acidosis listed are medical (or unusual physiological) conditions. Strenuous exercise can cause temporary metabolic acidosis due to the production of lactic acid. The last five causes result from the ingestion of specific substances. The active form of aspirin is its metabolite, sulfasalicylic acid. An overdose of aspirin causes acidosis due to the acidity of this metabolite. Metabolic acidosis can also result from uremia, which is the retention of urea and uric acid. Metabolic acidosis can also arise from diabetic ketoacidosis, wherein an excess of ketones is present in the blood. Other causes of metabolic acidosis are a decrease in the excretion of hydrogen ions, which inhibits the conservation of bicarbonate ions, and excessive loss of bicarbonate ions through the gastrointestinal tract due to diarrhea.

Metabolic Alkalosis: Primary Bicarbonate Excess

Metabolic alkalosis is the opposite of metabolic acidosis. It occurs when the blood is too alkaline (pH above 7.45) due to too much bicarbonate (called primary bicarbonate excess). A transient excess of bicarbonate in the blood can follow ingestion of excessive amounts of bicarbonate, citrate, or antacids for conditions such as stomach acid reflux—known as heartburn. Cushing’s disease, which is the chronic hypersecretion of adrenocorticotrophic hormone (ACTH) by the anterior pituitary gland, can cause chronic metabolic alkalosis. The oversecretion of ACTH results in elevated aldosterone levels and an increased loss of potassium by urinary excretion. Other causes of metabolic alkalosis include the loss of hydrochloric acid from the stomach through vomiting, potassium depletion due to the use of diuretics for hypertension, and the excessive use of laxatives.

Respiratory Acidosis: Primary Carbonic Acid/CO2 Excess

Respiratory acidosis occurs when the blood is overly acidic due to an excess of carbonic acid, resulting from too much CO2 in the blood. Respiratory acidosis can result from anything that interferes with respiration, such as pneumonia, emphysema, or congestive heart failure.

Respiratory Alkalosis: Primary Carbonic Acid/CO2 Deficiency

Respiratory alkalosis occurs when the blood is overly alkaline due to a deficiency in carbonic acid and CO2 levels in the blood. This condition usually occurs when too much CO2 is exhaled from the lungs, as occurs in hyperventilation, which is breathing that is deeper or more frequent than normal. An elevated respiratory rate leading to hyperventilation can be due to extreme emotional upset or fear, fever, infections, hypoxia, or abnormally high levels of catecholamines, such as epinephrine and norepinephrine. Surprisingly, aspirin overdose—salicylate toxicity—can result in respiratory alkalosis as the body tries to compensate for initial acidosis.

Compensation Mechanisms

Various compensatory mechanisms exist to maintain blood pH within a narrow range, including buffers, respiration, and renal mechanisms. Although compensatory mechanisms usually work very well, when one of these mechanisms is not working properly (like kidney failure or respiratory disease), they have their limits. If the pH and bicarbonate to carbonic acid ratio are changed too drastically, the body may not be able to compensate. Moreover, extreme changes in pH can denature proteins. Extensive damage to proteins in this way can result in disruption of normal metabolic processes, serious tissue damage, and ultimately death.

Respiratory Compensation

Respiratory compensation for metabolic acidosis increases the respiratory rate to drive off CO2 and readjust the bicarbonate to carbonic acid ratio to the 20:1 level. This adjustment can occur within minutes. Respiratory compensation for metabolic alkalosis is not as adept as its compensation for acidosis. The normal response of the respiratory system to elevated pH is to increase the amount of CO2 in the blood by decreasing the respiratory rate to conserve CO2. There is a limit to the decrease in respiration, however, that the body can tolerate. Hence, the respiratory route is less efficient at compensating for metabolic alkalosis than for acidosis.

Metabolic Compensation

Metabolic and renal compensation for respiratory diseases that can create acidosis revolves around the conservation of bicarbonate ions. In cases of respiratory acidosis, the kidney increases the conservation of bicarbonate and secretion of H+ through the exchange mechanism discussed earlier. These processes increase the concentration of bicarbonate in the blood, reestablishing the proper relative concentrations of bicarbonate and carbonic acid. In cases of respiratory alkalosis, the kidneys decrease the production of bicarbonate and reabsorb H+ from the tubular fluid. These processes can be limited by the exchange of potassium by the renal cells, which use a K+-H+ exchange mechanism (antiporter).

Diagnosing Acidosis and Alkalosis

Lab tests for pH, CO2 partial pressure (pCO2),and HCO3– can identify acidosis and alkalosis, indicating whether the imbalance is respiratory or metabolic, and the extent to which compensatory mechanisms are working. The blood pH value, as shown in Table 3, indicates whether the blood is in acidosis, the normal range, or alkalosis. The pCO2 and total HCO3– values aid in determining whether the condition is metabolic or respiratory, and whether the patient has been able to compensate for the problem. Table 3 lists the conditions and laboratory results that can be used to classify these conditions. Metabolic acid-base imbalances typically result from kidney disease, and the respiratory system usually responds to compensate.

Reference values (arterial): pH: 7.35–7.45; pCO2: male: 35–48 mm Hg, female: 32–45 mm Hg; total venous bicarbonate: 22–29 mM. N denotes normal; ↑ denotes a rising or increased value; and ↓ denotes a falling or decreased value.
Types of Acidosis and Alkalosis (Table 3)
pH pCO2 Total HCO3–
Metabolic acidosis ↓ N, then ↓ ↓
Respiratory acidosis ↓ ↑ N, then ↑
Metabolic alkalosis ↑ N, then↑ ↑
Respiratory alkalosis ↑ ↓ N, then ↓

Metabolic acidosis is problematic, as lower-than-normal amounts of bicarbonate are present in the blood. The pCO2 would be normal at first, but if compensation has occurred, it would decrease as the body reestablishes the proper ratio of bicarbonate and carbonic acid/CO2.

Respiratory acidosis is problematic, as excess CO2 is present in the blood. Bicarbonate levels would be normal at first, but if compensation has occurred, they would increase in an attempt to reestablish the proper ratio of bicarbonate and carbonic acid/CO2.

Alkalosis is characterized by a higher-than-normal pH. Metabolic alkalosis is problematic, as elevated pH and excess bicarbonate are present. The pCO2 would again be normal at first, but if compensation has occurred, it would increase as the body attempts to reestablish the proper ratios of bicarbonate and carbonic acid/CO2.

Respiratory alkalosis is problematic, as CO2 deficiency is present in the bloodstream. The bicarbonate concentration would be normal at first. When renal compensation occurs, however, the bicarbonate concentration in blood decreases as the kidneys attempt to reestablish the proper ratios of bicarbonate and carbonic acid/CO2 by eliminating more bicarbonate to bring the pH into the physiological range.

]]> 2762 2738 5 0 28.3 Fetal Development https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2807 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2807

Learning Objectives

By the end of this section, you will be able to:
  • Differentiate between the embryonic period and the fetal period
  • Briefly describe the process of sexual differentiation
  • Describe the fetal circulatory system and explain the role of the shunts
  • Trace the development of a fetus from the end of the embryonic period to birth

As you will recall, a developing human is called a fetus from the ninth week of gestation until birth. This 30-week period of development is marked by continued cell growth and differentiation, which fully develop the structures and functions of the immature organ systems formed during the embryonic period. The completion of fetal development results in a newborn who, although still immature in many ways, is capable of survival outside the womb.

Sexual Differentiation

Sexual differentiation does not begin until the fetal period, during weeks 9–12. Embryonic males and females, though genetically distinguishable, are morphologically identical (Figure 1). Bipotential gonads, or gonads that can develop into male or female sexual organs, are connected to a central cavity called the cloaca via Müllerian ducts and Wolffian ducts. (The cloaca is an extension of the primitive gut.) Several events lead to sexual differentiation during this period.

During male fetal development, the bipotential gonads become the testes and associated epididymis. The Müllerian ducts degenerate. The Wolffian ducts become the vas deferens, and the cloaca becomes the urethra and rectum.

During female fetal development, the bipotential gonads develop into ovaries. The Wolffian ducts degenerate. The Müllerian ducts become the uterine tubes and uterus, and the cloaca divides and develops into a vagina, a urethra, and a rectum.

[caption id="" align="aligncenter" width="350"]This flow chart shows how the sexual organs develop in embryos. The left side of the flow chart shows the development of male organs and the right side of the flow chart shows the development of female organs. Figure 1. Sexual Differentiation. Differentiation of the male and female reproductive systems does not occur until the fetal period of development.[/caption]

The Fetal Circulatory System

During prenatal development, the fetal circulatory system is integrated with the placenta via the umbilical cord so that the fetus receives both oxygen and nutrients from the placenta. However, after childbirth, the umbilical cord is severed, and the newborn’s circulatory system must be reconfigured. When the heart first forms in the embryo, it exists as two parallel tubes derived from mesoderm and lined with endothelium, which then fuse together. As the embryo develops into a fetus, the tube-shaped heart folds and further differentiates into the four chambers present in a mature heart. Unlike a mature cardiovascular system, however, the fetal cardiovascular system also includes circulatory shortcuts, or shunts. A shunt is an anatomical (or sometimes surgical) diversion that allows blood flow to bypass immature organs such as the lungs and liver until childbirth.

The placenta provides the fetus with necessary oxygen and nutrients via the umbilical vein. (Remember that veins carry blood toward the heart. In this case, the blood flowing to the fetal heart is oxygenated because it comes from the placenta. The respiratory system is immature and cannot yet oxygenate blood on its own.) From the umbilical vein, the oxygenated blood flows toward the inferior vena cava, all but bypassing the immature liver, via the ductus venosus shunt (Figure 2). The liver receives just a trickle of blood, which is all that it needs in its immature, semifunctional state. Blood flows from the inferior vena cava to the right atrium, mixing with fetal venous blood along the way.

Although the fetal liver is semifunctional, the fetal lungs are nonfunctional. The fetal circulation therefore bypasses the lungs by shifting some of the blood through the foramen ovale, a shunt that directly connects the right and left atria and avoids the pulmonary trunk altogether. Most of the rest of the blood is pumped to the right ventricle, and from there, into the pulmonary trunk, which splits into pulmonary arteries. However, a shunt within the pulmonary artery, the ductus arteriosus, diverts a portion of this blood into the aorta. This ensures that only a small volume of oxygenated blood passes through the immature pulmonary circuit, which has only minor metabolic requirements. Blood vessels of uninflated lungs have high resistance to flow, a condition that encourages blood to flow to the aorta, which presents much lower resistance. The oxygenated blood moves through the foramen ovale into the left atrium, where it mixes with the now deoxygenated blood returning from the pulmonary circuit. This blood then moves into the left ventricle, where it is pumped into the aorta. Some of this blood moves through the coronary arteries into the myocardium, and some moves through the carotid arteries to the brain.

The descending aorta carries partially oxygenated and partially deoxygenated blood into the lower regions of the body. It eventually passes into the umbilical arteries through branches of the internal iliac arteries. The deoxygenated blood collects waste as it circulates through the fetal body and returns to the umbilical cord. Thus, the two umbilical arteries carry blood low in oxygen and high in carbon dioxide and fetal wastes. This blood is filtered through the placenta, where wastes diffuse into the maternal circulation. Oxygen and nutrients from the mother diffuse into the placenta and from there into the fetal blood, and the process repeats.

[caption id="" align="aligncenter" width="525"]This figure shows a baby in the center of the image. To the left, is a panel showing the umbilical cord and how blood is supplied to the baby in the womb. Two panels on the right show the circulation of blood inside the baby’s body. Figure 2. Fetal Circulatory System. The fetal circulatory system includes three shunts to divert blood from undeveloped and partially functioning organs, as well as blood supply to and from the placenta.[/caption]

Other Organ Systems

During weeks 9–12 of fetal development, the brain continues to expand, the body elongates, and ossification continues. Fetal movements are frequent during this period, but are jerky and not well-controlled. The bone marrow begins to take over the process of erythrocyte production—a task that the liver performed during the embryonic period. The liver now secretes bile. The fetus circulates amniotic fluid by swallowing it and producing urine. The eyes are well-developed by this stage, but the eyelids are fused shut. The fingers and toes begin to develop nails. By the end of week 12, the fetus measures approximately 9 cm (3.5 in) from crown to rump.

Weeks 13–16 are marked by sensory organ development. The eyes move closer together; blinking motions begin, although the eyes remain sealed shut. The lips exhibit sucking motions. The ears move upward and lie flatter against the head. The scalp begins to grow hair. The excretory system is also developing: the kidneys are well-formed, and meconium, or fetal feces, begins to accumulate in the intestines. Meconium consists of ingested amniotic fluid, cellular debris, mucus, and bile.

During approximately weeks 16–20, as the fetus grows and limb movements become more powerful, the mother may begin to feel quickening, or fetal movements. However, space restrictions limit these movements and typically force the growing fetus into the “fetal position,” with the arms crossed and the legs bent at the knees. Sebaceous glands coat the skin with a waxy, protective substance called vernix caseosa that protects and moisturizes the skin and may provide lubrication during childbirth. A silky hair called lanugo also covers the skin during weeks 17–20, but it is shed as the fetus continues to grow. Extremely premature infants sometimes exhibit residual lanugo.

Developmental weeks 21–30 are characterized by rapid weight gain, which is important for maintaining a stable body temperature after birth. The bone marrow completely takes over erythrocyte synthesis, and the axons of the spinal cord begin to be myelinated, or coated in the electrically insulating glial cell sheaths that are necessary for efficient nervous system functioning. (The process of myelination is not completed until adolescence.) During this period, the fetus grows eyelashes. The eyelids are no longer fused and can be opened and closed. The lungs begin producing surfactant, a substance that reduces surface tension in the lungs and assists proper lung expansion after birth. Inadequate surfactant production in premature newborns may result in respiratory distress syndrome, and as a result, the newborn may require surfactant replacement therapy, supplemental oxygen, or maintenance in a continuous positive airway pressure (CPAP) chamber during their first days or weeks of life. In male fetuses, the testes descend into the scrotum near the end of this period. The fetus at 30 weeks measures 28 cm (11 in) from crown to rump and exhibits the approximate body proportions of a full-term newborn, but still is much leaner. [caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit this site for a summary of the stages of pregnancy, as experienced by the mother, and view the stages of development of the fetus throughout gestation.[/caption]

The fetus continues to lay down subcutaneous fat from week 31 until birth. The added fat fills out the hypodermis, and the skin transitions from red and wrinkled to soft and pink. Lanugo is shed, and the nails grow to the tips of the fingers and toes. Immediately before birth, the average crown-to-rump length is 35.5–40.5 cm (14–16 in), and the fetus weighs approximately 2.5–4 kg (5.5–8.8 lbs). Once born, the newborn is no longer confined to the fetal position, so subsequent measurements are made from head-to-toe instead of from crown-to-rump. At birth, the average length is approximately 51 cm (20 in).

Disorders of the…
Developing Fetus Throughout the second half of gestation, the fetal intestines accumulate a tarry, greenish black meconium. The newborn’s first stools consist almost entirely of meconium; they later transition to seedy yellow stools or slightly formed tan stools as meconium is cleared and replaced with digested breast milk or formula, respectively. Unlike these later stools, meconium is sterile; it is devoid of bacteria because the fetus is in a sterile environment and has not consumed any breast milk or formula. Typically, an infant does not pass meconium until after birth. However, in 5–20 percent of births, the fetus has a bowel movement in utero, which can cause major complications in the newborn.

The passage of meconium in the uterus signals fetal distress, particularly fetal hypoxia (i.e., oxygen deprivation). This may be caused by maternal drug abuse (especially tobacco or cocaine), maternal hypertension, depletion of amniotic fluid, long labor or difficult birth, or a defect in the placenta that prevents it from delivering adequate oxygen to the fetus. Meconium passage is typically a complication of full-term or post-term newborns because it is rarely passed before 34 weeks of gestation, when the gastrointestinal system has matured and is appropriately controlled by nervous system stimuli. Fetal distress can stimulate the vagus nerve to trigger gastrointestinal peristalsis and relaxation of the anal sphincter. Notably, fetal hypoxic stress also induces a gasping reflex, increasing the likelihood that meconium will be inhaled into the fetal lungs.

Although meconium is a sterile substance, it interferes with the antibiotic properties of the amniotic fluid and makes the newborn and mother more vulnerable to bacterial infections at birth and during the perinatal period. Specifically, inflammation of the fetal membranes, inflammation of the uterine lining, or neonatal sepsis (infection in the newborn) may occur. Meconium also irritates delicate fetal skin and can cause a rash.

The first sign that a fetus has passed meconium usually does not come until childbirth, when the amniotic sac ruptures. Normal amniotic fluid is clear and watery, but amniotic fluid in which meconium has been passed is stained greenish or yellowish. Antibiotics given to the mother may reduce the incidence of maternal bacterial infections, but it is critical that meconium is aspirated from the newborn before the first breath. Under these conditions, an obstetrician will extensively aspirate the infant’s airways as soon as the head is delivered, while the rest of the infant’s body is still inside the birth canal.

Aspiration of meconium with the first breath can result in labored breathing, a barrel-shaped chest, or a low Apgar score. An obstetrician can identify meconium aspiration by listening to the lungs with a stethoscope for a coarse rattling sound. Blood gas tests and chest X-rays of the infant can confirm meconium aspiration. Inhaled meconium after birth could obstruct a newborn’s airways leading to alveolar collapse, interfere with surfactant function by stripping it from the lungs, or cause pulmonary inflammation or hypertension. Any of these complications will make the newborn much more vulnerable to pulmonary infection, including pneumonia.

]]> 2807 2785 3 0 28.7 Patterns of Inheritance https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2825 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2825

Learning Objectives

By the end of this section, you will be able to:
  • Differentiate between genotype and phenotype
  • Describe how alleles determine a person’s traits
  • Summarize Mendel’s experiments and relate them to human genetics
  • Explain the inheritance of autosomal dominant and recessive and sex-linked genetic disorders

We have discussed the events that lead to the development of a newborn. But what makes each newborn unique? The answer lies, of course, in the DNA in the sperm and oocyte that combined to produce that first diploid cell, the human zygote.

From Genotype to Phenotype

Each human body cell has a full complement of DNA stored in 23 pairs of chromosomes. Figure 1 shows the pairs in a systematic arrangement called a karyotype. Among these is one pair of chromosomes, called the sex chromosomes, that determines the sex of the individual (XX in females, XY in males). The remaining 22 chromosome pairs are called autosomal chromosomes. Each of these chromosomes carries hundreds or even thousands of genes, each of which codes for the assembly of a particular protein—that is, genes are “expressed” as proteins. An individual’s complete genetic makeup is referred to as his or her genotype. The characteristics that the genes express, whether they are physical, behavioral, or biochemical, are a person’s phenotype.

You inherit one chromosome in each pair—a full complement of 23—from each parent. This occurs when the sperm and oocyte combine at the moment of your conception. Homologous chromosomes—those that make up a complementary pair—have genes for the same characteristics in the same location on the chromosome. Because one copy of a gene, an allele, is inherited from each parent, the alleles in these complementary pairs may vary. Take for example an allele that encodes for dimples. A child may inherit the allele encoding for dimples on the chromosome from the father and the allele that encodes for smooth skin (no dimples) on the chromosome from the mother.

[caption id="" align="aligncenter" width="480"]This figure show the 23 pairs of chromosomes in a male human being. Figure 1. Chromosomal Complement of a Male. Each pair of chromosomes contains hundreds to thousands of genes. The banding patterns are nearly identical for the two chromosomes within each pair, indicating the same organization of genes. As is visible in this karyotype, the only exception to this is the XY sex chromosome pair in males. (credit: National Human Genome Research Institute)[/caption]

Although a person can have two identical alleles for a single gene (a homozygous state), it is also possible for a person to have two different alleles (a heterozygous state). The two alleles can interact in several different ways. The expression of an allele can be dominant, for which the activity of this gene will mask the expression of a nondominant, or recessive, allele. Sometimes dominance is complete; at other times, it is incomplete. In some cases, both alleles are expressed at the same time in a form of expression known as codominance.

In the simplest scenario, a single pair of genes will determine a single heritable characteristic. However, it is quite common for multiple genes to interact to confer a feature. For instance, eight or more genes—each with their own alleles—determine eye color in humans. Moreover, although any one person can only have two alleles corresponding to a given gene, more than two alleles commonly exist in a population. This phenomenon is called multiple alleles. For example, there are three different alleles that encode ABO blood type; these are designated IA, IB, and i.

Over 100 years of theoretical and experimental genetics studies, and the more recent sequencing and annotation of the human genome, have helped scientists to develop a better understanding of how an individual’s genotype is expressed as their phenotype. This body of knowledge can help scientists and medical professionals to predict, or at least estimate, some of the features that an offspring will inherit by examining the genotypes or phenotypes of the parents. One important application of this knowledge is to identify an individual’s risk for certain heritable genetic disorders. However, most diseases have a multigenic pattern of inheritance and can also be affected by the environment, so examining the genotypes or phenotypes of a person’s parents will provide only limited information about the risk of inheriting a disease. Only for a handful of single-gene disorders can genetic testing allow clinicians to calculate the probability with which a child born to the two parents tested may inherit a specific disease.

Mendel’s Theory of Inheritance

Our contemporary understanding of genetics rests on the work of a nineteenth-century monk. Working in the mid-1800s, long before anyone knew about genes or chromosomes, Gregor Mendel discovered that garden peas transmit their physical characteristics to subsequent generations in a discrete and predictable fashion. When he mated, or crossed, two pure-breeding pea plants that differed by a certain characteristic, the first-generation offspring all looked like one of the parents. For instance, when he crossed tall and dwarf pure-breeding pea plants, all of the offspring were tall. Mendel called tallness dominant because it was expressed in offspring when it was present in a purebred parent. He called dwarfism recessive because it was masked in the offspring if one of the purebred parents possessed the dominant characteristic. Note that tallness and dwarfism are variations on the characteristic of height. Mendel called such a variation a trait. We now know that these traits are the expression of different alleles of the gene encoding height.

Mendel performed thousands of crosses in pea plants with differing traits for a variety of characteristics. And he repeatedly came up with the same results—among the traits he studied, one was always dominant, and the other was always recessive. (Remember, however, that this dominant–recessive relationship between alleles is not always the case; some alleles are codominant, and sometimes dominance is incomplete.)

Using his understanding of dominant and recessive traits, Mendel tested whether a recessive trait could be lost altogether in a pea lineage or whether it would resurface in a later generation. By crossing the second-generation offspring of purebred parents with each other, he showed that the latter was true: recessive traits reappeared in third-generation plants in a ratio of 3:1 (three offspring having the dominant trait and one having the recessive trait). Mendel then proposed that characteristics such as height were determined by heritable “factors” that were transmitted, one from each parent, and inherited in pairs by offspring.

In the language of genetics, Mendel’s theory applied to humans says that if an individual receives two dominant alleles, one from each parent, the individual’s phenotype will express the dominant trait. If an individual receives two recessive alleles, then the recessive trait will be expressed in the phenotype. Individuals who have two identical alleles for a given gene, whether dominant or recessive, are said to be homozygous for that gene (homo- = “same”). Conversely, an individual who has one dominant allele and one recessive allele is said to be heterozygous for that gene (hetero- = “different” or “other”). In this case, the dominant trait will be expressed, and the individual will be phenotypically identical to an individual who possesses two dominant alleles for the trait.

It is common practice in genetics to use capital and lowercase letters to represent dominant and recessive alleles. Using Mendel’s pea plants as an example, if a tall pea plant is homozygous, it will possess two tall alleles (TT). A dwarf pea plant must be homozygous because its dwarfism can only be expressed when two recessive alleles are present (tt). A heterozygous pea plant (Tt) would be tall and phenotypically indistinguishable from a tall homozygous pea plant because of the dominant tall allele. Mendel deduced that a 3:1 ratio of dominant to recessive would be produced by the random segregation of heritable factors (genes) when crossing two heterozygous pea plants. In other words, for any given gene, parents are equally likely to pass down either one of their alleles to their offspring in a haploid gamete, and the result will be expressed in a dominant–recessive pattern if both parents are heterozygous for the trait.

Because of the random segregation of gametes, the laws of chance and probability come into play when predicting the likelihood of a given phenotype. Consider a cross between an individual with two dominant alleles for a trait (AA) and an individual with two recessive alleles for the same trait (aa). All of the parental gametes from the dominant individual would be A, and all of the parental gametes from the recessive individual would be a (Figure 2). All of the offspring of that second generation, inheriting one allele from each parent, would have the genotype Aa, and the probability of expressing the phenotype of the dominant allele would be 4 out of 4, or 100 percent.

This seems simple enough, but the inheritance pattern gets interesting when the second-generation Aa individuals are crossed. In this generation, 50 percent of each parent’s gametes are A and the other 50 percent are a. By Mendel’s principle of random segregation, the possible combinations of gametes that the offspring can receive are AA, Aa, aA (which is the same as Aa), and aa. Because segregation and fertilization are random, each offspring has a 25 percent chance of receiving any of these combinations. Therefore, if an Aa × Aa cross were performed 1000 times, approximately 250 (25 percent) of the offspring would be AA; 500 (50 percent) would be Aa (that is, Aa plus aA); and 250 (25 percent) would be aa. The genotypic ratio for this inheritance pattern is 1:2:1. However, we have already established that AA and Aa (and aA) individuals all express the dominant trait (i.e., share the same phenotype), and can therefore be combined into one group. The result is Mendel’s third-generation phenotype ratio of 3:1.

[caption id="" align="aligncenter" width="350"]This diagram shows the genetics experiment conducted by Mendel. The top panel shows the offspring from first generation cross and the bottom panel shows the offspring from the second generation cross. Figure 2. Random Segregation. In the formation of gametes, it is equally likely that either one of a pair alleles from one parent will be passed on to the offspring. This figure follows the possible combinations of alleles through two generations following a first-generation cross of homozygous dominant and homozygous recessive parents. The recessive phenotype, which is masked in the second generation, has a 1 in 4, or 25 percent, chance of reappearing in the third generation.[/caption]

Mendel’s observation of pea plants also included many crosses that involved multiple traits, which prompted him to formulate the principle of independent assortment. The law states that the members of one pair of genes (alleles) from a parent will sort independently from other pairs of genes during the formation of gametes. Applied to pea plants, that means that the alleles associated with the different traits of the plant, such as color, height, or seed type, will sort independently of one another. This holds true except when two alleles happen to be located close to one other on the same chromosome. Independent assortment provides for a great degree of diversity in offspring.

Mendelian genetics represent the fundamentals of inheritance, but there are two important qualifiers to consider when applying Mendel’s findings to inheritance studies in humans. First, as we’ve already noted, not all genes are inherited in a dominant–recessive pattern. Although all diploid individuals have two alleles for every gene, allele pairs may interact to create several types of inheritance patterns, including incomplete dominance and codominance.

Secondly, Mendel performed his studies using thousands of pea plants. He was able to identify a 3:1 phenotypic ratio in second-generation offspring because his large sample size overcame the influence of variability resulting from chance. In contrast, no human couple has ever had thousands of children. If we know that a man and woman are both heterozygous for a recessive genetic disorder, we would predict that one in every four of their children would be affected by the disease. In real life, however, the influence of chance could change that ratio significantly. For example, if a man and a woman are both heterozygous for cystic fibrosis, a recessive genetic disorder that is expressed only when the individual has two defective alleles, we would expect one in four of their children to have cystic fibrosis. However, it is entirely possible for them to have seven children, none of whom is affected, or for them to have two children, both of whom are affected. For each individual child, the presence or absence of a single gene disorder depends on which alleles that child inherits from his or her parents.

Autosomal Dominant Inheritance

In the case of cystic fibrosis, the disorder is recessive to the normal phenotype. However, a genetic abnormality may be dominant to the normal phenotype. When the dominant allele is located on one of the 22 pairs of autosomes (non-sex chromosomes), we refer to its inheritance pattern as autosomal dominant. An example of an autosomal dominant disorder is neurofibromatosis type I, a disease that induces tumor formation within the nervous system that leads to skin and skeletal deformities. Consider a couple in which one parent is heterozygous for this disorder (and who therefore has neurofibromatosis), Nn, and one parent is homozygous for the normal gene, nn. The heterozygous parent would have a 50 percent chance of passing the dominant allele for this disorder to his or her offspring, and the homozygous parent would always pass the normal allele. Therefore, four possible offspring genotypes are equally likely to occur: Nn, Nn, nn, and nn. That is, every child of this couple would have a 50 percent chance of inheriting neurofibromatosis. This inheritance pattern is shown in Figure 3, in a form called a Punnett square, named after its creator, the British geneticist Reginald Punnett.

[caption id="" align="aligncenter" width="350"]This 2-by-2 Punnet square shows fifty percent dominant and fifty percent recessive offspring. Figure 3. Autosomal Dominant Inheritance. Inheritance pattern of an autosomal dominant disorder, such as neurofibromatosis, is shown in a Punnett square.[/caption]

Other genetic diseases that are inherited in this pattern are achondroplastic dwarfism, Marfan syndrome, and Huntington’s disease. Because autosomal dominant disorders are expressed by the presence of just one gene, an individual with the disorder will know that he or she has at least one faulty gene. The expression of the disease may manifest later in life, after the childbearing years, which is the case in Huntington’s disease (discussed in more detail later in this section).

Autosomal Recessive Inheritance

When a genetic disorder is inherited in an autosomal recessive pattern, the disorder corresponds to the recessive phenotype. Heterozygous individuals will not display symptoms of this disorder, because their unaffected gene will compensate. Such an individual is called a carrier. Carriers for an autosomal recessive disorder may never know their genotype unless they have a child with the disorder.

An example of an autosomal recessive disorder is cystic fibrosis (CF), which we introduced earlier. CF is characterized by the chronic accumulation of a thick, tenacious mucus in the lungs and digestive tract. Decades ago, children with CF rarely lived to adulthood. With advances in medical technology, the average lifespan in developed countries has increased into middle adulthood. CF is a relatively common disorder that occurs in approximately 1 in 2000 Caucasians. A child born to two CF carriers would have a 25 percent chance of inheriting the disease. This is the same 3:1 dominant:recessive ratio that Mendel observed in his pea plants would apply here. The pattern is shown in Figure 4, using a diagram that tracks the likely incidence of an autosomal recessive disorder on the basis of parental genotypes.

On the other hand, a child born to a CF carrier and someone with two unaffected alleles would have a 0 percent probability of inheriting CF, but would have a 50 percent chance of being a carrier. Other examples of autosome recessive genetic illnesses include the blood disorder sickle-cell anemia, the fatal neurological disorder Tay–Sachs disease, and the metabolic disorder phenylketonuria.

[caption id="" align="aligncenter" width="550"]In this figure, the offspring of a carrier father and carrier mother are shown. The first generation has one unaffected son, one affected daughter and one carrier son and one carrier daughter. The second generation cross shows seventy five percent unaffected and twenty five percent affected with cystic fibrosis. Figure 4. Autosomal Recessive Inheritance. The inheritance pattern of an autosomal recessive disorder with two carrier parents reflects a 3:1 probability of expression among offspring. (credit: U.S. National Library of Medicine)[/caption]

X-linked Dominant or Recessive Inheritance

An X-linked transmission pattern involves genes located on the X chromosome of the 23rd pair (Figure 5). Recall that a male has one X and one Y chromosome. When a father transmits a Y chromosome, the child is male, and when he transmits an X chromosome, the child is female. A mother can transmit only an X chromosome, as both her sex chromosomes are X chromosomes.

When an abnormal allele for a gene that occurs on the X chromosome is dominant over the normal allele, the pattern is described as X-linked dominant. This is the case with vitamin D–resistant rickets: an affected father would pass the disease gene to all of his daughters, but none of his sons, because he donates only the Y chromosome to his sons (see Figure 5a). If it is the mother who is affected, all of her children—male or female—would have a 50 percent chance of inheriting the disorder because she can only pass an X chromosome on to her children (see Figure 5b). For an affected female, the inheritance pattern would be identical to that of an autosomal dominant inheritance pattern in which one parent is heterozygous and the other is homozygous for the normal gene.

[caption id="" align="aligncenter" width="410"]This image shows the generations resulting from an X-linked dominant, affected father in the top panel and the generations resulting from an X-linked dominant, affected mother in the bottom panel. Figure 5. X-Linked Patterns of Inheritance. A chart of X-linked dominant inheritance patterns differs depending on whether (a) the father or (b) the mother is affected with the disease. (credit: U.S. National Library of Medicine)[/caption]

X-linked recessive inheritance is much more common because females can be carriers of the disease yet still have a normal phenotype. Diseases transmitted by X-linked recessive inheritance include color blindness, the blood-clotting disorder hemophilia, and some forms of muscular dystrophy. For an example of X-linked recessive inheritance, consider parents in which the mother is an unaffected carrier and the father is normal. None of the daughters would have the disease because they receive a normal gene from their father. However, they have a 50 percent chance of receiving the disease gene from their mother and becoming a carrier. In contrast, 50 percent of the sons would be affected (Figure 6).

With X-linked recessive diseases, males either have the disease or are genotypically normal—they cannot be carriers. Females, however, can be genotypically normal, a carrier who is phenotypically normal, or affected with the disease. A daughter can inherit the gene for an X-linked recessive illness when her mother is a carrier or affected, or her father is affected. The daughter will be affected by the disease only if she inherits an X-linked recessive gene from both parents. As you can imagine, X-linked recessive disorders affect many more males than females. For example, color blindness affects at least 1 in 20 males, but only about 1 in 400 females.

[caption id="" align="aligncenter" width="410"]This figure shows the offspring from a carrier mother with the X-linked recessive inheritance. Figure 6. X-Linked Recessive Inheritance. Given two parents in which the father is normal and the mother is a carrier of an X-linked recessive disorder, a son would have a 50 percent probability of being affected with the disorder, whereas daughters would either be carriers or entirely unaffected. (credit: U.S. National Library of Medicine)[/caption]

Other Inheritance Patterns: Incomplete Dominance, Codominance, and Lethal Alleles

Not all genetic disorders are inherited in a dominant–recessive pattern. In incomplete dominance, the offspring express a heterozygous phenotype that is intermediate between one parent’s homozygous dominant trait and the other parent’s homozygous recessive trait. An example of this can be seen in snapdragons when red-flowered plants and white-flowered plants are crossed to produce pink-flowered plants. In humans, incomplete dominance occurs with one of the genes for hair texture. When one parent passes a curly hair allele (the incompletely dominant allele) and the other parent passes a straight-hair allele, the effect on the offspring will be intermediate, resulting in hair that is wavy.

Codominance is characterized by the equal, distinct, and simultaneous expression of both parents’ different alleles. This pattern differs from the intermediate, blended features seen in incomplete dominance. A classic example of codominance in humans is ABO blood type. People are blood type A if they have an allele for an enzyme that facilitates the production of surface antigen A on their erythrocytes. This allele is designated IA. In the same manner, people are blood type B if they express an enzyme for the production of surface antigen B. People who have alleles for both enzymes (IA and IB) produce both surface antigens A and B. As a result, they are blood type AB. Because the effect of both alleles (or enzymes) is observed, we say that the IA and IB alleles are codominant. There is also a third allele that determines blood type. This allele (i) produces a nonfunctional enzyme. People who have two i alleles do not produce either A or B surface antigens: they have type O blood. If a person has IA and i alleles, the person will have blood type A. Notice that it does not make any difference whether a person has two IA alleles or one IA and one i allele. In both cases, the person is blood type A. Because IA masks i, we say that IA is dominant to i. Table 4 summarizes the expression of blood type.

Expression of Blood Types (Table 4)
Blood type Genotype Pattern of inheritance
A IAIA or IAi IAis dominant to i
B IBIB orIBi IB is dominant to i
AB IAIB IA is co-dominant to IB
O ii Two recessive alleles

Certain combinations of alleles can be lethal, meaning they prevent the individual from developing in utero, or cause a shortened life span. In recessive lethal inheritance patterns, a child who is born to two heterozygous (carrier) parents and who inherited the faulty allele from both would not survive. An example of this is Tay–Sachs, a fatal disorder of the nervous system. In this disorder, parents with one copy of the allele for the disorder are carriers. If they both transmit their abnormal allele, their offspring will develop the disease and will die in childhood, usually before age 5.

Dominant lethal inheritance patterns are much more rare because neither heterozygotes nor homozygotes survive. Of course, dominant lethal alleles that arise naturally through mutation and cause miscarriages or stillbirths are never transmitted to subsequent generations. However, some dominant lethal alleles, such as the allele for Huntington’s disease, cause a shortened life span but may not be identified until after the person reaches reproductive age and has children. Huntington’s disease causes irreversible nerve cell degeneration and death in 100 percent of affected individuals, but it may not be expressed until the individual reaches middle age. In this way, dominant lethal alleles can be maintained in the human population. Individuals with a family history of Huntington’s disease are typically offered genetic counseling, which can help them decide whether or not they wish to be tested for the faulty gene.

Mutations

A mutation is a change in the sequence of DNA nucleotides that may or may not affect a person’s phenotype. Mutations can arise spontaneously from errors during DNA replication, or they can result from environmental insults such as radiation, certain viruses, or exposure to tobacco smoke or other toxic chemicals. Because genes encode for the assembly of proteins, a mutation in the nucleotide sequence of a gene can change amino acid sequence and, consequently, a protein’s structure and function. Spontaneous mutations occurring during meiosis are thought to account for many spontaneous abortions (miscarriages).

Chromosomal Disorders

Sometimes a genetic disease is not caused by a mutation in a gene, but by the presence of an incorrect number of chromosomes. For example, Down syndrome is caused by having three copies of chromosome 21. This is known as trisomy 21. The most common cause of trisomy 21 is chromosomal nondisjunction during meiosis. The frequency of nondisjunction events appears to increase with age, so the frequency of bearing a child with Down syndrome increases in women over 36. The age of the father matters less because nondisjunction is much less likely to occur in a sperm than in an egg.

Whereas Down syndrome is caused by having three copies of a chromosome, Turner syndrome is caused by having just one copy of the X chromosome. This is known as monosomy. The affected child is always female. Women with Turner syndrome are sterile because their sexual organs do not mature.

Career Connections

Genetic Counselor Given the intricate orchestration of gene expression, cell migration, and cell differentiation during prenatal development, it is amazing that the vast majority of newborns are healthy and free of major birth defects. When a woman over 35 is pregnant or intends to become pregnant, or her partner is over 55, or if there is a family history of a genetic disorder, she and her partner may want to speak to a genetic counselor to discuss the likelihood that their child may be affected by a genetic or chromosomal disorder. A genetic counselor can interpret a couple’s family history and estimate the risks to their future offspring.

For many genetic diseases, a DNA test can determine whether a person is a carrier. For instance, carrier status for Fragile X, an X-linked disorder associated with mental retardation, or for cystic fibrosis can be determined with a simple blood draw to obtain DNA for testing. A genetic counselor can educate a couple about the implications of such a test and help them decide whether to undergo testing. For chromosomal disorders, the available testing options include a blood test, amniocentesis (in which amniotic fluid is tested), and chorionic villus sampling (in which tissue from the placenta is tested). Each of these has advantages and drawbacks. A genetic counselor can also help a couple cope with the news that either one or both partners is a carrier of a genetic illness, or that their unborn child has been diagnosed with a chromosomal disorder or other birth defect.

To become a genetic counselor, one needs to complete a 4-year undergraduate program and then obtain a Master of Science in Genetic Counseling from an accredited university. Board certification is attained after passing examinations by the American Board of Genetic Counseling. Genetic counselors are essential professionals in many branches of medicine, but there is a particular demand for preconception and prenatal genetic counselors.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit the National Society of Genetic Counselors website for more information about genetic counselors.[/caption]

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit the American Board of Genetic Counselors, Inc., website for more information about genetic counselors.[/caption]

]]> 2825 2785 7 0 1.3 Functions of Human Life https://pressbooks.bccampus.ca/apdouglas2017/chapter/1-3-functions-of-human-life/ Fri, 14 Jul 2017 21:50:10 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1844

Learning Objectives

By the end of this section, you will be able to:
  • Explain the importance of organization to the function of the human organism
  • Distinguish between metabolism, anabolism, and catabolism
  • Provide at least two examples of human responsiveness and human movement
  • Compare and contrast growth, differentiation, and reproduction

The different organ systems each have different functions and therefore unique roles to perform in physiology. These many functions can be summarized in terms of a few that we might consider definitive of human life: organization, metabolism, responsiveness, movement, development, and reproduction.

Organization

A human body consists of trillions of cells organized in a way that maintains distinct internal compartments. These compartments keep body cells separated from external environmental threats and keep the cells moist and nourished. They also separate internal body fluids from the countless microorganisms that grow on body surfaces, including the lining of certain tracts, or passageways. The intestinal tract, for example, is home to even more bacteria cells than the total of all human cells in the body, yet these bacteria are outside the body and cannot be allowed to circulate freely inside the body.

Cells, for example, have a cell membrane (also referred to as the plasma membrane) that keeps the intracellular environment—the fluids and organelles—separate from the extracellular environment. Blood vessels keep blood inside a closed circulatory system, and nerves and muscles are wrapped in connective tissue sheaths that separate them from surrounding structures. In the chest and abdomen, a variety of internal membranes keep major organs such as the lungs, heart, and kidneys separate from others.

The body’s largest organ system is the integumentary system, which includes the skin and its associated structures, such as hair and nails. The surface tissue of skin is a barrier that protects internal structures and fluids from potentially harmful microorganisms and other toxins.

Metabolism

The first law of thermodynamics holds that energy can neither be created nor destroyed—it can only change form. Your basic function as an organism is to consume (ingest) energy and molecules in the foods you eat, convert some of it into fuel for movement, sustain your body functions, and build and maintain your body structures. There are two types of reactions that accomplish this: anabolism and catabolism.

  • Anabolism is the process whereby smaller, simpler molecules are combined into larger, more complex substances. Your body can assemble, by utilizing energy, the complex chemicals it needs by combining small molecules derived from the foods you eat
  • Catabolism is the process by which larger more complex substances are broken down into smaller simpler molecules. Catabolism releases energy. The complex molecules found in foods are broken down so the body can use their parts to assemble the structures and substances needed for life.

Taken together, these two processes are called metabolism. Metabolism is the sum of all anabolic and catabolic reactions that take place in the body (Figure 1). Both anabolism and catabolism occur simultaneously and continuously to keep you alive.

[caption id="" align="aligncenter" width="300"]This illustration shows food entering a cell and being broken down into smaller particles of different colors. This is catabolism, which releases energy. In anabolism, the different colored particles are combined with each other to form larger, multi-colored structures. Anabolism requires an energy input. Figure 1. Metabolism. Anabolic reactions are building reactions, and they consume energy. Catabolic reactions break materials down and release energy. Metabolism includes both anabolic and catabolic reactions.[/caption]

Every cell in your body makes use of a chemical compound, adenosine triphosphate (ATP), to store and release energy. The cell stores energy in the synthesis (anabolism) of ATP, then moves the ATP molecules to the location where energy is needed to fuel cellular activities. Then the ATP is broken down (catabolism) and a controlled amount of energy is released, which is used by the cell to perform a particular job.

[caption id="attachment_2943" align="aligncenter" width="123"] Watch this CrashCourse video to learn more about metabolism![/caption]
[caption id="" align="aligncenter" width="120"]QR Code representing a URL View this animation to learn more about metabolic processes. What kind of catabolism occurs in the heart?[/caption]

Responsiveness

Responsiveness is the ability of an organism to adjust to changes in its internal and external environments. An example of responsiveness to external stimuli could include moving toward sources of food and water and away from perceived dangers. Changes in an organism’s internal environment, such as increased body temperature, can cause the responses of sweating and the dilation of blood vessels in the skin in order to decrease body temperature, as shown by the runners in Figure 2.

Movement

Human movement includes not only actions at the joints of the body, but also the motion of individual organs and even individual cells. As you read these words, red and white blood cells are moving throughout your body, muscle cells are contracting and relaxing to maintain your posture and to focus your vision, and glands are secreting chemicals to regulate body functions. Your body is coordinating the action of entire muscle groups to enable you to move air into and out of your lungs, to push blood throughout your body, and to propel the food you have eaten through your digestive tract. Consciously, of course, you contract your skeletal muscles to move the bones of your skeleton to get from one place to another (as the runners are doing in Figure 2), and to carry out all of the activities of your daily life.

[caption id="" align="aligncenter" width="480"]This photo shows three young men running in a competitive marathon. Figure 2. Marathon Runners. Runners demonstrate two characteristics of living humans—responsiveness and movement. Anatomic structures and physiological processes allow runners to coordinate the action of muscle groups and sweat in response to rising internal body temperature. (credit: Phil Roeder/flickr)[/caption]

Development, growth and reproduction

Development is all of the changes the body goes through in life. Development includes the process of differentiation, in which unspecialized cells become specialized in structure and function to perform certain tasks in the body. Development also includes the processes of growth and repair, both of which involve cell differentiation.

Growth is the increase in body size. Humans, like all multicellular organisms, grow by increasing the number of existing cells, increasing the amount of non-cellular material around cells (such as mineral deposits in bone), and, within very narrow limits, increasing the size of existing cells.

Reproduction is the formation of a new organism from parent organisms. In humans, reproduction is carried out by the male and female reproductive systems. Because death will come to all complex organisms, without reproduction, the line of organisms would end.

]]> 1844 1834 3 0 1.5 Homeostasis https://pressbooks.bccampus.ca/apdouglas2017/chapter/1-5-homeostasis/ Fri, 14 Jul 2017 21:55:44 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1851

Learning Objectives

By the end of this section, you will be able to:
  • Discuss the role of homeostasis in healthy functioning
  • Contrast negative and positive feedback, giving one physiologic example of each mechanism

Maintaining homeostasis requires that the body continuously monitor its internal conditions. From body temperature to blood pressure to levels of certain nutrients, each physiological condition has a particular set point. A set point is the physiological value around which the normal range fluctuates. A normal range is the restricted set of values that is optimally healthful and stable. For example, the set point for normal human body temperature is approximately 37°C (98.6°F) Physiological parameters, such as body temperature and blood pressure, tend to fluctuate within a normal range a few degrees above and below that point. Control centers in the brain and other parts of the body monitor and react to deviations from homeostasis using negative feedback. Negative feedback is a mechanism that reverses a deviation from the set point. Therefore, negative feedback maintains body parameters within their normal range. The maintenance of homeostasis by negative feedback goes on throughout the body at all times, and an understanding of negative feedback is thus fundamental to an understanding of human physiology.

Negative Feedback

A negative feedback system has three basic components (Figure 1a). A sensor, also referred to a receptor, is a component of a feedback system that monitors a physiological value. This value is reported to the control center. The control center is the component in a feedback system that compares the value to the normal range. If the value deviates too much from the set point, then the control center activates an effector. An effector is the component in a feedback system that causes a change to reverse the situation and return the value to the normal range.

[caption id="" align="aligncenter" width="450"]This figure shows three flow charts labeled A, B, and C. Chart A shows a general negative feedback loop. The loop starts with a stimulus. Information about the stimulus is perceived by a sensor which sends that information to a control center. The control center sends a signal to an effector, which then feeds back to the top of the flow chart by inhibiting the stimulus. Part B shows body temperature regulation as an example of negative feedback system. Here, the stimulus is body temperature exceeding 37 degrees Celsius. The sensor is a set of nerve cells in the skin and brain and the control center is the temperature regulatory center of the brain. The effectors are sweat glands throughout the body which inhibit the rising body temperature. Figure 1. Negative Feedback Loop. In a negative feedback loop, a stimulus—a deviation from a set point—is resisted through a physiological process that returns the body to homeostasis. (a) A negative feedback loop has four basic parts. (b) Body temperature is regulated by negative feedback.[/caption]

In order to set the system in motion, a stimulus must drive a physiological parameter beyond its normal range (that is, beyond homeostasis). This stimulus is “heard” by a specific sensor. For example, in the control of blood glucose, specific endocrine cells in the pancreas detect excess glucose (the stimulus) in the bloodstream. These pancreatic beta cells respond to the increased level of blood glucose by releasing the hormone insulin into the bloodstream. The insulin signals skeletal muscle fibers, fat cells (adipocytes), and liver cells to take up the excess glucose, removing it from the bloodstream. As glucose concentration in the bloodstream drops, the decrease in concentration—the actual negative feedback—is detected by pancreatic alpha cells, and insulin release stops. This prevents blood sugar levels from continuing to drop below the normal range.

Humans have a similar temperature regulation feedback system that works by promoting either heat loss or heat gain (Figure 1b). When the brain’s temperature regulation center receives data from the sensors indicating that the body’s temperature exceeds its normal range, it stimulates a cluster of brain cells referred to as the “heat-loss center.” This stimulation has three major effects:

  • Blood vessels in the skin begin to dilate allowing more blood from the body core to flow to the surface of the skin allowing the heat to radiate into the environment.
  • As blood flow to the skin increases, sweat glands are activated to increase their output. As the sweat evaporates from the skin surface into the surrounding air, it takes heat with it.
  • The depth of respiration increases, and a person may breathe through an open mouth instead of through the nasal passageways. This further increases heat loss from the lungs.

In contrast, activation of the brain’s heat-gain center by exposure to cold reduces blood flow to the skin, and blood returning from the limbs is diverted into a network of deep veins. This arrangement traps heat closer to the body core and restricts heat loss. If heat loss is severe, the brain triggers an increase in random signals to skeletal muscles, causing them to contract and producing shivering. The muscle contractions of shivering release heat while using up ATP. The brain triggers the thyroid gland in the endocrine system to release thyroid hormone, which increases metabolic activity and heat production in cells throughout the body. The brain also signals the adrenal glands to release epinephrine (adrenaline), a hormone that causes the breakdown of glycogen into glucose, which can be used as an energy source. The breakdown of glycogen into glucose also results in increased metabolism and heat production.

[caption id="" align="aligncenter" width="119"]QR Code representing a URL Watch this video to learn more about water concentration in the body.[/caption] [caption id="attachment_2946" align="aligncenter" width="119"] Watch this CrashCourse video to learn more about homeostasis![/caption]

Positive Feedback

Positive feedback intensifies a change in the body’s physiological condition rather than reversing it. A deviation from the normal range results in more change, and the system moves farther away from the normal range. Positive feedback in the body is normal only when there is a definite end point. Childbirth and the body’s response to blood loss are two examples of positive feedback loops that are normal but are activated only when needed.

Childbirth at full term is an example of a situation in which the maintenance of the existing body state is not desired. Enormous changes in the mother’s body are required to expel the baby at the end of pregnancy. And the events of childbirth, once begun, must progress rapidly to a conclusion or the life of the mother and the baby are at risk. The extreme muscular work of labor and delivery are the result of a positive feedback system (Figure 2).

[caption id="" align="aligncenter" width="380"]This diagram shows the steps of a positive feedback loop as a series of stepwise arrows looping around a diagram of an infant within the uterus of a pregnant woman. Initially the head of the baby pushes against the cervix, transmitting nerve impulses from the cervix to the brain. Next the brain stimulates the pituitary gland to secrete oxytocin which is carried in the bloodstream to the uterus. Finally, the oxytocin simulates uterine contractions and pushes the baby harder into the cervix. As the head of the baby pushes against the cervix with greater and greater force, the uterine contractions grow stronger and more frequent. This mechanism is a positive feedback loop. Figure 2. Positive Feedback Loop. Normal childbirth is driven by a positive feedback loop. A positive feedback loop results in a change in the body’s status, rather than a return to homeostasis.[/caption]
The first contractions of labor (the stimulus) push the baby toward the cervix (the lowest part of the uterus). The cervix contains stretch-sensitive nerve cells that monitor the degree of stretching (the sensors). These nerve cells send messages to the brain, which in turn causes the pituitary gland at the base of the brain to release the hormone oxytocin into the bloodstream. Oxytocin causes stronger contractions of the smooth muscles in of the uterus (the effectors), pushing the baby further down the birth canal. This causes even greater stretching of the cervix. The cycle of stretching, oxytocin release, and increasingly more forceful contractions stops only when the baby is born. At this point, the stretching of the cervix halts, stopping the release of oxytocin.

A second example of positive feedback centers on reversing extreme damage to the body. Following a penetrating wound, the most immediate threat is excessive blood loss. Less blood circulating means reduced blood pressure and reduced perfusion (penetration of blood) to the brain and other vital organs. If perfusion is severely reduced, vital organs will shut down and the person will die. The body responds to this potential catastrophe by releasing substances in the injured blood vessel wall that begin the process of blood clotting. As each step of clotting occurs, it stimulates the release of more clotting substances. This accelerates the processes of clotting and sealing off the damaged area. Clotting is contained in a local area based on the tightly controlled availability of clotting proteins. This is an adaptive, life-saving cascade of events.

]]> 1851 1834 5 0 2.2 Chemical Bonds https://pressbooks.bccampus.ca/apdouglas2017/chapter/2-2-chemical-bonds/ Fri, 14 Jul 2017 22:02:30 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1916

Ions and Ionic Bonds

Recall that an atom typically has the same number of positively charged protons and negatively charged electrons. As long as this situation remains, the atom is electrically neutral. But when an atom participates in a chemical reaction that results in the donation or acceptance of one or more electrons, the atom will then become positively or negatively charged. This happens frequently for most atoms in order to have a full valence shell, as described previously. This can happen either by gaining electrons to fill a shell that is more than half-full, or by giving away electrons to empty a shell than is less than half-full, thereby leaving the next smaller electron shell as the new, full, valence shell. An atom that has an electrical charge—whether positive or negative—is an ion.

[caption id="" align="alignright" width="120"]QR Code representing a URL Visit this website to learn about electrical energy and the attraction/repulsion of charges.[/caption]

Potassium (K), for instance, is an important element in all body cells. Its atomic number is 19. It has just one electron in its valence shell. This characteristic makes potassium highly likely to participate in chemical reactions in which it donates one electron. (It is easier for potassium to donate one electron than to gain seven electrons.) The loss will cause the positive charge of potassium’s protons to be more influential than the negative charge of potassium’s electrons. In other words, the resulting potassium ion will be slightly positive. A potassium ion is written K+, indicating that it has lost a single electron. A positively charged ion is known as a cation.

Now consider fluorine (F), a component of bones and teeth. Its atomic number is nine, and it has seven electrons in its valence shell. Thus, it is highly likely to bond with other atoms in such a way that fluorine accepts one electron (it is easier for fluorine to gain one electron than to donate seven electrons). When it does, its electrons will outnumber its protons by one, and it will have an overall negative charge. The ionized form of fluorine is called fluoride, and is written as F–. A negatively charged ion is known as an anion.

Atoms that have more than one electron to donate or accept will end up with stronger positive or negative charges. A cation that has donated two electrons has a net charge of +2. Using magnesium (Mg) as an example, this can be written Mg++ or Mg2+. An anion that has accepted two electrons has a net charge of –2. The ionic form of selenium (Se), for example, is typically written Se2–.

[caption id="" align="alignright" width="420"]The top panel of this figure shows the orbit model of a sodium atom and a chlorine atom and arrows pointing towards the transfer of electrons from sodium to chlorine to form sodium and chlorine ions. The bottom panel shows sodium and chloride ions in a crystal structure. Figure 1. Ionic Bonding. (a) Sodium readily donates the solitary electron in its valence shell to chlorine, which needs only one electron to have a full valence shell. (b) The opposite electrical charges of the resulting sodium cation and chloride anion result in the formation of a bond of attraction called an ionic bond. (c) The attraction of many sodium and chloride ions results in the formation of large groupings called crystals.[/caption]

The opposite charges of cations and anions exert a moderately strong mutual attraction that keeps the atoms in close proximity forming an ionic bond. An ionic bond is an ongoing, close association between ions of opposite charge. The table salt you sprinkle on your food owes its existence to ionic bonding. As shown in Figure 1, sodium commonly donates an electron to chlorine, becoming the cation Na+. When chlorine accepts the electron, it becomes the chloride anion, Cl–. With their opposing charges, these two ions strongly attract each other.

Water is an essential component of life because it is able to break the ionic bonds in salts to free the ions. In fact, in biological fluids, most individual atoms exist as ions. These dissolved ions produce electrical charges within the body. The behavior of these ions produces the tracings of heart and brain function observed as waves on an electrocardiogram (EKG or ECG) or an electroencephalogram (EEG). The electrical activity that derives from the interactions of the charged ions is why they are also called electrolytes.

Covalent Bonds

Unlike ionic bonds formed by the attraction between a cation’s positive charge and an anion’s negative charge, molecules formed by a covalent bond share electrons in a mutually stabilizing relationship. Like next-door neighbors whose kids hang out first at one home and then at the other, the atoms do not lose or gain electrons permanently. Instead, the electrons move back and forth between the elements. Because of the close sharing of pairs of electrons (one electron from each of two atoms), covalent bonds are stronger than ionic bonds.

Nonpolar Covalent Bonds

Figure 2 shows several common types of covalent bonds. Notice that the two covalently bonded atoms typically share just one or two electron pairs, though larger sharings are possible. The important concept to take from this is that in covalent bonds, electrons in the outermost valence shell are shared to fill the valence shells of both atoms, ultimately stabilizing both of the atoms involved. In a single covalent bond, a single electron is shared between two atoms, while in a double covalent bond, two pairs of electrons are shared between two atoms. There even are triple covalent bonds, where three atoms are shared.
[caption id="" align="alignleft" width="550"]The top panel in this figure shows two hydrogen atoms sharing two electrons. The middle panel shows two oxygen atoms sharing four electrons, and the bottom panel shows two oxygen atoms and one carbon atom sharing 2 pairs of electrons each. Figure 2. Covalent Bonding.[/caption]
You can see that the covalent bonds shown in Figure 2 are balanced. The sharing of the negative electrons is relatively equal, as is the electrical pull of the positive protons in the nucleus of the atoms involved. This is why covalently bonded molecules that are electrically balanced in this way are described as nonpolar; that is, no region of the molecule is either more positive or more negative than any other.

Polar Covalent Bonds

Groups of legislators with completely opposite views on a particular issue are often described as “polarized” by news writers. In chemistry, a polar molecule is a molecule that contains regions that have opposite electrical charges. Polar molecules occur when atoms share electrons unequally, in polar covalent bonds.

The most familiar example of a polar molecule is water (Figure 3). The molecule has three parts: one atom of oxygen, the nucleus of which contains eight protons, and two hydrogen atoms, whose nuclei each contain only one proton. Because every proton exerts an identical positive charge, a nucleus that contains eight protons exerts a charge eight times greater than a nucleus that contains one proton. This means that the negatively charged electrons present in the water molecule are more strongly attracted to the oxygen nucleus than to the hydrogen nuclei. Each hydrogen atom’s single negative electron therefore migrates toward the oxygen atom, making the oxygen end of their bond slightly more negative than the hydrogen end of their bond.

[caption id="" align="alignleft" width="380"]This figure shows the structure of a water molecule. The top panel shows two oxygen atoms and one hydrogen atom with electrons in orbit and the shared electrons. The middle panel shows a three-dimensional model of a water molecule and the bottom panel shows the structural formula for water. Figure 3. Polar Covalent Bonds in a Water Molecule.[/caption]

What is true for the bonds is true for the water molecule as a whole; that is, the oxygen region has a slightly negative charge and the regions of the hydrogen atoms have a slightly positive charge. These charges are often referred to as “partial charges” because the strength of the charge is less than one full electron, as would occur in an ionic bond. As shown in Figure 3, regions of weak polarity are indicated with the Greek letter delta (∂) and a plus (+) or minus (–) sign.

Even though a single water molecule is unimaginably tiny, it has mass, and the opposing electrical charges on the molecule pull that mass in such a way that it creates a shape somewhat like a triangular tent (see Figure 3b). The resulting dipole, with the positive charges at one end formed by the hydrogen atoms at the “bottom” of the tent and the negative charge at the opposite end (the oxygen atom at the “top” of the tent) makes the charged regions highly likely to interact with charged regions of other polar molecules. For human physiology, the resulting bond is one of the most important formed by water—the hydrogen bond.

Hydrogen Bonds

A hydrogen bond is formed when a weakly positive hydrogen atom already bonded to one electronegative atom (for example, the oxygen in the water molecule) is attracted to another electronegative atom from another molecule. In other words, hydrogen bonds always include hydrogen that is already part of a polar molecule.

The most common example of hydrogen bonding in the natural world occurs between molecules of water. It happens before your eyes whenever two raindrops merge into a larger bead, or a creek spills into a river. Hydrogen bonding occurs because the weakly negative oxygen atom in one water molecule is attracted to the weakly positive hydrogen atoms of two other water molecules (Figure 4).

[caption id="" align="alignright" width="280"]This figure shows three water molecules and the hydrogen bonds between them. Figure 4. Hydrogen Bonds between Water Molecules. Notice that the bonds occur between the weakly positive charge on the hydrogen atoms and the weakly negative charge on the oxygen atoms. Hydrogen bonds are relatively weak, and therefore are indicated with a dotted (rather than a solid) line.[/caption]

Water molecules also strongly attract other types of charged molecules as well as ions. This explains why “table lt,” for example, actually is a molecule called a “salt” in chemistry, which consists of equal numbers of positively-charged sodium (Na+) and negatively-charged chloride (Cl–), dissolves so readily in water, in this case forming dipole-ion bonds between the water and the electrically-charged ions (electrolytes). Water molecules also repel molecules with nonpolar covalent bonds, like fats, lipids, and oils. You can demonstrate this with a simple kitchen experiment: pour a teaspoon of vegetable oil, a compound formed by nonpolar covalent bonds, into a glass of water. Instead of instantly dissolving in the water, the oil forms a distinct bead because the polar water molecules repel the nonpolar oil.

[caption id="attachment_2948" align="aligncenter" width="150"] Watch this SciShow video to learn more about electrostatic forces![/caption] [caption id="attachment_2949" align="aligncenter" width="150"] Watch this CrashCourse video to learn more bonding![/caption]
]]> 1916 1901 3 0 2.4 Inorganic Compounds Essential to Human Functioning https://pressbooks.bccampus.ca/apdouglas2017/chapter/2-4-inorganic-compounds-essential-to-human-functioning/ Fri, 14 Jul 2017 22:07:16 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1921

Water

As much as 70 percent of an adult’s body weight is water. This water is contained both within the cells and between the cells that make up tissues and organs. Its several roles make water indispensable to human functioning.

Water as a Lubricant and Cushion

Water is a major component of many of the body’s lubricating fluids. Just as oil lubricates the hinge on a door, water in synovial fluid lubricates the actions of body joints, and water in pleural fluid helps the lungs expand and recoil with breathing. Watery fluids help keep food flowing through the digestive tract, and ensure that the movement of adjacent abdominal organs is friction free.

Water also protects cells and organs from physical trauma, cushioning the brain within the skull, for example, and protecting the delicate nerve tissue of the eyes. Water cushions a developing fetus in the mother’s womb as well.

Water as a Heat Sink

A heat sink is a substance or object that absorbs and dissipates heat but does not experience a corresponding increase in temperature. In the body, water absorbs the heat generated by chemical reactions without greatly increasing in temperature. Moreover, when the environmental temperature soars, the water stored in the body helps keep the body cool. This cooling effect happens as warm blood from the body’s core flows to the blood vessels just under the skin and is transferred to the environment. At the same time, sweat glands release warm water in sweat. As the water evaporates into the air, it carries away heat, and then the cooler blood from the periphery circulates back to the body core.

Water as a Component of Liquid Mixtures

A mixture is a combination of two or more substances, each of which maintains its own chemical identity. In other words, the constituent substances are not chemically bonded into a new, larger chemical compound. The concept is easy to imagine if you think of powdery substances such as flour and sugar; when you stir them together in a bowl, they obviously do not bond to form a new compound. The room air you breathe is a gaseous mixture, containing three discrete elements—nitrogen, oxygen, and argon—and one compound, carbon dioxide. There are three types of liquid mixtures, all of which contain water as a key component. These are solutions, colloids, and suspensions.

For cells in the body to survive, they must be kept moist in a water-based liquid called a solution. In chemistry, a liquid solution consists of a solvent that dissolves a substance called a solute. An important characteristic of solutions is that they are homogeneous; that is, the solute molecules are distributed evenly throughout the solution. If you were to stir a teaspoon of sugar into a glass of water, the sugar would dissolve into sugar molecules separated by water molecules. The ratio of sugar to water in the left side of the glass would be the same as the ratio of sugar to water in the right side of the glass. If you were to add more sugar, the ratio of sugar to water would change, but the distribution—provided you had stirred well—would still be even.

Water is considered the “universal solvent” and it is believed that life cannot exist without water because of this. Water is certainly the most abundant solvent in the body; essentially all of the body’s chemical reactions occur among compounds dissolved in water. Because water molecules are polar, with regions of positive and negative electrical charge, water readily dissolves ionic compounds and polar covalent compounds. Such compounds are referred to as hydrophilic, or “water-loving.” As mentioned above, sugar dissolves well in water. This is because sugar molecules contain regions of hydrogen-oxygen polar bonds, making it hydrophilic. Nonpolar molecules, which do not readily dissolve in water, are called hydrophobic, or “water-fearing.”

The Role of Water in Chemical Reactions

[caption id="" align="alignright" width="520"]The top panel in this figure shows a dehydration-synthesis reaction, and the bottom panel shows a hydrolysis reaction. Figure 1. Dehydration Synthesis and Hydrolysis. Monomers, the basic units for building larger molecules, form polymers (two or more chemically-bonded monomers). (a) In dehydration synthesis, two monomers are covalently bonded in a reaction in which one gives up a hydroxyl group and the other a hydrogen atom. A molecule of water is released as a byproduct during dehydration reactions. (b) In hydrolysis, the covalent bond between two monomers is split by the addition of a hydrogen atom to one and a hydroxyl group to the other, which requires the contribution of one molecule of water.[/caption]

Two types of chemical reactions involve the creation or the consumption of water: dehydration synthesis and hydrolysis.

  • In dehydration synthesis, one reactant gives up an atom of hydrogen and another reactant gives up a hydroxyl group (OH) in the synthesis of a new product. In the formation of their covalent bond, a molecule of water is released as a byproduct (Figure 1). This is also sometimes referred to as a condensation reaction.
  • In hydrolysis, a molecule of water disrupts a compound, breaking its bonds. The water is itself split into H and OH. One portion of the severed compound then bonds with the hydrogen atom, and the other portion bonds with the hydroxyl group.
These reactions are reversible, and play an important role in the chemistry of organic compounds (which will be discussed shortly).
[caption id="attachment_3037" align="aligncenter" width="150"] Watch this amoeba sisters video to learn more about the properties of water![/caption]
[caption id="attachment_2951" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about the importance of water and its chemical properties.[/caption]

Salts

[caption id="" align="alignright" width="480"]This figure shows a crystal lattice of sodium chloride interacting with water to form a hydrated sodium ion and a hydrated chloride ion. Figure 2. Dissociation of Sodium Chloride in Water. Notice that the crystals of sodium chloride dissociate not into molecules of NaCl, but into Na+ cations and Cl– anions, each completely surrounded by water molecules.[/caption] Recall that salts are formed when ions form ionic bonds. In these reactions, one atom gives up one or more electrons, and thus becomes positively charged, whereas the other accepts one or more electrons and becomes negatively charged. You can now define a salt as a substance that, when dissolved in water, dissociates into ions other than H+ or OH–. This fact is important in distinguishing salts from acids and bases, discussed next.

A typical salt, NaCl, dissociates completely in water (Figure 2). The positive and negative regions on the water molecule (the hydrogen and oxygen ends respectively) attract the negative chloride and positive sodium ions, pulling them away from each other. Again, whereas nonpolar and polar covalently bonded compounds break apart into molecules in solution, salts dissociate into ions. These ions are electrolytes; they are capable of conducting an electrical current in solution. This property is critical to the function of ions in transmitting nerve impulses and prompting muscle contraction.

Many other salts are important in the body. For example, bile salts produced by the liver help break apart dietary fats, and calcium phosphate salts form the mineral portion of teeth and bones.

Acids and Bases

Acids and bases, like salts, dissociate in water into electrolytes. Acids and bases can very much change the properties of the solutions in which they are dissolved.

Acids

An acid is a substance that releases hydrogen ions (H+) in solution (Figure 3a). Because an atom of hydrogen has just one proton and one electron, a positively charged hydrogen ion is simply a proton. This solitary proton is highly likely to participate in chemical reactions. Strong acids are compounds that release all of their H+ in solution; that is, they ionize completely. Hydrochloric acid (HCl), which is released from cells in the lining of the stomach, is a strong acid because it releases all of its H+ in the stomach’s watery environment. This strong acid aids in digestion and kills ingested microbes. Weak acids do not ionize completely; that is, some of their hydrogen ions remain bonded within a compound in solution. An example of a weak acid is vinegar, or acetic acid; and it is called acetate after it gives up a proton.
[caption id="" align="alignleft" width="420"]This figure shows four beakers containing different liquids. Figure 3. Acids and Bases. (a) In aqueous solution, an acid dissociates into hydrogen ions (H+) and anions. Nearly every molecule of a strong acid dissociates, producing a high concentration of H+. (b) In aqueous solution, a base dissociates into hydroxyl ions (OH–) and cations. Nearly every molecule of a strong base dissociates, producing a high concentration of OH–.[/caption]

Bases

A base is a substance that releases hydroxyl ions (OH–) in solution, or one that accepts H+ already present in solution (see Figure 3b). The hydroxyl ions (also known as hydroxide ions) or other basic substances combine with H+ present to form a water molecule, thereby removing H+ and reducing the solution’s acidity. Strong bases release most or all of their hydroxyl ions; weak bases release only some hydroxyl ions or absorb only a few H+. Food mixed with hydrochloric acid from the stomach would burn the small intestine, the next portion of the digestive tract after the stomach, if it were not for the release of bicarbonate (HCO3–), a weak base that attracts H+. Bicarbonate accepts some of the H+ protons, thereby reducing the acidity of the solution.

The Concept of pH

The relative acidity or alkalinity of a solution can be indicated by its pH. A solution’s pH is the negative, base-10 logarithm of the hydrogen ion (H+) concentration of the solution. As an example, a pH 4 solution has an H+ concentration that is ten times greater than that of a pH 5 solution. That is, a solution with a pH of 4 is ten times more acidic than a solution with a pH of 5. The concept of pH will begin to make more sense when you study the pH scale. The scale consists of a series of increments ranging from 0 to 14. A solution with a pH of 7 is considered neutral—neither acidic nor basic. Pure water has a pH of 7. The lower the number below 7, the more acidic the solution, or the greater the concentration of H+. The concentration of hydrogen ions at each pH value is 10 times different than the next pH. For instance, a pH value of 4 corresponds to a proton concentration of 10–4 M, or 0.0001M, while a pH value of 5 corresponds to a proton concentration of 10–5 M, or 0.00001M. The higher the number above 7, the more basic (alkaline) the solution, or the lower the concentration of H+. Human urine, for example, is ten times more acidic than pure water, and HCl is 10,000,000 times more acidic than water.
[caption id="" align="alignright" width="320"]This figure shows a vertical arrow with the top half showing the basic scale and the bottom half showing the acidic scale. Different chemicals and their pH are also shown. Figure 4. The pH Scale[/caption]

Buffers

The pH of human blood normally ranges from 7.35 to 7.45, although it is typically identified as pH 7.4. At this slightly basic pH, blood can reduce the acidity resulting from the carbon dioxide (CO2) constantly being released into the bloodstream by the trillions of cells in the body. Homeostatic mechanisms (along with exhaling CO2 while breathing) normally keep the pH of blood within this narrow range. This is critical, because fluctuations—either too acidic or too alkaline—can lead to life-threatening disorders.

All cells of the body depend on homeostatic regulation of acid–base balance at a pH of approximately 7.4. The body therefore has several mechanisms for this regulation, involving breathing, the excretion of chemicals in urine, and the internal release of chemicals collectively called buffers into body fluids. A buffer is a solution of a weak acid and its conjugate base. A buffer can neutralize small amounts of acids or bases in body fluids. For example, if there is even a slight decrease below 7.35 in the pH of a bodily fluid, the buffer in the fluid—in this case, acting as a weak base—will bind the excess hydrogen ions. In contrast, if pH rises above 7.45, the buffer will act as a weak acid and contribute hydrogen ions.

[caption id="attachment_2952" align="aligncenter" width="150"] For a more complex explanation of buffers, watch this CrashCourse video on acids, bases, and buffers.[/caption]
]]> 1921 1901 4 0 3.1 The Cell Membrane https://pressbooks.bccampus.ca/apdouglas2017/chapter/3-1-the-cell-membrane/ Fri, 14 Jul 2017 22:13:23 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1950

By the end of this section, you will be able to:
  • Describe the molecular components that make up the cell membrane
  • Explain the major features and properties of the cell membrane
  • Differentiate between materials that can and cannot diffuse through the lipid bilayer
  • Compare and contrast different types of passive transport with active transport, providing examples of each

Despite differences in structure and function, all living cells in multicellular organisms have a surrounding cell membrane. As the outer layer of your skin separates your body from its environment, the cell membrane (also known as the plasma membrane) separates the inner contents of a cell from its exterior environment. This cell membrane provides a protective barrier around the cell and regulates which materials can pass in or out.

Structure and Composition of the Cell Membrane

The cell membrane is an extremely pliable structure composed primarily of back-to-back phospholipids (a “bilayer”). Cholesterol is also present, which contributes to the fluidity of the membrane, and there are various proteins embedded within the membrane that have a variety of functions.

[caption id="" align="alignleft" width="280"]This diagram shows the structure of a phospholipid. The hydrophilic head group is shown as a pink sphere and the two tails are shown as yellow rectangles. Figure 1. Phospholipid Structure. A phospholipid molecule consists of a polar phosphate “head,” which is hydrophilic and a non-polar lipid “tail,” which is hydrophobic. Unsaturated fatty acids result in kinks in the hydrophobic tails.[/caption] A single phospholipid molecule has a phosphate group on one end, called the “head,” and two side-by-side chains of fatty acids that make up the lipid tails (Figure 1). The phosphate group is negatively charged, making the head polar and hydrophilic—or “water loving.” A hydrophilic molecule (or region of a molecule) is one that is attracted to water. The phosphate heads are thus attracted to the water molecules of both the extracellular and intracellular environments. The lipid tails, on the other hand, are uncharged, or nonpolar, and are hydrophobic—or “water fearing.” A hydrophobic molecule (or region of a molecule) repels and is repelled by water. Some lipid tails consist of saturated fatty acids and some contain unsaturated fatty acids. This combination adds to the fluidity of the tails that are constantly in motion. Phospholipids are thus amphipathic molecules. An amphipathic molecule is one that contains both a hydrophilic and a hydrophobic region. In fact, soap works to remove oil and grease stains because it has amphipathic properties. The hydrophilic portion can dissolve in water while the hydrophobic portion can trap grease in micelles that then can be washed away.
[caption id="" align="alignright" width="280"]This diagram shows a phospholipid bilayer. Two sets of phospholipids are arranged such that the hydrophobic tails are facing each other and the hydrophilic heads are facing the extracellular environment. Figure 2. Phospolipid Bilayer. The phospholipid bilayer consists of two adjacent sheets of phospholipids, arranged tail to tail. The hydrophobic tails associate with one another, forming the interior of the membrane. The polar heads contact the fluid inside and outside of the cell.[/caption] The cell membrane consists of two adjacent layers of phospholipids. The lipid tails of one layer face the lipid tails of the other layer, meeting at the interface of the two layers. The phospholipid heads face outward, one layer exposed to the interior of the cell and one layer exposed to the exterior (Figure 2). Because the phosphate groups are polar and hydrophilic, they are attracted to water in the intracellular fluid. Intracellular fluid (ICF) is the fluid interior of the cell. The phosphate groups are also attracted to the extracellular fluid. Extracellular fluid (ECF) is the fluid environment outside the enclosure of the cell membrane. Interstitial fluid (IF) is the term given to extracellular fluid not contained within blood vessels. Because the lipid tails are hydrophobic, they meet in the inner region of the membrane, excluding watery intracellular and extracellular fluid from this space. The cell membrane has many proteins, as well as other lipids (such as cholesterol), that are associated with the phospholipid bilayer. An important feature of the membrane is that it remains fluid; the lipids and proteins in the cell membrane are not rigidly locked in place.

Membrane Proteins

The lipid bilayer forms the basis of the cell membrane, but it is peppered throughout with various proteins. Two different types of proteins that are commonly associated with the cell membrane are the integral proteins and peripheral protein (Figure 3). As its name suggests, an integral protein is a protein that is embedded in the membrane. A channel protein is an example of an integral protein that selectively allows particular materials, such as certain ions, to pass into or out of the cell.

[caption id="" align="alignright" width="495"]This image shows a lipid bilayer with different types of proteins, lipids and cholesterol embedded in it. Figure 3. Cell Membrane. The cell membrane of the cell is a phospholipid bilayer containing many different molecular components, including proteins and cholesterol, some with carbohydrate groups attached.[/caption]

Another important group of integral proteins are cell recognition proteins, which serve to mark a cell’s identity so that it can be recognized by other cells. A receptor is a type of recognition protein that can selectively bind a specific molecule outside the cell, and this binding induces a chemical reaction within the cell. A ligand is the specific molecule that binds to and activates a receptor. Some integral proteins serve dual roles as both a receptor and an ion channel. One example of a receptor-ligand interaction is the receptors on nerve cells that bind neurotransmitters, such as dopamine. When a dopamine molecule binds to a dopamine receptor protein, a channel within the transmembrane protein opens to allow certain ions to flow into the cell.

Some integral membrane proteins are glycoproteins. A glycoprotein is a protein that has carbohydrate molecules attached, which extend into the extracellular matrix. The attached carbohydrate tags on glycoproteins aid in cell recognition. The carbohydrates that extend from membrane proteins and even from some membrane lipids collectively form the glycocalyx. The glycocalyx is a fuzzy-appearing coating around the cell formed from glycoproteins and other carbohydrates attached to the cell membrane. The glycocalyx can have various roles. For example, it may have molecules that allow the cell to bind to another cell, it may contain receptors for hormones, or it might have enzymes to break down nutrients. The glycocalyces found in a person’s body are products of that person’s genetic makeup. They give each of the individual’s trillions of cells the “identity” of belonging in the person’s body. This identity is the primary way that a person’s immune defense cells “know” not to attack the person’s own body cells, but it also is the reason organs donated by another person might be rejected.

Peripheral proteins are typically found on the inner or outer surface of the lipid bilayer but can also be attached to the internal or external surface of an integral protein. These proteins typically perform a specific function for the cell. Some peripheral proteins on the surface of intestinal cells, for example, act as digestive enzymes to break down nutrients to sizes that can pass through the cells and into the bloodstream.

Transport across the Cell Membrane

One of the great wonders of the cell membrane is its ability to regulate the concentration of substances inside the cell. These substances include ions such as Ca++, Na+, K+, and Cl–; nutrients including sugars, fatty acids, and amino acids; and waste products, particularly carbon dioxide (CO2), which must leave the cell.

The membrane’s lipid bilayer structure provides the first level of control. The phospholipids are tightly packed together, and the membrane has a hydrophobic interior. This structure causes the membrane to be selectively permeable. A membrane that has selective permeability allows only substances meeting certain criteria to pass through it unaided. In the case of the cell membrane, only relatively small, nonpolar materials can move through the lipid bilayer (remember, the lipid tails of the membrane are nonpolar). Some examples of these are other lipids, oxygen and carbon dioxide gases, and alcohol. However, water-soluble materials—like glucose, amino acids, and electrolytes—need some assistance to cross the membrane because they are repelled by the hydrophobic tails of the phospholipid bilayer. All substances that move through the membrane do so by one of two general methods, which are categorized based on whether or not energy is required. Passive transport is the movement of substances across the membrane without the expenditure of cellular energy. In contrast, active transport is the movement of substances across the membrane using energy from adenosine triphosphate (ATP).

Passive Transport

In order to understand how substances move passively across a cell membrane, it is necessary to understand concentration gradients and diffusion. A concentration gradient is the difference in concentration of a substance across a space. Molecules (or ions) will spread/diffuse from where they are more concentrated to where they are less concentrated until they are equally distributed in that space. (When molecules move in this way, they are said to move down their concentration gradient.) Diffusion is the movement of particles from an area of higher concentration to an area of lower concentration. A couple of common examples will help to illustrate this concept. Imagine being inside a closed bathroom. If a bottle of perfume were sprayed, the scent molecules would naturally diffuse from the spot where they left the bottle to all corners of the bathroom, and this diffusion would go on until no more concentration gradient remains. Another example is a spoonful of sugar placed in a cup of tea. Eventually the sugar will diffuse throughout the tea until no concentration gradient remains. In both cases, if the room is warmer or the tea hotter, diffusion occurs even faster as the molecules are bumping into each other and spreading out faster than at cooler temperatures. Having an internal body temperature around 98.6° F thus also aids in diffusion of particles within the body.

Whenever a substance exists in greater concentration on one side of a semipermeable membrane, such as the cell membranes, any substance that can move down its concentration gradient across the membrane will do so. Consider substances that can easily diffuse through the lipid bilayer of the cell membrane, such as the gases oxygen (O2) and CO2. O2 generally diffuses into cells because it is more concentrated outside of them, and CO2 typically diffuses out of cells because it is more concentrated inside of them.  In both these examples the molecules rely on their own kinetic energy to move, so neither of these examples requires any energy output from the cell.  The movement of molecules across a cell membrane without the expenditure of cellular energy is referred to as passive transport, or diffusion.

[caption id="" align="alignright" width="380"]This figure shows the simple diffusion of small non-polar molecules across the plasma membrane. A red horizontal arrow pointing towards the right indicates the progress of time. The nonpolar molecules are shown in blue and are present in higher numbers in the extracellular fluid. There are a few nonpolar molecules in the cytoplasm and their number increases with time. Figure 4. Simple Diffusion across the Cell (Plasma) Membrane. The structure of the lipid bilayer allows small, uncharged substances such as oxygen and carbon dioxide, and hydrophobic molecules such as lipids, to pass through the cell membrane, down their concentration gradient, by simple diffusion.[/caption] Before moving on, you need to review the gases that can diffuse across a cell membrane. Because cells rapidly use up oxygen during metabolism, there is typically a lower concentration of O2 inside the cell than outside. As a result, oxygen will diffuse from the interstitial fluid into the cytoplasm within the cell. On the other hand, because cells produce CO2 as a byproduct of metabolism, CO2 concentrations rise within the cytoplasm; therefore, CO2 will move from the cell into the interstitial fluid, where its concentration is lower. Both these molecules are small and nonpolar, which means they can easily interact with the hydrophobic core of a lipid bilayer and move between the molecules to get from one side to the other.  This mechanism of small, nonpolar molecules slipping between the lipid tails of a cell membrane from the side where they are more concentrated to the side where they are less concentrated is a form of passive transport called simple diffusion (Figure 4). [caption id="" align="alignright" width="300"]This diagram shows the different means of facilitated diffusion across the plasma membrane. In the top panel, a channel protein is shown to allow the transport of solutes across the membrane. In the bottom panel, the membrane contains carrier proteins in addition to channel proteins. Figure 5. Facilitated Diffusion. (a) Facilitated diffusion of substances crossing the cell (plasma) membrane takes place with the help of proteins such as channel proteins and carrier proteins. Channel proteins are less selective than carrier proteins, and usually mildly discriminate between their cargo based on size and charge. (b) Carrier proteins are more selective, often only allowing one particular type of molecule to cross.[/caption] Large polar or ionic molecules, which are hydrophilic, cannot easily cross the phospholipid bilayer.  Charged atoms or molecules of any size cannot cross the cell membrane via simple diffusion as the charges are repelled by the hydrophobic tails in the interior of the phospholipid bilayer. Solutes dissolved in water on either side of the cell membrane will tend to diffuse down their concentration gradients, but because most substances cannot pass freely through the lipid bilayer of the cell membrane, their movement is restricted to protein channels and specialized transport mechanisms in the membrane. Facilitated diffusion is the diffusion process used for those substances that cannot cross the lipid bilayer due to their size, charge, and/or polarity (Figure 5). A common example of facilitated diffusion is the movement of glucose into the cell, where it is used to make ATP. Although glucose can be more concentrated outside of a cell, it cannot cross the lipid bilayer via simple diffusion because it is both large and polar. To resolve this, a specialized carrier protein called the glucose transporter will transfer glucose molecules into the cell to facilitate its inward diffusion.

As an example, even though sodium ions (Na+) are highly concentrated outside of cells, these electrolytes are charged and cannot pass through the nonpolar lipid bilayer of the membrane. Their diffusion is facilitated by membrane proteins that form sodium channels (or “pores”), so that Na+ ions can move down their concentration gradient from outside the cells to inside the cells. There are many other solutes that must undergo facilitated diffusion to move into a cell, such as amino acids, or to move out of a cell, such as wastes. Because facilitated diffusion is a passive process, it does not require energy expenditure by the cell.

[caption id="" align="alignleft" width="280"]This figure shows the diffusion of water through osmosis. The left panel shows a beaker with water and different solute concentrations. A semipermeable membrane is present in the middle of the beaker. In the right panel, the water concentration is higher to the right of the semipermeable membrane. Figure 6. Osmosis. Osmosis is the diffusion of water through a semipermeable membrane down its concentration gradient. If a membrane is permeable to water, though not to a solute, water will equalize its own concentration by diffusing to the side of lower water concentration (and thus the side of higher solute concentration). In the beaker on the left, the solution on the right side of the membrane is hypertonic relative to the solution on the left side of the membrane.[/caption] Very small polar molecules, including water, can cross a phospholipid bilayer via simple diffusion due to their small size.  The rate at which water can move across cell membranes is increased by the presence of membrane proteins called aquaporins that form channels through which water molecules (but not solutes) can pass.  Osmosis refers to the passive movement of water across a semipermeable membrane (Figure 6). Osmosis across a cell membrane therefore includes the movement of water molecules by either simple diffusion or facilitated diffusion or both.

The movement of water across a cell membrane cannot be directly determined by cells, so it is important that cells are exposed to an environment in which the concentration of solutes outside of the cells (in the extracellular fluid) is equal to the concentration of solutes inside the cells (in the cytoplasm). Two solutions that have the same concentration of solutes are said to be isotonic (equal tension). When cells and their extracellular environments are isotonic, the concentration of water molecules is the same outside and inside the cells, and the cells maintain their normal shape (and function).

[caption id="" align="alignright" width="280"]This image shows how a red blood cell responds to the tonicity of solution. The left panel shows the hypertonic case, the middle panel shows the isotonic case and the right panel shows the hypotonic case. Figure 7. Concentration of Solutions. A hypertonic solution has a solute concentration higher than another solution. An isotonic solution has a solute concentration equal to another solution. A hypotonic solution has a solute concentration lower than another solution.[/caption] Osmosis occurs when there is an imbalance of solutes outside of a cell versus inside the cell. A solution that has a higher concentration of solutes than another solution is said to be hypertonic, and water molecules tend to diffuse into a hypertonic solution (Figure 7). Cells in a hypertonic solution will shrivel as water leaves the cell via osmosis. In contrast, a solution that has a lower concentration of solutes than another solution is said to be hypotonic, and water molecules tend to diffuse out of a hypotonic solution. Cells in a hypotonic solution will take on too much water and swell, with the risk of eventually bursting. A critical aspect of homeostasis in living things is to create an internal environment in which all of the body’s cells are in an isotonic solution. Various organ systems, particularly the kidneys, work to maintain this homeostasis.

Another mechanism besides diffusion to passively transport materials between compartments is filtration. Unlike diffusion of a substance from where it is more concentrated to less concentrated, filtration uses a hydrostatic pressure gradient that pushes the fluid—and the solutes within it—from a higher pressure area to a lower pressure area. Filtration is an extremely important process in the body. For example, the circulatory system uses filtration to move plasma and substances across the endothelial lining of capillaries and into surrounding tissues, supplying cells with the nutrients. Filtration pressure in the kidneys provides the mechanism to remove wastes from the bloodstream.

Active Transport

For all of the transport methods described above, the cell expends no energy. Membrane proteins that aid in the passive transport of substances do so without the use of ATP. During active transport, ATP is required to move a substance across a membrane, often with the help of carrier proteins, and usually against the concentration gradient of the substance being moved.

One of the most common types of active transport involves proteins that serve as pumps. The word “pump” probably conjures up thoughts of using energy to pump up the tire of a bicycle or a basketball. Similarly, energy from ATP is required for these membrane proteins to transport substances—molecules or ions—across the membrane, usually against their concentration gradients (from an area of low concentration to an area of high concentration).

The sodium-potassium pump, which is also called Na+/K+ ATPase, transports sodium out of a cell while moving potassium into the cell. The Na+/K+ pump is an important ion pump found in the membranes of many types of cells. These pumps are particularly abundant in nerve cells, which are constantly pumping out sodium ions and pulling in potassium ions to maintain an electrical gradient across their cell membranes. An electrical gradient is a difference in electrical charge across a space. In the case of nerve cells, for example, the electrical gradient exists between the inside and outside of the cell, with the inside being negatively-charged (at around -70 mV) relative to the outside. The negative electrical gradient is maintained because each Na+/K+ pump moves three Na+ ions out of the cell and two K+ ions into the cell for each ATP molecule that is used (Figure 8). This process is so important for nerve cells that it accounts for the majority of their ATP usage.

[caption id="" align="alignleft" width="550"]This diagram shows many sodium potassium pumps embedded in the membrane. Potassium is pumped into the cytoplasm and sodium is pumped out of the cytoplasm. Figure 8. Sodium-Potassium Pump. The sodium-potassium pump is found in many cell (plasma) membranes. Powered by ATP, the pump moves sodium and potassium ions in opposite directions, each against its concentration gradient. In a single cycle of the pump, three sodium ions are extruded from and two potassium ions are imported into the cell.[/caption]

Active transport pumps can also work together with other active or passive transport systems to move substances across the membrane. For example, the sodium-potassium pump maintains a high concentration of sodium ions outside of the cell. Therefore, if the cell needs sodium ions, all it has to do is open a passive sodium channel, as the concentration gradient of the sodium ions will drive them to diffuse into the cell. In this way, the action of an active transport pump (the sodium-potassium pump) powers the passive transport of sodium ions by creating a concentration gradient. When active transport powers the transport of another substance in this way, it is called secondary active transport.

Symporters are secondary active transporters that move two substances in the same direction. For example, the sodium-glucose symporter uses sodium ions to “pull” glucose molecules into the cell. Because cells store glucose for energy, glucose is typically at a higher concentration inside of the cell than outside. However, due to the action of the sodium-potassium pump, sodium ions will easily diffuse into the cell when the symporter is opened. The flood of sodium ions through the symporter provides the energy that allows glucose to move through the symporter and into the cell, against its concentration gradient.

Conversely, antiporters are secondary active transport systems that transport substances in opposite directions. For example, the sodium-hydrogen ion antiporter uses the energy from the inward flood of sodium ions to move hydrogen ions (H+) out of the cell. The sodium-hydrogen antiporter is used to maintain the pH of the cell's interior.

[caption id="" align="alignleft" width="380"]This image shows the three different types of endocytosis. The left panel shows phagocytosis, where a large particle is seen to be engulfed by the membrane into a vacuole. In the middle panel, pinocytosis is shown, where a small particle is engulfed into a vesicle. In the right panel, receptor-mediated endocytosis is shown; the ligand binds to the receptor and is then engulfed into a coated vesicle. Figure 9. Three Forms of Endocytosis. Endocytosis is a form of active transport in which a cell envelopes extracellular materials using its cell membrane. (a) In phagocytosis, which is relatively nonselective, the cell takes in a large particle. (b) In pinocytosis, the cell takes in small particles in fluid. (c) In contrast, receptor-mediated endocytosis is quite selective. When external receptors bind a specific ligand, the cell responds by endocytosing the ligand.[/caption] Other forms of active transport do not involve membrane carriers. Endocytosis (bringing “into the cell”) is the process of a cell ingesting material by enveloping it in a portion of its cell membrane, and then pinching off that portion of membrane (Figure 9). Once pinched off, the portion of membrane and its contents becomes an independent, intracellular vesicle. A vesicle is a membranous sac—a spherical and hollow organelle bounded by a lipid bilayer membrane. Endocytosis often brings materials into the cell that must to be broken down or digested. Phagocytosis (“cell eating”) is the endocytosis of large particles. Many immune cells engage in phagocytosis of invading pathogens. Like little Pac-men, their job is to patrol body tissues for unwanted matter, such as invading bacterial cells, phagocytize them, and digest them. In contrast to phagocytosis, pinocytosis (“cell drinking”) brings fluid containing dissolved substances into a cell through membrane vesicles.

Phagocytosis and pinocytosis take in large portions of extracellular material, and they are typically not highly selective in the substances they bring in. Cells regulate the endocytosis of specific substances via receptor-mediated endocytosis. Receptor-mediated endocytosis is endocytosis by a portion of the cell membrane that contains many receptors that are specific for a certain substance. Once the surface receptors have bound sufficient amounts of the specific substance (the receptor’s ligand), the cell will endocytose the part of the cell membrane containing the receptor-ligand complexes. Iron, a required component of hemoglobin, is endocytosed by red blood cells in this way. Iron is bound to a protein called transferrin in the blood. Specific transferrin receptors on red blood cell surfaces bind the iron-transferrin molecules, and the cell endocytoses the receptor-ligand complexes.

In contrast with endocytosis, exocytosis (taking “out of the cell”) is the process of a cell exporting material using vesicular transport (Figure 10). Many cells manufacture substances that must be secreted, like a factory manufacturing a product for export. These substances are typically packaged into membrane-bound vesicles within the cell. When the vesicle membrane fuses with the cell membrane, the vesicle releases it contents into the interstitial fluid. The vesicle membrane then becomes part of the cell membrane. Cells of the stomach and pancreas produce and secrete digestive enzymes through exocytosis (Figure 11). Endocrine cells produce and secrete hormones that are sent throughout the body, and certain immune cells produce and secrete large amounts of histamine, a chemical important for immune responses.

[caption id="attachment_3050" align="aligncenter" width="150"] Watch this amoeba sisters video to learn more about cell transport![/caption]
[caption id="attachment_2954" align="aligncenter" width="150"] Watch this CrashCourse video on membranes and transport![/caption] [caption id="attachment_2955" align="aligncenter" width="150"] Check out the Khan Academy membranes and transport section to find out more[/caption]
[caption id="" align="alignright" width="242"]This figure shows the process of exocytosis. A vesicle is shown fusing with the membrane and then releasing its contents into the extracellular fluid. Figure 10. Exocytosis. Exocytosis is much like endocytosis in reverse. Material destined for export is packaged into a vesicle inside the cell. The membrane of the vesicle fuses with the cell membrane, and the contents are released into the extracellular space.[/caption]
[caption id="" align="aligncenter" width="370"]This micrograph shows the structure of a pancreatic acinar cell and the location of secretory vesicles. Figure 11. Pancreatic Cells' Enzyme Products. The pancreatic acinar cells produce and secrete many enzymes that digest food. The tiny black granules in this electron micrograph are secretory vesicles filled with enzymes that will be exported from the cells via exocytosis. LM × 2900. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]
]]> 1950 1936 2 0 3.5 Cell Growth and Division https://pressbooks.bccampus.ca/apdouglas2017/chapter/3-5-cell-growth-and-division/ Fri, 14 Jul 2017 22:18:34 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1972

Learning Objectives

By the end of this section, you will be able to:
  • Describe the stages of the cell cycle
  • Discuss how the cell cycle is regulated
  • Describe the implications of losing control over the cell cycle
  • Describe the stages of mitosis and cytokinesis, in order

So far in this chapter, you have read numerous times of the importance and prevalence of cell division. While there are a few cells in the body that do not undergo cell division (such as gametes, red blood cells, most neurons, and some muscle cells), most somatic cells divide regularly. A somatic cell is a general term for a body cell, and all human cells, except for the cells that produce eggs and sperm (which are referred to as germ cells), are somatic cells. Somatic cells contain two copies of each of their chromosomes (one copy received from each parent). A homologous pair of chromosomes is the two copies of a single chromosome found in each somatic cell. The human is a diploid organism, having 23 homologous pairs of chromosomes in each of the somatic cells. The condition of having pairs of chromosomes is known as diploidy.

Cells in the body replace themselves over the lifetime of a person. For example, the cells lining the gastrointestinal tract must be frequently replaced when constantly “worn off” by the movement of food through the gut. But what triggers a cell to divide, and how does it prepare for and complete cell division? The cell cycle is the sequence of events in the life of the cell from the moment it is created at the end of a previous cycle of cell division until it then divides itself, generating two new cells.

The Cell Cycle

One “turn” or cycle of the cell cycle consists of two general phases: interphase, followed by mitosis and cytokinesis. Interphase is the period of the cell cycle during which the cell is not dividing. The majority of cells are in interphase most of the time. Mitosis is the division of genetic material, during which the cell nucleus breaks down and two new, fully functional, nuclei are formed. Cytokinesis divides the cytoplasm into two distinctive cells.

Interphase

A cell grows and carries out all normal metabolic functions and processes in a period called G1 (Figure 1). G1 phase (gap 1 phase) is the first gap, or growth phase in the cell cycle. For cells that will divide again, G1 is followed by replication of the DNA, during the S phase. The S phase (synthesis phase) is period during which a cell replicates its DNA.

[caption id="" align="aligncenter" width="280"]This figure shows the different stages of the cell cycle. The G0 phase where the cells are not actively dividing is also labeled. Figure 1. Cell Cycle. The two major phases of the cell cycle include mitosis (cell division), and interphase, when the cell grows and performs all of its normal functions. Interphase is further subdivided into G1, S, and G2 phases.[/caption]

After the synthesis phase, the cell proceeds through the G2 phase. The G2 phase is a second gap phase, during which the cell continues to grow and makes the necessary preparations for mitosis. Between G1, S, and G2 phases, cells will vary the most in their duration of the G1 phase. It is here that a cell might spend a couple of hours, or many days. The S phase typically lasts between 8-10 hours and the G2 phase approximately 5 hours. In contrast to these phases, the G0 phase is a resting phase of the cell cycle. Cells that have temporarily stopped dividing and are resting (a common condition) and cells that have permanently ceased dividing (like nerve cells) are said to be in G0.

The Structure of Chromosomes

Billions of cells in the human body divide every day. During the synthesis phase (S, for DNA synthesis) of interphase, the amount of DNA within the cell precisely doubles. Therefore, after DNA replication but before cell division, each cell actually contains two copies of each chromosome. Each copy of the chromosome is referred to as a sister chromatid and is physically bound to the other copy. The centromere is the structure that attaches one sister chromatid to another. Because a human cell has 46 chromosomes, during this phase, there are 92 chromatids (46 × 2) in the cell. Make sure not to confuse the concept of a pair of chromatids (one chromosome and its exact copy attached during mitosis) and a homologous pair of chromosomes (two paired chromosomes which were inherited separately, one from each parent) (Figure 2).

[caption id="" align="aligncenter" width="250"]This image shows a pair of chromosomes. The major parts such as the homologous chromosomes, kinetochore and the sister chromatids are labeled. Figure 2. A Homologous Pair of Chromosomes with their Attached Sister Chromatids. The red and blue colors correspond to a homologous pair of chromosomes. Each member of the pair was separately inherited from one parent. Each chromosome in the homologous pair is also bound to an identical sister chromatid, which is produced by DNA replication, and results in the familiar “X” shape.[/caption]

Mitosis and Cytokinesis

The mitotic phase of the cell typically takes between 1 and 2 hours. During this phase, a cell undergoes two major processes. First, it completes mitosis, during which the contents of the nucleus are equitably pulled apart and distributed between its two halves. Cytokinesis then occurs, dividing the cytoplasm and cell body into two new cells. Mitosis is divided into four major stages that take place after interphase (Figure 3) and in the following order: prophase, metaphase, anaphase, and telophase. The process is then followed by cytokinesis.

[caption id="attachment_1512" align="aligncenter" width="600"]This tabular image shows the different stages of mitosis and cytokinesis using both drawings and text. The top panel is a series of schematics for each step, followed by text listing the important aspects of that step. The bottom panel shows fluorescent micrographs for the corresponding stage. Figure 3. Cell Division: Mitosis Followed by Cytokinesis. The stages of cell division oversee the separation of identical genetic material into two new nuclei, followed by the division of the cytoplasm.[/caption]
Prophase is the first phase of mitosis, during which the loosely packed chromatin coils and condenses into visible chromosomes. During prophase, each chromosome becomes visible with its identical partner attached, forming the familiar X-shape of sister chromatids. The nucleolus disappears early during this phase, and the nuclear envelope also disintegrates.A major occurrence during prophase concerns a very important structure that contains the origin site for microtubule growth. Recall the cellular structures called centrioles that serve as origin points from which microtubules extend. These tiny structures also play a very important role during mitosis. A centrosome is a pair of centrioles together. The cell contains two centrosomes side-by-side, which begin to move apart during prophase. As the centrosomes migrate to two different sides of the cell, microtubules begin to extend from each like long fingers from two hands extending toward each other. The mitotic spindle is the structure composed of the centrosomes and their emerging microtubules.

Near the end of prophase there is an invasion of the nuclear area by microtubules from the mitotic spindle. The nuclear membrane has disintegrated, and the microtubules attach themselves to the centromeres that adjoin pairs of sister chromatids. The kinetochore is a protein structure on the centromere that is the point of attachment between the mitotic spindle and the sister chromatids. This stage is referred to as late prophase or “prometaphase” to indicate the transition between prophase and metaphase.

Metaphase is the second stage of mitosis. During this stage, the sister chromatids, with their attached microtubules, line up along a linear plane in the middle of the cell. A metaphase plate forms between the centrosomes that are now located at either end of the cell. The metaphase plate is the name for the plane through the center of the spindle on which the sister chromatids are positioned. The microtubules are now poised to pull apart the sister chromatids and bring one from each pair to each side of the cell.

Anaphase is the third stage of mitosis. Anaphase takes place over a few minutes, when the pairs of sister chromatids are separated from one another, forming individual chromosomes once again. These chromosomes are pulled to opposite ends of the cell by their kinetochores, as the microtubules shorten. Each end of the cell receives one partner from each pair of sister chromatids, ensuring that the two new daughter cells will contain identical genetic material.

Telophase is the final stage of mitosis. Telophase is characterized by the formation of two new daughter nuclei at either end of the dividing cell. These newly formed nuclei surround the genetic material, which uncoils such that the chromosomes return to loosely packed chromatin. Nucleoli also reappear within the new nuclei, and the mitotic spindle breaks apart, each new cell receiving its own complement of DNA, organelles, membranes, and centrioles. At this point, the cell is already beginning to split in half as cytokinesis begins.

The cleavage furrow is a contractile band made up of microfilaments that forms around the midline of the cell during cytokinesis. (Recall that microfilaments consist of actin.) This contractile band squeezes the two cells apart until they finally separate. Two new cells are now formed. One of these cells (the “stem cell”) enters its own cell cycle; able to grow and divide again at some future time. The other cell transforms into the functional cell of the tissue, typically replacing an “old” cell there.

Imagine a cell that completed mitosis but never underwent cytokinesis. In some cases, a cell may divide its genetic material and grow in size, but fail to undergo cytokinesis. This results in larger cells with more than one nucleus. Usually this is an unwanted aberration and can be a sign of cancerous cells.

Cell Cycle Control

A very elaborate and precise system of regulation controls direct the way cells proceed from one phase to the next in the cell cycle and begin mitosis. The control system involves molecules within the cell as well as external triggers. These internal and external control triggers provide “stop” and “advance” signals for the cell. Precise regulation of the cell cycle is critical for maintaining the health of an organism, and loss of cell cycle control can lead to cancer.

Mechanisms of Cell Cycle Control

As the cell proceeds through its cycle, each phase involves certain processes that must be completed before the cell should advance to the next phase. A checkpoint is a point in the cell cycle at which the cycle can be signaled to move forward or stopped. At each of these checkpoints, different varieties of molecules provide the stop or go signals, depending on certain conditions within the cell. A cyclin is one of the primary classes of cell cycle control molecules (Figure 4). A cyclin-dependent kinase (CDK) is one of a group of molecules that work together with cyclins to determine progression past cell checkpoints. By interacting with many additional molecules, these triggers push the cell cycle forward unless prevented from doing so by “stop” signals, if for some reason the cell is not ready. At the G1 checkpoint, the cell must be ready for DNA synthesis to occur. At the G2 checkpoint the cell must be fully prepared for mitosis. Even during mitosis, a crucial stop and go checkpoint in metaphase ensures that the cell is fully prepared to complete cell division. The metaphase checkpoint ensures that all sister chromatids are properly attached to their respective microtubules and lined up at the metaphase plate before the signal is given to separate them during anaphase.

[caption id="" align="aligncenter" width="350"]This image shows the different stages of the cell cycle along with the checkpoints between them and the cyclins responsible for the checkpoint at each stage. Figure 4. Control of the Cell Cycle. Cells proceed through the cell cycle under the control of a variety of molecules, such as cyclins and cyclin-dependent kinases. These control molecules determine whether or not the cell is prepared to move into the following stage.[/caption]

The Cell Cycle Out of Control: Implications

Most people understand that cancer or tumors are caused by abnormal cells that multiply continuously. If the abnormal cells continue to divide unstopped, they can damage the tissues around them, spread to other parts of the body, and eventually result in death. In healthy cells, the tight regulation mechanisms of the cell cycle prevent this from happening, while failures of cell cycle control can cause unwanted and excessive cell division. Failures of control may be caused by inherited genetic abnormalities that compromise the function of certain “stop” and “go” signals. Environmental insult that damages DNA can also cause dysfunction in those signals. Often, a combination of both genetic predisposition and environmental factors lead to cancer.

The process of a cell escaping its normal control system and becoming cancerous may actually happen throughout the body quite frequently. Fortunately, certain cells of the immune system are capable of recognizing cells that have become cancerous and destroying them. However, in certain cases the cancerous cells remain undetected and continue to proliferate. If the resulting tumor does not pose a threat to surrounding tissues, it is said to be benign and can usually be easily removed. If capable of damage, the tumor is considered malignant and the patient is diagnosed with cancer.
Homeostatic Imbalances

Cancer Arises from Homeostatic Imbalances Cancer is an extremely complex condition, capable of arising from a wide variety of genetic and environmental causes. Typically, mutations or aberrations in a cell’s DNA that compromise normal cell cycle control systems lead to cancerous tumors. Cell cycle control is an example of a homeostatic mechanism that maintains proper cell function and health. While progressing through the phases of the cell cycle, a large variety of intracellular molecules provide stop and go signals to regulate movement forward to the next phase. These signals are maintained in an intricate balance so that the cell only proceeds to the next phase when it is ready. This homeostatic control of the cell cycle can be thought of like a car’s cruise control. Cruise control will continually apply just the right amount of acceleration to maintain a desired speed, unless the driver hits the brakes, in which case the car will slow down. Similarly, the cell includes molecular messengers, such as cyclins, that push the cell forward in its cycle.

In addition to cyclins, a class of proteins that are encoded by genes called proto-oncogenes provide important signals that regulate the cell cycle and move it forward. Examples of proto-oncogene products include cell-surface receptors for growth factors, or cell-signaling molecules, two classes of molecules that can promote DNA replication and cell division. In contrast, a second class of genes known as tumor suppressor genes sends stop signals during a cell cycle. For example, certain protein products of tumor suppressor genes signal potential problems with the DNA and thus stop the cell from dividing, while other proteins signal the cell to die if it is damaged beyond repair. Some tumor suppressor proteins also signal a sufficient surrounding cellular density, which indicates that the cell need not presently divide. The latter function is uniquely important in preventing tumor growth: normal cells exhibit a phenomenon called “contact inhibition;” thus, extensive cellular contact with neighboring cells causes a signal that stops further cell division.

These two contrasting classes of genes, proto-oncogenes and tumor suppressor genes, are like the accelerator and brake pedal of the cell’s own “cruise control system,” respectively. Under normal conditions, these stop and go signals are maintained in a homeostatic balance. Generally speaking, there are two ways that the cell’s cruise control can lose control: a malfunctioning (overactive) accelerator, or a malfunctioning (underactive) brake. When compromised through a mutation, or otherwise altered, proto-oncogenes can be converted to oncogenes, which produce oncoproteins that push a cell forward in its cycle and stimulate cell division even when it is undesirable to do so. For example, a cell that should be programmed to self-destruct (a process called apoptosis) due to extensive DNA damage might instead be triggered to proliferate by an oncoprotein. On the other hand, a dysfunctional tumor suppressor gene may fail to provide the cell with a necessary stop signal, also resulting in unwanted cell division and proliferation.

A delicate homeostatic balance between the many proto-oncogenes and tumor suppressor genes delicately controls the cell cycle and ensures that only healthy cells replicate. Therefore, a disruption of this homeostatic balance can cause aberrant cell division and cancerous growths.

[caption id="attachment_3039" align="aligncenter" width="150"] Watch this amoeba sisters video to learn more about mitosis![/caption] [caption id="attachment_3040" align="aligncenter" width="150"] Watch this amoeba sisters video to learn about the process of meiosis![/caption]
[caption id="attachment_2957" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about mitosis![/caption] The life of cell consists of stages that make up the cell cycle. After a cell is born, it passes through an interphase before it is ready to replicate itself and produce daughter cells. This interphase includes two gap phases (G1 and G2), as well as an S phase, during which its DNA is replicated in preparation for cell division. The cell cycle is under precise regulation by chemical messengers both inside and outside the cell that provide “stop” and “go” signals for movement from one phase to the next. Failures of these signals can result in cells that continue to divide uncontrollably, which can lead to cancer.

Once a cell has completed interphase and is ready for cell division, it proceeds through four separate stages of mitosis (prophase, metaphase, anaphase, and telophase). Telophase is followed by the division of the cytoplasm (cytokinesis), which generates two daughter cells. This process takes place in all normally dividing cells of the body except for the germ cells that produce eggs and sperm.

]]> 1972 1936 6 0 4.2 Epithelial Tissue https://pressbooks.bccampus.ca/apdouglas2017/chapter/4-2-epithelial-tissue/ Fri, 14 Jul 2017 22:21:15 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1992

Learning Objectives

By the end of this section, you will be able to:
  • Explain the structure and function of epithelial tissue
  • Distinguish between tight junctions, anchoring junctions, and gap junctions
  • Distinguish between simple epithelia and stratified epithelia, as well as between squamous, cuboidal, and columnar epithelia
  • Describe the structure and function of endocrine and exocrine glands and their respective secretions

Most epithelial tissues are essentially large sheets of cells covering all the surfaces of the body exposed to the outside world and lining the outside of organs. Epithelium also forms much of the glandular tissue of the body. Skin is not the only area of the body exposed to the outside. Other areas include the airways, the digestive tract, as well as the urinary and reproductive systems, all of which are lined by an epithelium. Hollow organs and body cavities that do not connect to the exterior of the body, which includes, blood vessels and serous membranes, are lined by endothelium (plural = endothelia), which is a type of epithelium.

Epithelial cells derive from all three major embryonic layers. The epithelia lining the skin, parts of the mouth and nose, and the anus develop from the ectoderm. Cells lining the airways and most of the digestive system originate in the endoderm. The epithelium that lines vessels in the lymphatic and cardiovascular system derives from the mesoderm and is called an endothelium.

All epithelia share some important structural and functional features. This tissue is highly cellular, with little or no extracellular material present between cells. Adjoining cells form a specialized intercellular connection between their cell membranes called a cell junction. The epithelial cells exhibit polarity with differences in structure and function between the exposed or apical facing surface of the cell and the basal surface close to the underlying body structures. The basal lamina, a mixture of glycoproteins and collagen, provides an attachment site for the epithelium, separating it from underlying connective tissue. The basal lamina attaches to a reticular lamina, which is secreted by the underlying connective tissue, forming a basement membrane that helps hold it all together.

Epithelial tissues are nearly completely avascular. For instance, no blood vessels cross the basement membrane to enter the tissue, and nutrients must come by diffusion or absorption from underlying tissues or the surface. Many epithelial tissues are capable of rapidly replacing damaged and dead cells. Sloughing off of damaged or dead cells is a characteristic of surface epithelium and allows our airways and digestive tracts to rapidly replace damaged cells with new cells.

Generalized Functions of Epithelial Tissue

Epithelial tissues provide the body’s first line of protection from physical, chemical, and biological wear and tear. The cells of an epithelium act as gatekeepers of the body controlling permeability and allowing selective transfer of materials across a physical barrier. All substances that enter the body must cross an epithelium. Some epithelia often include structural features that allow the selective transport of molecules and ions across their cell membranes.

Many epithelial cells are capable of secretion and release mucous and specific chemical compounds onto their apical surfaces. The epithelium of the small intestine releases digestive enzymes, for example. Cells lining the respiratory tract secrete mucous that traps incoming microorganisms and particles. A glandular epithelium contains many secretory cells.

The Epithelial Cell

Epithelial cells are typically characterized by the polarized distribution of organelles and membrane-bound proteins between their basal and apical surfaces. Particular structures found in some epithelial cells are an adaptation to specific functions. Certain organelles are segregated to the basal sides, whereas other organelles and extensions, such as cilia, when present, are on the apical surface.

Cilia are microscopic extensions of the apical cell membrane that are supported by microtubules. They beat in unison and move fluids as well as trapped particles. Ciliated epithelium lines the ventricles of the brain where it helps circulate the cerebrospinal fluid. The ciliated epithelium of your airway forms a mucociliary escalator that sweeps particles of dust and pathogens trapped in the secreted mucous toward the throat. It is called an escalator because it continuously pushes mucous with trapped particles upward. In contrast, nasal cilia sweep the mucous blanket down towards your throat. In both cases, the transported materials are usually swallowed, and end up in the acidic environment of your stomach.

Cell to Cell Junctions

[caption id="" align="alignright" width="500"]These three illustrations each show the edges of two vertical cell membranes. The cell membranes are viewed partially from the side so that the inside edge of the right cell membrane is visible. The upper left image shows a tight junction. The two cell membranes are bound by transmembrane protein strands. The proteins travel the inside edge of the right cell membrane and cross over to the left cell membrane, cinching the two membranes together. The cell membranes are still somewhat separated in between neighboring strands, creating intercellular spaces. The upper right diagram shows a gap junction. The gap junctions are composed of two interlocking connexins, which are round, hollow tubes that extend through the cell membranes. Two connexins, one from the left cell membrane and the other from the right cell membrane, meet between the two cells, forming a connexon. Even at the site of the connexon, there is a small gap between the cell membranes. On the inside edge of the right cell membrane, the gap junction appears as a depression. Three connexins are embedded into the membranes like buttons on a shirt. The bottom images show the three types of anchoring junctions. The left image shows a desmosome. Here, the inside edge of both the right and left cell membranes have brown, round plaques. Each plaque has tentacle-like intermediate filaments (keratin) that extend into each cell’s cytoplasm. The two plaques are connected across the intercellular space by several interlocking transmembrane glycoproteins (cadherin). The connected glycoproteins look similar to a zipped-up zipper between the right and left cell membranes. The right image shows an adheren. These are similar to desmosomes, with two plaques on the inside edge of each cell membrane connected across the intercellular space by glycoproteins. However, the plaques do not contain the tentacle-like intermediate filaments branching into the cytoplasm. Instead, the plaques are ribbed with green actin filaments. The filaments are neatly arranged in parallel, horizontal strands on the surface of the plaque facing the cytoplasm. The bottom image shows a hemidesmosome. Rather than located between two neighboring cells, the hemidesmosome is located between the bottom of a cell and the basement membrane. A hemidesmosome contains a single plaque on the inside edge of the cell membrane. Like the desmosome, intermediate filaments project from the plaque into the cytoplasm. The opposite side of the plaque has purple, knob-shaped integrins extending out to the basal lamina of the basement membrane. Figure 1. Types of Cell Junctions. The three basic types of cell-to-cell junctions are tight junctions, gap junctions, and anchoring junctions.[/caption]

Cells of epithelia are closely connected and are not separated by intracellular material. Three basic types of connections allow varying degrees of interaction between the cells: tight junctions, anchoring junctions, and gap junctions (Figure 1).

At one end of the spectrum is the tight junction, which separates the cells into apical and basal compartments. An anchoring junction includes several types of cell junctions that help stabilize epithelial tissues. Anchoring junctions are common on the lateral and basal surfaces of cells where they provide strong and flexible connections. There are three types of anchoring junctions: desmosomes, hemidesmosomes, and adherens. Desmosomes occur in patches on the membranes of cells. The patches are structural proteins on the inner surface of the cell’s membrane. The adhesion molecule, cadherin, is embedded in these patches and projects through the cell membrane to link with the cadherin molecules of adjacent cells. These connections are especially important in holding cells together. Hemidesmosomes, which look like half a desmosome, link cells to the extracellular matrix, for example, the basal lamina. While similar in appearance to desmosomes, they include the adhesion proteins called integrins rather than cadherins. Adherens junctions use either cadherins or integrins depending on whether they are linking to other cells or matrix. The junctions are characterized by the presence of the contractile protein actin located on the cytoplasmic surface of the cell membrane. The actin can connect isolated patches or form a belt-like structure inside the cell. These junctions influence the shape and folding of the epithelial tissue.

In contrast with the tight and anchoring junctions, a gap junction forms an intercellular passageway between the membranes of adjacent cells to facilitate the movement of small molecules and ions between the cytoplasm of adjacent cells. These junctions allow electrical and metabolic coupling of adjacent cells, which coordinates function in large groups of cells.

Classification of Epithelial Tissues

[caption id="" align="alignright" width="520"]This figure is a table showing the appearance of squamous, cuboidal and columnar epithelial tissues. Simple and compound forms are shown for each tissue type. In a simple squamous epithelium, the cells are flattened and single layered. In a simple cuboidal epithelium, the cells are cube shaped and single layered. In a simple columnar epithelium, the cells are rectangular and are attached to the basement membrane on one of their narrow sides, so that each cell is standing up like a column. There is only one layer of cells. In a pseudostratified columnar epithelium, the cells are column-like in appearance, but they vary in height. The taller cells bend over the tops of the shorter cells so that the top of the epithelial tissue is continuous. There is only one layer of cells. A stratified squamous epithelium contains many layers of flattened cells. Stratified cuboidal epithelium contains many layers of cube-shaped cells. Stratified columnar epithelium contains many layers of rectangular, column-shaped cells. Figure 2. Cells of Epithelial Tissue. Simple epithelial tissue is organized as a single layer of cells and stratified epithelial tissue is formed by several layers of cells.[/caption]

Epithelial tissues are classified according to the shape of the cells and number of the cell layers formed (Figure 2). Cell shapes can be squamous (flattened and thin), cuboidal (boxy, as wide as it is tall), or columnar (rectangular, taller than it is wide). Similarly, the number of cell layers in the tissue can be one—where every cell rests on the basal lamina—which is a simple epithelium, or more than one, which is a stratified epithelium and only the basal layer of cells rests on the basal lamina. Pseudostratified (pseudo- = “false”) describes tissue with a single layer of irregularly shaped cells that give the appearance of more than one layer. Transitional describes a form of specialized stratified epithelium in which the shape of the cells can vary.

Simple Epithelium

The shape of the cells in the single cell layer of simple epithelium reflects the functioning of those cells. The cells in simple squamous epithelium have the appearance of thin scales. Squamous cell nuclei tend to be flat, horizontal, and elliptical, mirroring the form of the cell. The endothelium is the epithelial tissue that lines vessels of the lymphatic and cardiovascular system, and it is made up of a single layer of squamous cells. Simple squamous epithelium, because of the thinness of the cell, is present where rapid passage of chemical compounds is observed. The alveoli of lungs where gases diffuse, segments of kidney tubules, and the lining of capillaries are also made of simple squamous epithelial tissue. The mesothelium is a simple squamous epithelium that forms the surface layer of the serous membrane that lines body cavities and internal organs. Its primary function is to provide a smooth and protective surface. Mesothelial cells are squamous epithelial cells that secrete a fluid that lubricates the mesothelium.

In simple cuboidal epithelium, the nucleus of the box-like cells appears round and is generally located near the center of the cell. These epithelia are active in the secretion and absorptions of molecules. Simple cuboidal epithelia are observed in the lining of the kidney tubules and in the ducts of glands.

[caption id="" align="alignleft" width="250"]This illustration shows a diagram of a goblet cell. The goblet cell is shaped roughly like an upside down vase. The enlarged end at the top contains six finger like projections labeled microvilli. Between the microvilli, secretary vesicles containing mucin are moving from the upper half of the cell toward the microvilli. Below the secretory vesicles are several rough endoplasmic reticula and an irregularly shaped Golgi apparatus with secretory vesicles budding off of it. The narrow, lower half of the cell contains the oval-shaped nucleus as well as a few mitochondria and segments of the endoplasmic reticulum. Figure 3. Goblet Cell. (a) In the lining of the small intestine, columnar epithelium cells are interspersed with goblet cells. (b) The arrows in this micrograph point to the mucous-secreting goblet cells. LM × 1600. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

The second image is a micrograph of the innermost lining of the small intestine. This innermost lining is a simple columnar epithelium, with a single layer of rectangular cells oriented in a line. Occasionally, the line of epithelial cells is interrupted by a goblet cell. Goblet cells are thinner than the epithelial cells and appear roughly pill shaped. In this micrograph, the cells did not stain as darkly as the epithelial cells.In simple columnar epithelium, the nucleus of the tall column-like cells tends to be elongated and located in the basal end of the cells. Like the cuboidal epithelia, this epithelium is active in the absorption and secretion of molecules. Simple columnar epithelium forms the lining of some sections of the digestive system and parts of the female reproductive tract. Ciliated columnar epithelium is composed of simple columnar epithelial cells with cilia on their apical surfaces. These epithelial cells are found in the lining of the fallopian tubes and parts of the respiratory system, where the beating of the cilia helps remove particulate matter.

Pseudostratified columnar epithelium is a type of epithelium that appears to be stratified but instead consists of a single layer of irregularly shaped and differently sized columnar cells. In pseudostratified epithelium, nuclei of neighboring cells appear at different levels rather than clustered in the basal end. The arrangement gives the appearance of stratification; but in fact all the cells are in contact with the basal lamina, although some do not reach the apical surface. Pseudostratified columnar epithelium is found in the respiratory tract, where some of these cells have cilia.

Both simple and pseudostratified columnar epithelia are heterogeneous epithelia because they include additional types of cells interspersed among the epithelial cells. For example, a goblet cell is a mucous-secreting unicellular “gland” interspersed between the columnar epithelial cells of mucous membranes (Figure 3).

Stratified Epithelium

[caption id="" align="alignright" width="500"]This figure is a table with three columns and eight rows. The leftmost column is titled cells, and contains a drawing in each row showing how epithelial cells are arranged above a basement membrane. The middle column is titled location, while the rightmost column is titled function. In a simple squamous epithelium, the cells are flattened and single-layered. Simple squamous cells are found in the air sacs of the lungs, in the lining of the heart, blood vessels and lymphatic vessels. Their function is to allow materials to pass through by diffusion and filtration, as well as to secrete lubricating substances. In a simple cuboidal epithelium, the cells are cube shaped and single layered and located in ducts and secretory portions of small glands as well as in the kidney tubules. The function of simple cuboidal epithelium is to secrete and absorb. In a simple columnar epithelium, the cells are rectangular and are attached to the basement membrane on one of their narrow sides, so that each cell is standing up like a column. There is only one layer of cells. Simple columnar epithelium is found in ciliated tissues including the bronchi, uterine tubes, and uterus, as well as in smooth, nonciliated tissues such as the digestive tract bladder. The function of simple columnar epithelium is to absorb substances but also to secrete mucous and enzymes. In a pseudostratified columnar epithelium, the cells are column-like in appearance, but they vary in height. The taller cells bend over the tops of the shorter cells so that the top of the epithelial tissue is continuous. There is only one layer of cells. Pseudostratified columnar epithelium lines the trachea and much of the upper respiratory tract. The function of pseudostratified columnar epithelium is to secrete mucous and also move that mucus using the hair like cilia projecting from the top of each cell. A stratified squamous epithelium contains many layers of flattened cells. Stratified squamous epithelium lines the esophagus, mouth, and vagina. The function of stratified squamous epithelium is to protect against abrasion. Stratified cuboidal epithelium contains many layers of cube-shaped cells. Stratified cuboidal epithelium is found in the sweat glands, salivary glands, and mammary glands. The function of stratified cuboidal epithelium is to protect other tissues of the body. Stratified columnar epithelium contains many layers of rectangular, column-shaped cells. Stratified columnar epithelium is located in the male urethra and the ducts of some glands. The function of stratified columnar epithelium is to secrete and protect. Transitional epithelium consists of many layers of irregularly shaped cells with diverse sizes. Transitional epithelium is found lining the bladder, urethra and ureters. The function of transitional epithelium is to allow the urinary organs to expand and stretch. Figure 4. Summary of Epithelial Tissue Cells.[/caption]

A stratified epithelium consists of several stacked layers of cells. This epithelium protects against physical and chemical wear and tear. The stratified epithelium is named by the shape of the most apical layer of cells, closest to the free space. Stratified squamous epithelium is the most common type of stratified epithelium in the human body. The apical cells are squamous, whereas the basal layer contains either columnar or cuboidal cells. The top layer may be covered with dead cells filled with keratin. Mammalian skin is an example of this dry, keratinized, stratified squamous epithelium. The lining of the mouth cavity is an example of an unkeratinized, stratified squamous epithelium. Stratified cuboidal epithelium and stratified columnar epithelium can also be found in certain glands and ducts, but are uncommon in the human body.

Another kind of stratified epithelium is transitional epithelium, so-called because of the gradual changes in the shapes of the apical cells as the bladder fills with urine. It is found only in the urinary system, specifically the ureters and urinary bladder. When the bladder is empty, this epithelium is convoluted and has cuboidal apical cells with convex, umbrella shaped, apical surfaces. As the bladder fills with urine, this epithelium loses its convolutions and the apical cells transition from cuboidal to squamous. It appears thicker and more multi-layered when the bladder is empty, and more stretched out and less stratified when the bladder is full and distended. Figure 4 summarizes the different categories of epithelial cell tissue cells. [caption id="attachment_2959" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about epithelial histology.[/caption]

Glandular Epithelium

A gland is a structure made up of one or more cells modified to synthesize and secrete chemical substances. Most glands consist of groups of epithelial cells. A gland can be classified as an endocrine gland, a ductless gland that releases secretions directly into surrounding tissues and fluids (endo- = “inside”), or an exocrine gland whose secretions leave through a duct that opens directly, or indirectly, to the external environment (exo- = “outside”).

Endocrine Glands

The secretions of endocrine glands are called hormones. Hormones are released into the interstitial fluid, diffused into the bloodstream, and delivered to targets, in other words, cells that have receptors to bind the hormones. The endocrine system is part of a major regulatory system coordinating the regulation and integration of body responses. A few examples of endocrine glands include the anterior pituitary, thymus, adrenal cortex, and gonads.

Exocrine Glands

Exocrine glands release their contents through a duct that leads to the epithelial surface. Mucous, sweat, saliva, and breast milk are all examples of secretions from exocrine glands. They are all discharged through tubular ducts. Secretions into the lumen of the gastrointestinal tract, technically outside of the body, are of the exocrine category.

Glandular Structure

[caption id="" align="alignright" width="550"]This table shows the different types of exocrine glands: alveolar (acinar) versus tubular and those with simple ducts versus compound ducts. Each diagram shows a single layer of columnar epithelial cells with a line of cells travelling along the surface of a tissue (surface epithelium) and then dipping into a hole in the tissue. The cells travel down the right side of the hole until they reach the bottom, then curve around the bottom of the hole and then travel up the left side. Finally, the cells emerge back onto the surface of the tissue. The surface epithelial cells are those that are on the surface of the tissue; the duct cells are those that line both walls of the hole. The gland cells are those that line the bottom of the hole. The shape of the hole differs in each gland. In the simple alvelolar (acinar) gland, the duct and gland cells are bulb shaped with the gland cells being the larger end of the bulb. Simple alveolar glands are not found in adults, as these represent an early developmental stage of simple, branched glands. In simple tubular glands, the duct and gland cells are U shaped. Simple tubular glands are found in the intestinal glands. In simple branched alveolar glands, the gland cells form three bulbs at the end of the duct, similar in appearance to a clover leaf. The sebaceous (oil) glands are examples of simple branched alveolar glands. In simple coiled tubular glands, the duct and gland cells form a U, however, the bottom of the U, which is all gland cells, is curved up to the right. Merocrine sweat glands are examples of simple coiled tubular glands. In simple branched tubular glands, the duct is very short and the gland cells divide into three lobes, similar in appearance to a bird’s foot. The gastric glands of the stomach and mucous glands of the esophagus, tongue and duodenum are examples of simple branched tubular glands. Among the glands with compound ducts, compound alveolar (acinar) glands have three sets of clover leaf bulbs, for a total of six bulbs. Two of the clover leaf shaped structures extend parallel to the surface epithelium in opposite directions to each other. The third clover leaf extends down into the tissue, perpendicular to the surface. The duct is cross-shaped. The mammary glands are an example of compound alveolar glands. Compound tubular glands have a similar structure to compound alveolar glands. However, instead of three cloverleaf shaped bulbs, the compound tubular gland has three bird’s foot shaped bulbs. The duct is also cross-shaped in the compound tubular gland. The mucous glands of the mouth and the bulbourethral glands of the male reproductive system are examples of compound tubular glands, which are also found in the seminiferous tubules of the testis. Compound tubuloalveolar glands are a hybrid between the compound alveolar gland and the compound tubular gland. The two sets of bulbs that run parallel to the surface are bird-foot shaped; however, the set of bulbs that runs perpendicularly below the surface is cloverleaf shaped. The salivary glands, glands of the respiratory passages and glands of the pancreas are all compound tubuloalveolar glands. Figure 5. Types of Exocrine Glands. Exocrine glands are classified by their structure.[/caption]

Exocrine glands are classified as either unicellular or multicellular. The unicellular glands are scattered single cells, such as goblet cells, found in the mucous membranes of the small and large intestine.

The multicellular exocrine glands known as serous glands develop from simple epithelium to form a secretory surface that secretes directly into an inner cavity. These glands line the internal cavities of the abdomen and chest and release their secretions directly into the cavities. Other multicellular exocrine glands release their contents through a tubular duct. The duct is single in a simple gland but in compound glands is divided into one or more branches (Figure 5). In tubular glands, the ducts can be straight or coiled, whereas tubes that form pockets are alveolar (acinar), such as the exocrine portion of the pancreas. Combinations of tubes and pockets are known as tubuloalveolar (tubuloacinar) compound glands. In a branched gland, a duct is connected to more than one secretory group of cells.

Methods and Types of Secretion

[caption id="" align="alignleft" width="400"]These three diagrams show the three modes of secretion. All three diagrams show three orange cells in a line with attached to a basement membrane. Each cell has a large nucleus in its lower half. The upper half of each cell contains a Golgi apparatus, which appears like an upside down jellyfish. Yellow secretory vesicles are budding from the top end of the Golgi apparatus. Each vesicle contains several orange circles, which are the secreted substance. In merocrine secretion, the secretory vesicles travel to the top edge of the cells and release the secretion from the cell by melding with the cell membrane. In apocrine secretion, the top third of the cell, which contains the secretory vesicles, pinches in at the sides and then completely disconnects above the Golgi complex. The pinched off portion of the cell is the secretion, as it contains the majority of the secretory vesicles. In holocrine secretion, the upper third of the cell, just above the Golgi complex, forms many finger like projections. Each projection contains several vesicles. The tips of the projections that contain secretory vesicles bud off from the cell. In this method of secretion, the mature cell eventually dies and becomes the secretory product. Figure 6. Modes of Glandular Secretion. (a) In merocrine secretion, the cell remains intact. (b) In apocrine secretion, the apical portion of the cell is released, as well. (c) In holocrine secretion, the cell is destroyed as it releases its product and the cell itself becomes part of the secretion.[/caption]

Exocrine glands can be classified by their mode of secretion and the nature of the substances released, as well as by the structure of the glands and shape of ducts (Figure 6). Merocrine secretion is the most common type of exocrine secretion. The secretions are enclosed in vesicles that move to the apical surface of the cell where the contents are released by exocytosis. For example, watery mucous containing the glycoprotein mucin, a lubricant that offers some pathogen protection is a merocrine secretion. The eccrine glands that produce and secrete sweat are another example.

Apocrine secretion accumulates near the apical portion of the cell. That portion of the cell and its secretory contents pinch off from the cell and are released. The sweat glands of the armpit are classified as apocrine glands. Both merocrine and apocrine glands continue to produce and secrete their contents with little damage caused to the cell because the nucleus and golgi regions remain intact after secretion.

In contrast, the process of holocrine secretion involves the rupture and destruction of the entire gland cell. The cell accumulates its secretory products and releases them only when it bursts. New gland cells differentiate from cells in the surrounding tissue to replace those lost by secretion. The sebaceous glands that produce the oils on the skin and hair are holocrine glands/cells (Figure 7).

[caption id="" align="alignright" width="520"]Image A depicts a cross section of the skin layers. The surface of the skin is at the top of the diagram, with the outer layer occupying about one fifth of the cross section. The outer layer has an irregular border with the inner skin layer, which occupies the remainder of the cross section. A hair follicle is embedded within the inner layer. However, the outer layer actually invaginates into the inner layer around the outside of the follicle, completely sheathing the follicle. The follicle has a bulb at its bottom that is connected to blood vessels. The hair projects from the bulb and travels through the sheath to erupt from the skin surface. The sebaceous gland is an irregular, yellow structure attached at the midpoint of the hair shaft near the border between the inner and outer layers of skin. Its duct actually connects into the side of the hair follicle. Image B shows a micrograph of a sebaceous gland connected to a hair follicle. The bulb of the hair follicle is evident in the micrograph as a bundle of cell surrounding the growing hair at its center. The sebaceous gland is connected to the right of the follicle bulb. The gland appears as an oval shaped mass of pink staining, cube shaped cells with purple nuclei. Figure 7. Sebaceous Glands. These glands secrete oils that lubricate and protect the skin. They are holocrine glands and they are destroyed after releasing their contents. New glandular cells form to replace the cells that are lost. LM × 400. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Glands are also named after the products they produce. The serous gland produces watery, blood-plasma-like secretions rich in enzymes such as alpha amylase, whereas the mucous gland releases watery to viscous products rich in the glycoprotein mucin. Both serous and mucous glands are common in the salivary glands of the mouth. Mixed exocrine glands contain both serous and mucous glands and release both types of secretions.

]]> 1992 1977 3 0 4.4 Muscle Tissue and Motion https://pressbooks.bccampus.ca/apdouglas2017/chapter/4-4-muscle-tissue-and-motion/ Fri, 14 Jul 2017 22:22:14 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2001

Learning Objectives

By the end of this section, you will be able to:
  • Identify the three types of muscle tissue
  • Compare and contrast the functions of each muscle tissue type
  • Explain how muscle tissue can enable motion

Muscle tissue is characterized by properties that allow movement. Muscle cells are excitable; they respond to a stimulus. They are contractile, meaning they can shorten and generate a pulling force. When attached between two movable objects, in other words, bones, contractions of the muscles cause the bones to move. Some muscle movement is voluntary, which means it is under conscious control. For example, a person decides to open a book and read a chapter on anatomy. Other movements are involuntary, meaning they are not under conscious control, such as the contraction of your pupil in bright light. Muscle tissue is classified into three types according to structure and function: skeletal, cardiac, and smooth (Table 1).

Comparison of Structure and Properties of Muscle Tissue Types (Table 1)
Tissue Histology Function Location
Skeletal Long cylindrical fiber, striated, many peripherally located nuclei Voluntary movement, produces heat, protects organs Attached to bones and around entrance points to body (e.g., mouth, anus)
Cardiac Short, branched, striated, single central nucleus Contracts to pump blood Heart
Smooth Short, spindle-shaped, no evident striation, single nucleus in each fiber Involuntary movement, moves food, involuntary control of respiration, moves secretions, regulates flow of blood in arteries by contraction Walls of major organs and passageways
[caption id="" align="alignright" width="280"]This shows three micrographs, each depicting one of the three muscle tissues. Picture A shows skeletal muscle tissue, which is dense strips of pink tissue that somewhat resemble bacon in appearance. Many small nuclei are dispersed throughout the tissues. The nuclei are flat and elongated, with multiple nuclei clustered into each cell. Picture B shows smooth muscle, which is densely packed and looks similar to skeletal muscle except that each cell only has one oval-shaped nucleus. Picture C shows cardiac muscle. Unlike skeletal and smooth muscle cells, cardiac muscle cells are not densely packed. The cardiac cells are branched, creating a large amount of space between each muscle cell. Figure 1. Muscle Tissue. (a) Skeletal muscle cells have prominent striation and nuclei on their periphery. (b) Smooth muscle cells have a single nucleus and no visible striations. (c) Cardiac muscle cells appear striated and have a single nucleus. From top, LM × 1600, LM × 1600, LM × 1600. (Micrographs provided by the Regents of University of Michigan Medical School © 2012)[/caption] Skeletal muscle is attached to bones and its contraction makes possible locomotion, facial expressions, posture, and other voluntary movements of the body. Forty percent of your body mass is made up of skeletal muscle. Skeletal muscles generate heat as a byproduct of their contraction and thus participate in thermal homeostasis. Shivering is an involuntary contraction of skeletal muscles in response to perceived lower than normal body temperature. The muscle cell, or myocyte, develops from myoblasts derived from the mesoderm. Myocytes and their numbers remain relatively constant throughout life. Skeletal muscle tissue is arranged in bundles surrounded by connective tissue. Under the light microscope, muscle cells appear striated with many nuclei squeezed along the membranes. The striation is due to the regular alternation of the contractile proteins actin and myosin, along with the structural proteins that couple the contractile proteins to connective tissues. The cells are multinucleated as a result of the fusion of the many myoblasts that fuse to form each long muscle fiber.

Cardiac muscle forms the contractile walls of the heart. The cells of cardiac muscle, known as cardiomyocytes, also appear striated under the microscope. Unlike skeletal muscle fibers, cardiomyocytes are single cells typically with a single centrally located nucleus. A principal characteristic of cardiomyocytes is that they contract on their own intrinsic rhythms without any external stimulation. Cardiomyocyte attach to one another with specialized cell junctions called intercalated discs. Intercalated discs have both anchoring junctions and gap junctions. Attached cells form long, branching cardiac muscle fibers that are, essentially, a mechanical and electrochemical syncytium allowing the cells to synchronize their actions. The cardiac muscle pumps blood through the body and is under involuntary control. The attachment junctions hold adjacent cells together across the dynamic pressures changes of the cardiac cycle.

Smooth muscle tissue contraction is responsible for involuntary movements in the internal organs. It forms the contractile component of the digestive, urinary, and reproductive systems as well as the airways and arteries. Each cell is spindle shaped with a single nucleus and no visible striations (Figure 1).
[caption id="attachment_2961" align="aligncenter" width="150"] Watch this CrashCourse video on tissues to learn more about muscle tissue.[/caption]
]]> 2001 1977 5 0 5.1 Layers of the Skin https://pressbooks.bccampus.ca/apdouglas2017/chapter/5-1-layers-of-the-skin/ Fri, 14 Jul 2017 22:34:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2016

Learning Objectives

By the end of this section, you will be able to:
  • Identify the components of the integumentary system
  • Describe the layers of the skin and the functions of each layer
  • Identify and describe the hypodermis and deep fascia
  • Describe the role of keratinocytes and their life cycle
  • Describe the role of melanocytes in skin pigmentation
[caption id="" align="alignright" width="500"]This illustration shows a cross section of skin tissue. The outermost layer is called the epidermis, and occupies one fifth of the cross section. Several hairs are emerging from the surface. The epidermis dives around one of the hairs, forming a follicle. The middle layer is called the dermis, which occupies four fifths of the cross section. The dermis contains an erector pilli muscle connected to one of the follicles. The dermis also contains an eccrine sweat gland, composed of a bunch of tubules. One tubule travels up from the bunch, through the epidermis, opening onto the surface a pore. There are two string-like nerves travelling vertically through the dermis. The right nerve is attached to a Pacinian corpuscle, which is a yellow structure consisting of concentric ovals similar to an onion. The lowest level of the skin, the hypodermis, contains fatty tissue, arteries, and veins. Blood vessels travel from the hypodermis and connect to hair follicles and erector pilli muscle in the dermis. Figure 1. Layers of Skin. The skin is composed of two main layers: the epidermis, made of closely packed epithelial cells, and the dermis, made of dense, irregular connective tissue that houses blood vessels, hair follicles, sweat glands, and other structures. Beneath the dermis lies the hypodermis, which is composed mainly of loose connective and fatty tissues.[/caption] Although you may not typically think of the skin as an organ, it is in fact made of tissues that work together as a single structure to perform unique and critical functions. The skin and its accessory structures make up the integumentary system, which provides the body with overall protection. The skin is made of multiple layers of cells and tissues, which are held to underlying structures by connective tissue (Figure 1). The deeper layer of skin is well vascularized (has numerous blood vessels). It also has numerous sensory, and autonomic and sympathetic nerve fibers ensuring communication to and from the brain. [caption id="attachment_2963" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about the integumentary system.[/caption]
[caption id="" align="alignleft" width="280"]Part A is a micrograph showing a cross section of thin skin. The topmost layer is a thin, translucent layer with irregular texture and areas where cells are sloughing off. The deepest layer is dark purple and extends into the third layer with finger like projections. The third light purple layer contains thin bands of fibers and small, dark cells. The fourth, and deepest layer, is darker than the third layer, but is still light purple. It contains thick fiber bands that are loosely packed. Part B is a magnified view of the epidermis of thick skin. It shows the topmost layer is five times thicker than the topmost layer of thin skin. The topmost layer of thick skin is also denser and less translucent than the topmost layer of thin skin. Figure 2. Thin Skin versus Thick Skin. These slides show cross-sections of the epidermis and dermis of (a) thin and (b) thick skin. Note the significant difference in the thickness of the epithelial layer of the thick skin. From top, LM × 40, LM × 40. (Micrographs provided by the Regents of University of Michigan Medical School © 2012)[/caption]

The Epidermis

The epidermis is composed of keratinized, stratified squamous epithelium. It is made of four or five layers of epithelial cells, depending on its location in the body. It does not have any blood vessels within it (i.e., it is avascular). Skin that has four layers of cells is referred to as “thin skin.” From deep to superficial, these layers are the stratum basale, stratum spinosum, stratum granulosum, and stratum corneum. Most of the skin can be classified as thin skin. “Thick skin” is found only on the palms of the hands and the soles of the feet. It has a fifth layer, called the stratum lucidum, located between the stratum corneum and the stratum granulosum (Figure 2).

[caption id="" align="alignright" width="280"]The outer layer of cells in this micrograph is the thinnest layer and stained deep purple due to full keratinization of dead cells. The next layer occupies one quarter of the micrograph, is lightly stained, and is a dense collection of cells. The third layer from the top is mostly white, with lightly stained, loosely-packed strands radiating in random directions. The bottom-most layer is densely-packed, with thick bands of highly organized muscle tissue that are darkly stained. Figure 3. Epidermis. The epidermis is epithelium composed of multiple layers of cells. The basal layer consists of cuboidal cells, whereas the outer layers are squamous, keratinized cells, so the whole epithelium is often described as being keratinized stratified squamous epithelium. LM × 40. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption] The cells in all of the layers except the stratum basale are called keratinocytes. A keratinocyte is a cell that manufactures and stores the protein keratin. Keratin is an intracellular fibrous protein that gives hair, nails, and skin their hardness and water-resistant properties. The keratinocytes in the stratum corneum are dead and regularly slough away, being replaced by cells from the deeper layers (Figure 3).

Stratum Basale

The stratum basale (also called the stratum germinativum) is the deepest epidermal layer and attaches the epidermis to the basal lamina, below which lie the layers of the dermis. The cells in the stratum basale bond to the dermis via intertwining collagen fibers, referred to as the basement membrane. A finger-like projection, or fold, known as the dermal papilla (plural = dermal papillae) is found in the superficial portion of the dermis. Dermal papillae increase the strength of the connection between the epidermis and dermis; the greater the folding, the stronger the connections made (Figure 4).

[caption id="" align="aligncenter" width="500"]This illustration shows a cross section of the epidermis. The cells of the innermost layer, the stratum basale, are large and have a purple nucleus. The stratum basale curls around the dermis, which projects into the epidermis. The stratum basale contains four layers of large, triangle-shaped keratinocytes. Fibers are visible within the spaces between keratinocytes in the stratum basale. A melanocyte is also present in this layer. The melanocyte possesses finger-like projections extending from its main cell body. The projections branch through the extracellular spaces between nearby keratinocytes. Above the stratum basale is the stratum spinosum which consists of 8 layers of oval-shaped keratinocytes. The nucleus is present in these keratinocytes, but has faded to a lighter purple. The stratum granulosum contains five layers of keratinocytes, each containing spots in its cytoplasm, labeled the lamellar granules. The stratum lucidium contains 4 layers of diamond-shaped cells with no nucleus. The stratum corneum contains 9 layers of keratinocytes with no nucleus , nor cytoplasm. A few of the cells in the topmost layer of the stratum corneum are flaking off from the skin. Figure 4. Layers of the Epidermis. The epidermis of thick skin has five layers: stratum basale, stratum spinosum, stratum granulosum, stratum lucidum, and stratum corneum.[/caption]

The stratum basale is a single layer of cells primarily made of basal cells. A basal cell is a cuboidal-shaped stem cell that is a precursor of the keratinocytes of the epidermis. All of the keratinocytes are produced from this single layer of cells, which are constantly going through mitosis to produce new cells. As new cells are formed, the existing cells are pushed superficially away from the stratum basale. Two other cell types are found dispersed among the basal cells in the stratum basale. The first is a Merkel cell, which functions as a receptor and is responsible for stimulating sensory nerves that the brain perceives as touch. These cells are especially abundant on the surfaces of the hands and feet. The second is a melanocyte, a cell that produces the pigment melanin. Melanin gives hair and skin its color, and also helps protect the living cells of the epidermis from ultraviolet (UV) radiation damage.

In a growing fetus, fingerprints form where the cells of the stratum basale meet the papillae of the underlying dermal layer (papillary layer), resulting in the formation of the ridges on your fingers that you recognize as fingerprints. Fingerprints are unique to each individual and are used for forensic analyses because the patterns do not change with the growth and aging processes.

[caption id="" align="alignright" width="500"]This micrograph of the epidermis shows stratum corneum as a rough, darkened layer. The next layer, the stratum granulosum, contains white cells with areas of black in their cytoplasm, equal in thickness to the stratum corneum. The third layer, the stratum spinosum, contains large, grayish cells. The stratum spinosum is the thickest layer, occupying half of the micrograph. A hair follicle is embedded in this layer, which is a round structure with black, concentric spots. The fourth layer is the stratum basalis, which contains grayish cells with clear, dark nuclei, similar in thickness to the stratum corneum. The dermis is the deepest layer, and is lightly-colored with interspersed gray cells. A cross-section of a capillary is visible within the dermis. Figure 5. Cells of the Epidermis. The cells in the different layers of the epidermis originate from basal cells located in the stratum basale, yet the cells of each layer are distinctively different. EM × 2700. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Stratum Spinosum

As the name suggests, the stratum spinosum is spiny in appearance due to the protruding cell processes that join the cells via a structure called a desmosome. The desmosomes interlock with each other and strengthen the bond between the cells. It is interesting to note that the “spiny” nature of this layer is an artifact of the staining process. Unstained epidermis samples do not exhibit this characteristic appearance. The stratum spinosum is composed of eight to 10 layers of keratinocytes, formed as a result of cell division in the stratum basale (Figure 5). Interspersed among the keratinocytes of this layer is a type of dendritic cell called the Langerhans cell, which functions as a macrophage by engulfing bacteria, foreign particles, and damaged cells that occur in this layer.
[caption id="" align="alignleft" width="120"]QR Code representing a URL View the University of Michigan WebScope at http://virtualslides.med.umich.edu/Histology/EMsmallCharts/3%20Image%20Scope%20finals/065%20-%20Epidermis_001.svs/view.apml to explore the tissue sample in greater detail.[/caption]

The keratinocytes in the stratum spinosum begin the synthesis of keratin and release a water-repelling glycolipid that helps prevent water loss from the body, making the skin relatively waterproof. As new keratinocytes are produced atop the stratum basale, the keratinocytes of the stratum spinosum are pushed into the stratum granulosum.

Stratum Granulosum

The stratum granulosum has a grainy appearance due to further changes to the keratinocytes as they are pushed from the stratum spinosum. The cells (three to five layers deep) become flatter, their cell membranes thicken, and they generate large amounts of the proteins keratin, which is fibrous, and keratohyalin, which accumulates as lamellar granules within the cells (see Figure 4). These two proteins make up the bulk of the keratinocyte mass in the stratum granulosum and give the layer its grainy appearance. The nuclei and other cell organelles disintegrate as the cells die, leaving behind the keratin, keratohyalin, and cell membranes that will form the stratum lucidum, the stratum corneum, and the accessory structures of hair and nails.

Stratum Lucidum

The stratum lucidum is a smooth, seemingly translucent layer of the epidermis located just above the stratum granulosum and below the stratum corneum. This thin layer of cells is found only in the thick skin of the palms, soles, and digits. The keratinocytes that compose the stratum lucidum are dead and flattened (see Figure 4). These cells are densely packed with eleiden, a clear protein rich in lipids, derived from keratohyalin, which gives these cells their transparent (i.e., lucid) appearance and provides a barrier to water.

Stratum Corneum

The stratum corneum is the most superficial layer of the epidermis and is the layer exposed to the outside environment (see Figure 4). The increased keratinization (also called cornification) of the cells in this layer gives it its name. There are usually 15 to 30 layers of cells in the stratum corneum. This dry, dead layer helps prevent the penetration of microbes and the dehydration of underlying tissues, and provides a mechanical protection against abrasion for the more delicate, underlying layers. Cells in this layer are shed periodically and are replaced by cells pushed up from the stratum granulosum (or stratum lucidum in the case of the palms and soles of feet). The entire layer is replaced during a period of about 4 weeks. Cosmetic procedures, such as microdermabrasion, help remove some of the dry, upper layer and aim to keep the skin looking “fresh” and healthy.

[caption id="" align="alignleft" width="320"]This micrograph shows layers of skin in a cross section. The papillary layer of the dermis extends between the downward fingers of the darkly stained epidermis. The papillary layer appears finer than the reticular layer, consisting of smaller, densely-packed fibers. The reticular layer is three times thicker than the papillary layer and contains larger, thicker fibers. The fibers seem more loosely packed than those of the papillary layer, with some separated by empty spaces. Both layers of the dermis contain cells with darkly stained nuclei. Figure 6. Layers of the Dermis. This stained slide shows the two components of the dermis—the papillary layer and the reticular layer. Both are made of connective tissue with fibers of collagen extending from one to the other, making the border between the two somewhat indistinct. The dermal papillae extending into the epidermis belong to the papillary layer, whereas the dense collagen fiber bundles below belong to the reticular layer. LM × 10. (credit: modification of work by “kilbad”/Wikimedia Commons)[/caption]

Dermis

The dermis might be considered the “core” of the integumentary system (derma- = “skin”), as distinct from the epidermis (epi- = “upon” or “over”) and hypodermis (hypo- = “below”). It contains blood and lymph vessels, nerves, and other structures, such as hair follicles and sweat glands. The dermis is made of two layers of connective tissue that compose an interconnected mesh of elastin and collagenous fibers, produced by fibroblasts (Figure 6).

Papillary Layer

The papillary layer is made of loose, areolar connective tissue, which means the collagen and elastin fibers of this layer form a loose mesh. This superficial layer of the dermis projects into the stratum basale of the epidermis to form finger-like dermal papillae (see Figure 6). Within the papillary layer are fibroblasts, a small number of fat cells (adipocytes), and an abundance of small blood vessels. In addition, the papillary layer contains phagocytes, defensive cells that help fight bacteria or other infections that have breached the skin. This layer also contains lymphatic capillaries, nerve fibers, and touch receptors called the Meissner corpuscles.

Reticular Layer

Underlying the papillary layer is the much thicker reticular layer, composed of dense, irregular connective tissue. This layer is well vascularized and has a rich sensory and sympathetic nerve supply. The reticular layer appears reticulated (net-like) due to a tight meshwork of fibers. Elastin fibers provide some elasticity to the skin, enabling movement. Collagen fibers provide structure and tensile strength, with strands of collagen extending into both the papillary layer and the hypodermis. In addition, collagen binds water to keep the skin hydrated. Collagen injections and Retin-A creams help restore skin turgor by either introducing collagen externally or stimulating blood flow and repair of the dermis, respectively.

Hypodermis

The hypodermis (also called the subcutaneous layer or superficial fascia) is a layer directly below the dermis and serves to connect the skin to the underlying fascia (fibrous tissue) of the bones and muscles. It is not strictly a part of the skin, although the border between the hypodermis and dermis can be difficult to distinguish. The hypodermis consists of well-vascularized, loose, areolar connective tissue and adipose tissue, which functions as a mode of fat storage and provides insulation and cushioning for the integument.

Pigmentation

The color of skin is influenced by a number of pigments, including melanin, carotene, and hemoglobin. Recall that melanin is produced by cells called melanocytes, which are found scattered throughout the stratum basale of the epidermis. The melanin is transferred into the keratinocytes via a cellular vesicle called a melanosome (Figure 7).

[caption id="" align="aligncenter" width="550"]This figure consists of two diagrams side by side. The right diagram shows development of light colored skin; the left shows development of dark-colored skin. In both, a brown melanocyte sits at the border between the dermis and epidermis. The melanocyte has a large nucleus and six finger-like extensions. These reach between cells of the stratum basalis. Sections of the extensions detach and travel through the skins. These are melanosomes. In the left diagram, both the melanocyte and melanosomes contain melanin particles, shown as dark dots. Melanosomes travel upwards to outer skin layers, releasing melanin. As a result, keratinocytes in the left diagram contain several melanin particles that darken skin color. In light colored skin, the melanocyte contains no melanin. It still releases melanosomes into upper layers of the skin; however, these melanosomes contain no melanin. Therefore, the skin does not darken and remains light. Figure 7. Skin Pigmentation. The relative coloration of the skin depends of the amount of melanin produced by melanocytes in the stratum basale and taken up by keratinocytes.[/caption]

Melanin occurs in two primary forms. Eumelanin exists as black and brown, whereas pheomelanin provides a red color. Dark-skinned individuals produce more melanin than those with pale skin. Exposure to the UV rays of the sun or a tanning salon causes melanin to be manufactured and built up in keratinocytes, as sun exposure stimulates keratinocytes to secrete chemicals that stimulate melanocytes. The accumulation of melanin in keratinocytes results in the darkening of the skin, or a tan. This increased melanin accumulation protects the DNA of epidermal cells from UV ray damage and the breakdown of folic acid, a nutrient necessary for our health and well-being. In contrast, too much melanin can interfere with the production of vitamin D, an important nutrient involved in calcium absorption. Thus, the amount of melanin present in our skin is dependent on a balance between available sunlight and folic acid destruction, and protection from UV radiation and vitamin D production.

It requires about 10 days after initial sun exposure for melanin synthesis to peak, which is why pale-skinned individuals tend to suffer sunburns of the epidermis initially. Dark-skinned individuals can also get sunburns, but are more protected than are pale-skinned individuals. Melanosomes are temporary structures that are eventually destroyed by fusion with lysosomes; this fact, along with melanin-filled keratinocytes in the stratum corneum sloughing off, makes tanning impermanent.

Too much sun exposure can eventually lead to wrinkling due to the destruction of the cellular structure of the skin, and in severe cases, can cause sufficient DNA damage to result in skin cancer. When there is an irregular accumulation of melanocytes in the skin, freckles appear. Moles are larger masses of melanocytes, and although most are benign, they should be monitored for changes that might indicate the presence of cancer (Figure 8).
[caption id="" align="aligncenter" width="380"]Five photos of moles. The three upper photos show moles that are small, flat, and dark brown. The bottom left photo shows a dark black mole that is raised above the skin. The bottom right photo shows a large, raised, reddish mole with protruding hairs. Figure 8. Moles. Moles range from benign accumulations of melanocytes to melanomas. These structures populate the landscape of our skin. (credit: the National Cancer Institute)[/caption]
]]> 2016 2005 2 0 5.2 Accessory Structures of the Skin https://pressbooks.bccampus.ca/apdouglas2017/chapter/5-2-accessory-structures-of-the-skin/ Fri, 14 Jul 2017 22:35:42 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2021

Learning Objectives

By the end of this section, you will be able to:
  • Identify the accessory structures of the skin
  • Describe the structure and function of hair and nails
  • Describe the structure and function of sweat glands and sebaceous glands

Accessory structures of the skin include hair, nails, sweat glands, and sebaceous glands. These structures embryologically originate from the epidermis and can extend down through the dermis into the hypodermis.

[caption id="attachment_2965" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about the accessory structures![/caption]
[caption id="" align="alignright" width="300"]This diagram shows a cross section of the skin containing a hair follicle. The follicle is teardrop shaped. Its enlarged base, labeled the hair bulb, is embedded in the hypodermis. The outermost layer of the follicle is the epidermis, which invaginates from the skin surface to envelope the follicle. Within the epidermis is the outer root sheath, which is only present on the hair bulb. It does not extend up the shaft of the hair. Within the outer root sheath is the inner root sheath. The inner root sheath extends about half of the way up the hair shaft, ending midway through the dermis. The hair matrix is the innermost layer. The hair matrix surrounds the bottom of the hair shaft where it is embedded within the hair bulb. The hair shaft, in itself, contains three layers: the outermost cuticle, a middle layer called the cortex, and an innermost layer called the medulla. Figure 1. Hair. Hair follicles originate in the epidermis and have many different parts.[/caption]

Hair

Hair is a keratinous filament growing out of the epidermis. It is primarily made of dead, keratinized cells. Strands of hair originate in an epidermal penetration of the dermis called the hair follicle. The hair shaft is the part of the hair not anchored to the follicle, and much of this is exposed at the skin’s surface. The rest of the hair, which is anchored in the follicle, lies below the surface of the skin and is referred to as the hair root. The hair root ends deep in the dermis at the hair bulb, and includes a layer of mitotically active basal cells called the hair matrix. The hair bulb surrounds the hair papilla, which is made of connective tissue and contains blood capillaries and nerve endings from the dermis (Figure 1).

Just as the basal layer of the epidermis forms the layers of epidermis that get pushed to the surface as the dead skin on the surface sheds, the basal cells of the hair bulb divide and push cells outward in the hair root and shaft as the hair grows. The medulla forms the central core of the hair, which is surrounded by the cortex, a layer of compressed, keratinized cells that is covered by an outer layer of very hard, keratinized cells known as the cuticle. These layers are depicted in a longitudinal cross-section of the hair follicle (Figure 2), although not all hair has a medullary layer. [caption id="" align="alignleft" width="380"]This micrograph is of the base of a hair follicle. The protruding hair is largely transparent, with only its dark outline visible. The inner root sheath is visible surrounding the very bottom of the hair as a circle of cells with dark-staining nuclei. The inner sheath extends up the hair shaft. The outer root sheath is much thicker than the inner root sheath, consisting of a large oval of lighter staining cells. The oval surrounds the bottom of the hair and extends into the hypodermis. Figure 2. Hair Follicle. The slide shows a cross-section of a hair follicle. Basal cells of the hair matrix in the center differentiate into cells of the inner root sheath. Basal cells at the base of the hair root form the outer root sheath. LM × 4. (credit: modification of work by “kilbad”/Wikimedia Commons)[/caption] Hair texture (straight, curly) is determined by the shape and structure of the cortex, and to the extent that it is present, the medulla. The shape and structure of these layers are, in turn, determined by the shape of the hair follicle. Hair growth begins with the production of keratinocytes by the basal cells of the hair bulb. As new cells are deposited at the hair bulb, the hair shaft is pushed through the follicle toward the surface. Keratinization is completed as the cells are pushed to the skin surface to form the shaft of hair that is externally visible. The external hair is completely dead and composed entirely of keratin. For this reason, our hair does not have sensation. Furthermore, you can cut your hair or shave without damaging the hair structure because the cut is superficial. Most chemical hair removers also act superficially; however, electrolysis and yanking both attempt to destroy the hair bulb so hair cannot grow.
The wall of the hair follicle is made of three concentric layers of cells. The cells of the internal root sheath surround the root of the growing hair and extend just up to the hair shaft. They are derived from the basal cells of the hair matrix. The external root sheath, which is an extension of the epidermis, encloses the hair root. It is made of basal cells at the base of the hair root and tends to be more keratinous in the upper regions. The glassy membrane is a thick, clear connective tissue sheath covering the hair root, connecting it to the tissue of the dermis.

Hair serves a variety of functions, including protection, sensory input, thermoregulation, and communication. For example, hair on the head protects the skull from the sun. The hair in the nose and ears, and around the eyes (eyelashes) defends the body by trapping and excluding dust particles that may contain allergens and microbes. Hair of the eyebrows prevents sweat and other particles from dripping into and bothering the eyes. Hair also has a sensory function due to sensory innervation by a hair root plexus surrounding the base of each hair follicle. Hair is extremely sensitive to air movement or other disturbances in the environment, much more so than the skin surface. This feature is also useful for the detection of the presence of insects or other potentially damaging substances on the skin surface. Each hair root is connected to a smooth muscle called the arrector pili that contracts in response to nerve signals from the sympathetic nervous system, making the external hair shaft “stand up.” The primary purpose for this is to trap a layer of air to add insulation. This is visible in humans as goose bumps and even more obvious in animals, such as when a frightened cat raises its fur. Of course, this is much more obvious in organisms with a heavier coat than most humans, such as dogs and cats.

Hair Growth

Hair grows and is eventually shed and replaced by new hair. This occurs in three phases. The first is the anagen phase, during which cells divide rapidly at the root of the hair, pushing the hair shaft up and out. The length of this phase is measured in years, typically from 2 to 7 years. The catagen phase lasts only 2 to 3 weeks, and marks a transition from the hair follicle’s active growth. Finally, during the telogen phase, the hair follicle is at rest and no new growth occurs. At the end of this phase, which lasts about 2 to 4 months, another anagen phase begins. The basal cells in the hair matrix then produce a new hair follicle, which pushes the old hair out as the growth cycle repeats itself. Hair typically grows at the rate of 0.3 mm per day during the anagen phase. On average, 50 hairs are lost and replaced per day. Hair loss occurs if there is more hair shed than what is replaced and can happen due to hormonal or dietary changes. Hair loss can also result from the aging process, or the influence of hormones.

Hair Color

Similar to the skin, hair gets its color from the pigment melanin, produced by melanocytes in the hair papilla. Different hair color results from differences in the type of melanin, which is genetically determined. As a person ages, the melanin production decreases, and hair tends to lose its color and becomes gray and/or white.

Nails

[caption id="" align="alignright" width="520"]These two images show anatomy of the fingernail region. The top image shows a dorsal view of a finger. The proximal nail fold is the part underneath where the skin of the finger connects with the edge of the nail. The eponychium is a thin, pink layer between the white proximal edge of the nail (the lunula), and the edge of the finger skin. The lunula appears as a crescent-shaped white area at the proximal edge of the pink-shaded nail. The lateral nail folds are where the sides of the nail contact the finger skin. The distal edge of the nail is white and is called the free edge. An arrow indicates that the nail grows distally out from the proximal nail fold. The lower image shows a lateral view of the nail bed anatomy. In this view, one can see how the edge of the nail is located just proximal to the nail fold. This end of the nail, from which the nail grows, is called the nail root. Figure 3. Nails. The nail is an accessory structure of the integumentary system.[/caption] The nail bed is a specialized structure of the epidermis that is found at the tips of our fingers and toes. The nail body is formed on the nail bed, and protects the tips of our fingers and toes as they are the farthest extremities and the parts of the body that experience the maximum mechanical stress (Figure 3). In addition, the nail body forms a back-support for picking up small objects with the fingers. The nail body is composed of densely packed dead keratinocytes. The epidermis in this part of the body has evolved a specialized structure upon which nails can form. The nail body forms at the nail root, which has a matrix of proliferating cells from the stratum basale that enables the nail to grow continuously. The lateral nail fold overlaps the nail on the sides, helping to anchor the nail body. The nail fold that meets the proximal end of the nail body forms the nail cuticle, also called the eponychium. The nail bed is rich in blood vessels, making it appear pink, except at the base, where a thick layer of epithelium over the nail matrix forms a crescent-shaped region called the lunula (the “little moon”). The area beneath the free edge of the nail, furthest from the cuticle, is called the hyponychium. It consists of a thickened layer of stratum corneum.
[caption id="" align="alignleft" width="350"]This diagram shows an eccrine sweat gland embedded in a cross section of skin tissue. The eccrine sweat gland is a bundle of white tubes embedded in the dermis. A single white tube travels up from the bundle and opens on to the surface of the epidermis. The opening is called a pore. There are several pores on the small block of skin portrayed in this diagram. Figure 4. Eccrine Gland. Eccrine glands are coiled glands in the dermis that release sweat that is mostly water.[/caption]

Sweat Glands

When the body becomes warm, sudoriferous glands produce sweat to cool the body. Sweat glands develop from epidermal projections into the dermis and are classified as merocrine glands; that is, the secretions are excreted by exocytosis through a duct without affecting the cells of the gland. There are two types of sweat glands, each secreting slightly different products.

An eccrine sweat gland is type of gland that produces a hypotonic sweat for thermoregulation. These glands are found all over the skin’s surface, but are especially abundant on the palms of the hand, the soles of the feet, and the forehead (Figure 4). They are coiled glands lying deep in the dermis, with the duct rising up to a pore on the skin surface, where the sweat is released. This type of sweat, released by exocytosis, is hypotonic and composed mostly of water, with some salt, antibodies, traces of metabolic waste, and dermicidin, an antimicrobial peptide. Eccrine glands are a primary component of thermoregulation in humans and thus help to maintain homeostasis.

An apocrine sweat gland is usually associated with hair follicles in densely hairy areas, such as armpits and genital regions. Apocrine sweat glands are larger than eccrine sweat glands and lie deeper in the dermis, sometimes even reaching the hypodermis, with the duct normally emptying into the hair follicle. In addition to water and salts, apocrine sweat includes organic compounds that make the sweat thicker and subject to bacterial decomposition and subsequent smell. The release of this sweat is under both nervous and hormonal control, and plays a role in the poorly understood human pheromone response. Most commercial antiperspirants use an aluminum-based compound as their primary active ingredient to stop sweat. When the antiperspirant enters the sweat gland duct, the aluminum-based compounds precipitate due to a change in pH and form a physical block in the duct, which prevents sweat from coming out of the pore.

Sebaceous Glands

A sebaceous gland is a type of oil gland that is found all over the body and helps to lubricate and waterproof the skin and hair. Most sebaceous glands are associated with hair follicles. They generate and excrete sebum, a mixture of lipids, onto the skin surface, thereby naturally lubricating the dry and dead layer of keratinized cells of the stratum corneum, keeping it pliable. The fatty acids of sebum also have antibacterial properties, and prevent water loss from the skin in low-humidity environments. The secretion of sebum is stimulated by hormones, many of which do not become active until puberty. Thus, sebaceous glands are relatively inactive during childhood.

]]> 2021 2005 3 0 9.1 Classification of Joints https://pressbooks.bccampus.ca/apdouglas2017/chapter/9-1-classification-of-joints/ Fri, 14 Jul 2017 22:46:43 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2114

Learning Objectives

By the end of this section, you will be able to:
  • Distinguish between the functional and structural classifications for joints
  • Describe the three functional types of joints and give an example of each
  • List the three types of diarthrodial joints

A joint, also called an articulation, is any place where adjacent bones or bone and cartilage come together (articulate with each other) to form a connection. Joints are classified both structurally and functionally. Structural classifications of joints take into account whether the adjacent bones are strongly anchored to each other by fibrous connective tissue or cartilage, or whether the adjacent bones articulate with each other within a fluid-filled space called a joint cavity. Functional classifications describe the degree of movement available between the bones, ranging from immobile, to slightly mobile, to freely moveable joints. The amount of movement available at a particular joint of the body is related to the functional requirements for that joint. Thus immobile or slightly moveable joints serve to protect internal organs, give stability to the body, and allow for limited body movement. In contrast, freely moveable joints allow for much more extensive movements of the body and limbs.

Structural Classification of Joints

The structural classification of joints is based on whether the articulating surfaces of the adjacent bones are directly connected by fibrous connective tissue or cartilage, or whether the articulating surfaces contact each other within a fluid-filled joint cavity. These differences serve to divide the joints of the body into three structural classifications. A fibrous joint is where the adjacent bones are united by fibrous connective tissue. At a cartilaginous joint, the bones are joined by hyaline cartilage or fibrocartilage. At a synovial joint, the articulating surfaces of the bones are not directly connected, but instead come into contact with each other within a joint cavity that is filled with a lubricating fluid. Synovial joints allow for free movement between the bones and are the most common joints of the body.

Functional Classification of Joints

The functional classification of joints is determined by the amount of mobility found between the adjacent bones. Joints are thus functionally classified as a synarthrosis or immobile joint, an amphiarthrosis or slightly moveable joint, or as a diarthrosis, which is a freely moveable joint (arthroun = “to fasten by a joint”). Depending on their location, fibrous joints may be functionally classified as a synarthrosis (immobile joint) or an amphiarthrosis (slightly mobile joint). Cartilaginous joints are also functionally classified as either a synarthrosis or an amphiarthrosis joint. All synovial joints are functionally classified as a diarthrosis joint.

Synarthrosis

An immobile or nearly immobile joint is called a synarthrosis. The immobile nature of these joints provide for a strong union between the articulating bones. This is important at locations where the bones provide protection for internal organs. Examples include sutures, the fibrous joints between the bones of the skull that surround and protect the brain (Figure 1), and the manubriosternal joint, the cartilaginous joint that unites the manubrium and body of the sternum for protection of the heart.

[caption id="" align="aligncenter" width="340"]This image shows the lateral view of the human skeleton. The lambdoid, coronal, and squamous sutures are labeled. Figure 1. Suture Joints of Skull. The suture joints of the skull are an example of a synarthrosis, an immobile or essentially immobile joint.[/caption]

Amphiarthrosis

An amphiarthrosis is a joint that has limited mobility. An example of this type of joint is the cartilaginous joint that unites the bodies of adjacent vertebrae. Filling the gap between the vertebrae is a thick pad of fibrocartilage called an intervertebral disc (Figure 2). Each intervertebral disc strongly unites the vertebrae but still allows for a limited amount of movement between them. However, the small movements available between adjacent vertebrae can sum together along the length of the vertebral column to provide for large ranges of body movements.

Another example of an amphiarthrosis is the pubic symphysis of the pelvis. This is a cartilaginous joint in which the pubic regions of the right and left hip bones are strongly anchored to each other by fibrocartilage. This joint normally has very little mobility. The strength of the pubic symphysis is important in conferring weight-bearing stability to the pelvis.
[caption id="" align="aligncenter" width="330"]This image shows the lateral view of the intervertebral disc located between two vertebral discs. Figure 2. Intervertebral Disc. An intervertebral disc unites the bodies of adjacent vertebrae within the vertebral column. Each disc allows for limited movement between the vertebrae and thus functionally forms an amphiarthrosis type of joint. Intervertebral discs are made of fibrocartilage and thereby structurally form a symphysis type of cartilaginous joint.[/caption]

Diarthrosis

A freely mobile joint is classified as a diarthrosis. These types of joints include all synovial joints of the body, which provide the majority of body movements. Most diarthrotic joints are found in the appendicular skeleton and thus give the limbs a wide range of motion. These joints are divided into three categories, based on the number of axes of motion provided by each. An axis in anatomy is described as the movements in reference to the three anatomical planes: transverse, frontal, and sagittal. Thus, diarthroses are classified as uniaxial (for movement in one plane), biaxial (for movement in two planes), or multiaxial joints (for movement in all three anatomical planes).

A uniaxial joint only allows for a motion in a single plane (around a single axis). The elbow joint, which only allows for bending or straightening, is an example of a uniaxial joint. A biaxial joint allows for motions within two planes. An example of a biaxial joint is a metacarpophalangeal joint (knuckle joint) of the hand. The joint allows for movement along one axis to produce bending or straightening of the finger, and movement along a second axis, which allows for spreading of the fingers away from each other and bringing them together. A joint that allows for the several directions of movement is called a multiaxial joint (polyaxial or triaxial joint). This type of diarthrotic joint allows for movement along three axes (Figure 3). The shoulder and hip joints are multiaxial joints. They allow the upper or lower limb to move in an anterior-posterior direction and a medial-lateral direction. In addition, the limb can also be rotated around its long axis. This third movement results in rotation of the limb so that its anterior surface is moved either toward or away from the midline of the body.

[caption id="" align="aligncenter" width="450"]This image shows a multiaxial joint. The left panel shows the acetabulum of the hip bone and the head of the femur. The right panel shows a simplified ball-and-socket joint structure to illustrate the movement of the hip joint. Figure 3. Multiaxial Joint. A multiaxial joint, such as the hip joint, allows for three types of movement: anterior-posterior, medial-lateral, and rotational.[/caption] [caption id="attachment_2967" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about joints![/caption]
]]> 2114 2108 2 0 10.2 Skeletal Muscle https://pressbooks.bccampus.ca/apdouglas2017/chapter/10-2-skeletal-muscle/ Fri, 14 Jul 2017 22:48:20 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2147

Learning Objectives

By the end of this section, you will be able to:
  • Describe the layers of connective tissues packaging skeletal muscle
  • Explain how muscles work with tendons to move the body
  • Identify areas of the skeletal muscle fibers
  • Describe excitation-contraction coupling

The best-known feature of skeletal muscle is its ability to contract and cause movement. Skeletal muscles act not only to produce movement but also to stop movement, such as resisting gravity to maintain posture. Small, constant adjustments of the skeletal muscles are needed to hold a body upright or balanced in any position. Muscles also prevent excess movement of the bones and joints, maintaining skeletal stability and preventing skeletal structure damage or deformation. Joints can become misaligned or dislocated entirely by pulling on the associated bones; muscles work to keep joints stable. Skeletal muscles are located throughout the body at the openings of internal tracts to control the movement of various substances. These muscles allow functions, such as swallowing, urination, and defecation, to be under voluntary control. Skeletal muscles also protect internal organs (particularly abdominal and pelvic organs) by acting as an external barrier or shield to external trauma and by supporting the weight of the organs.

Skeletal muscles contribute to the maintenance of homeostasis in the body by generating heat. Muscle contraction requires energy, and when ATP is broken down, heat is produced. This heat is very noticeable during exercise, when sustained muscle movement causes body temperature to rise, and in cases of extreme cold, when shivering produces random skeletal muscle contractions to generate heat.

Each skeletal muscle is an organ that consists of various integrated tissues. These tissues include the skeletal muscle fibers, blood vessels, nerve fibers, and connective tissue. Each skeletal muscle has three layers of connective tissue (called “mysia”) that enclose it and provide structure to the muscle as a whole, and also compartmentalize the muscle fibers within the muscle (Figure 1). Each muscle is wrapped in a sheath of dense, irregular connective tissue called the epimysium, which allows a muscle to contract and move powerfully while maintaining its structural integrity. The epimysium also separates muscle from other tissues and organs in the area, allowing the muscle to move independently.

[caption id="" align="aligncenter" width="420"]This figure shows the structure of muscle fibers. The top panel shows a skeleton muscle fiber, and a magnified view of the muscle fascicles are shown. The middle panel shows a magnified view of the muscle fascicles with the muscle fibers, perimysium and the endomysium. The bottom panel shows the structure of the muscle fiber with the sarcolemma highlighted. Figure 1. The Three Connective Tissue Layers. Bundles of muscle fibers, called fascicles, are covered by the perimysium. Muscle fibers are covered by the endomysium.[/caption]

Inside each skeletal muscle, muscle fibers are organized into individual bundles, each called a fascicle, by a middle layer of connective tissue called the perimysium. This fascicular organization is common in muscles of the limbs; it allows the nervous system to trigger a specific movement of a muscle by activating a subset of muscle fibers within a bundle, or fascicle of the muscle. Inside each fascicle, each muscle fiber is encased in a thin connective tissue layer of collagen and reticular fibers called the endomysium. The endomysium contains the extracellular fluid and nutrients to support the muscle fiber. These nutrients are supplied via blood to the muscle tissue.

In skeletal muscles that work with tendons to pull on bones, the collagen in the three tissue layers (the mysia) intertwines with the collagen of a tendon. At the other end of the tendon, it fuses with the periosteum coating the bone. The tension created by contraction of the muscle fibers is then transferred though the mysia, to the tendon, and then to the periosteum to pull on the bone for movement of the skeleton. In other places, the mysia may fuse with a broad, tendon-like sheet called an aponeurosis, or to fascia, the connective tissue between skin and bones. The broad sheet of connective tissue in the lower back that the latissimus dorsi muscles (the “lats”) fuse into is an example of an aponeurosis.

Every skeletal muscle is also richly supplied by blood vessels for nourishment, oxygen delivery, and waste removal. In addition, every muscle fiber in a skeletal muscle is supplied by the axon branch of a somatic motor neuron, which signals the fiber to contract. Unlike cardiac and smooth muscle, the only way to functionally contract a skeletal muscle is through signaling from the nervous system.

Skeletal Muscle Fibers

Because skeletal muscle cells are long and cylindrical, they are commonly referred to as muscle fibers. Skeletal muscle fibers can be quite large for human cells, with diameters up to 100 μm and lengths up to 30 cm (11.8 in) in the Sartorius of the upper leg. During early development, embryonic myoblasts, each with its own nucleus, fuse with up to hundreds of other myoblasts to form the multinucleated skeletal muscle fibers. Multiple nuclei mean multiple copies of genes, permitting the production of the large amounts of proteins and enzymes needed for muscle contraction.

Some other terminology associated with muscle fibers is rooted in the Greek sarco, which means “flesh.” The plasma membrane of muscle fibers is called the sarcolemma, the cytoplasm is referred to as sarcoplasm, and the specialized smooth endoplasmic reticulum, which stores, releases, and retrieves calcium ions (Ca++) is called the sarcoplasmic reticulum (SR) (Figure 2). As will soon be described, the functional unit of a skeletal muscle fiber is the sarcomere, a highly organized arrangement of the contractile myofilaments actin (thin filament) and myosin (thick filament), along with other support proteins.

[caption id="" align="aligncenter" width="420"]This figure shows the structure of the muscle fibers. In the top panel, a sarcolemma is shown with the major parts labeled. In the bottom panel, a magnified view of a single myofibril is shown and the major parts are labeled. Figure 2. Muscle Fiber. A skeletal muscle fiber is surrounded by a plasma membrane called the sarcolemma, which contains sarcoplasm, the cytoplasm of muscle cells. A muscle fiber is composed of many fibrils, which give the cell its striated appearance.[/caption]

The Sarcomere

The striated appearance of skeletal muscle fibers is due to the arrangement of the myofilaments of actin and myosin in sequential order from one end of the muscle fiber to the other. Each packet of these microfilaments and their regulatory proteins, troponin and tropomyosin (along with other proteins) is called a sarcomere.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this video to learn more about macro- and microstructures of skeletal muscles.[/caption] [caption id="attachment_2969" align="aligncenter" width="150"] Watch this CrashCourse video to find out more about skeletal muscle structure.[/caption]

The sarcomere is the functional unit of the muscle fiber. The sarcomere itself is bundled within the myofibril that runs the entire length of the muscle fiber and attaches to the sarcolemma at its end. As myofibrils contract, the entire muscle cell contracts. Because myofibrils are only approximately 1.2 μm in diameter, hundreds to thousands (each with thousands of sarcomeres) can be found inside one muscle fiber. Each sarcomere is approximately 2 μm in length with a three-dimensional cylinder-like arrangement and is bordered by structures called Z-discs (also called Z-lines, because pictures are two-dimensional), to which the actin myofilaments are anchored (Figure 3). Because the actin and its troponin-tropomyosin complex (projecting from the Z-discs toward the center of the sarcomere) form strands that are thinner than the myosin, it is called the thin filament of the sarcomere. Likewise, because the myosin strands and their multiple heads (projecting from the center of the sarcomere, toward but not all to way to, the Z-discs) have more mass and are thicker, they are called the thick filament of the sarcomere.

[caption id="" align="aligncenter" width="390"]This figure shows the structure of thick and thin filaments. On the top of the image a sarcomere is shown with the H zone, Z line and M lines labeled. To the right of the bottom panel, the structure of the thick filament is shown in detail. To the left of the bottom panel, the structure of a thin filament is shown in detail. Figure 3. The Sarcomere. The sarcomere, the region from one Z-line to the next Z-line, is the functional unit of a skeletal muscle fiber.[/caption]

The Neuromuscular Junction

Another specialization of the skeletal muscle is the site where a motor neuron’s terminal meets the muscle fiber—called the neuromuscular junction (NMJ). This is where the muscle fiber first responds to signaling by the motor neuron. Every skeletal muscle fiber in every skeletal muscle is innervated by a motor neuron at the NMJ. Excitation signals from the neuron are the only way to functionally activate the fiber to contract.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this video to learn more about what happens at the NMJ.[/caption]

Excitation-Contraction Coupling

All living cells have membrane potentials, or electrical gradients across their membranes. The inside of the membrane is usually around -60 to -90 mV, relative to the outside. This is referred to as a cell’s membrane potential. Neurons and muscle cells can use their membrane potentials to generate electrical signals. They do this by controlling the movement of charged particles, called ions, across their membranes to create electrical currents. This is achieved by opening and closing specialized proteins in the membrane called ion channels. Although the currents generated by ions moving through these channel proteins are very small, they form the basis of both neural signaling and muscle contraction.

Both neurons and skeletal muscle cells are electrically excitable, meaning that they are able to generate action potentials. An action potential is a special type of electrical signal that can travel along a cell membrane as a wave. This allows a signal to be transmitted quickly and faithfully over long distances.

Although the term excitation-contraction coupling confuses or scares some students, it comes down to this: for a skeletal muscle fiber to contract, its membrane must first be “excited”—in other words, it must be stimulated to fire an action potential. The muscle fiber action potential, which sweeps along the sarcolemma as a wave, is “coupled” to the actual contraction through the release of calcium ions (Ca++) from the SR. Once released, the Ca++ interacts with the shielding proteins, forcing them to move aside so that the actin-binding sites are available for attachment by myosin heads. The myosin then pulls the actin filaments toward the center, shortening the muscle fiber.

In skeletal muscle, this sequence begins with signals from the somatic motor division of the nervous system. In other words, the “excitation” step in skeletal muscles is always triggered by signaling from the nervous system (Figure 4).

[caption id="" align="aligncenter" width="380"]Alt text to come. Figure 4. Motor End-Plate and Innervation. At the NMJ, the axon terminal releases ACh. The motor end-plate is the location of the ACh-receptors in the muscle fiber sarcolemma. When ACh molecules are released, they diffuse across a minute space called the synaptic cleft and bind to the receptors.[/caption]

The motor neurons that tell the skeletal muscle fibers to contract originate in the spinal cord, with a smaller number located in the brainstem for activation of skeletal muscles of the face, head, and neck. These neurons have long processes, called axons, which are specialized to transmit action potentials long distances— in this case, all the way from the spinal cord to the muscle itself (which may be up to three feet away). The axons of multiple neurons bundle together to form nerves, like wires bundled together in a cable.

Signaling begins when a neuronal action potential travels along the axon of a motor neuron, and then along the individual branches to terminate at the NMJ. At the NMJ, the axon terminal releases a chemical messenger, or neurotransmitter, called acetylcholine (ACh). The ACh molecules diffuse across a minute space called the synaptic cleft and bind to ACh receptors located within the motor end-plate of the sarcolemma on the other side of the synapse. Once ACh binds, a channel in the ACh receptor opens and positively charged ions can pass through into the muscle fiber, causing it to depolarize, meaning that the membrane potential of the muscle fiber becomes less negative (closer to zero.)

As the membrane depolarizes, another set of ion channels called voltage-gated sodium channels are triggered to open. Sodium ions enter the muscle fiber, and an action potential rapidly spreads (or “fires”) along the entire membrane to initiate excitation-contraction coupling.

Things happen very quickly in the world of excitable membranes (just think about how quickly you can snap your fingers as soon as you decide to do it). Immediately following depolarization of the membrane, it repolarizes, re-establishing the negative membrane potential. Meanwhile, the ACh in the synaptic cleft is degraded by the enzyme acetylcholinesterase (AChE) so that the ACh cannot rebind to a receptor and reopen its channel, which would cause unwanted extended muscle excitation and contraction.

Propagation of an action potential along the sarcolemma is the excitation portion of excitation-contraction coupling. Recall that this excitation actually triggers the release of calcium ions (Ca++) from its storage in the cell’s SR. For the action potential to reach the membrane of the SR, there are periodic invaginations in the sarcolemma, called T-tubules (“T” stands for “transverse”). You will recall that the diameter of a muscle fiber can be up to 100 μm, so these T-tubules ensure that the membrane can get close to the SR in the sarcoplasm. The arrangement of a T-tubule with the membranes of SR on either side is called a triad (Figure 5). The triad surrounds the cylindrical structure called a myofibril, which contains actin and myosin.

[caption id="" align="aligncenter" width="350"]Alt text to come. Figure 5. The T-tubule. Narrow T-tubules permit the conduction of electrical impulses. The SR functions to regulate intracellular levels of calcium. Two terminal cisternae (where enlarged SR connects to the T-tubule) and one T-tubule comprise a triad—a “threesome” of membranes, with those of SR on two sides and the T-tubule sandwiched between them.[/caption]

The T-tubules carry the action potential into the interior of the cell, which triggers the opening of calcium channels in the membrane of the adjacent SR, causing Ca++ to diffuse out of the SR and into the sarcoplasm. It is the arrival of Ca++ in the sarcoplasm that initiates contraction of the muscle fiber by its contractile units, or sarcomeres.

]]> 2147 2135 3 0 18.1 An Overview of Blood https://pressbooks.bccampus.ca/apdouglas2017/chapter/18-1-an-overview-of-blood/ Fri, 14 Jul 2017 22:56:02 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2200

Learning Objectives

By the end of this section, you will be able to:
  • Identify the primary functions of blood in transportation, defense, and maintenance of homeostasis
  • Name the fluid component of blood and the three major types of formed elements, and identify their relative proportions in a blood sample
  • Discuss the unique physical characteristics of blood
  • Identify the composition of blood plasma, including its most important solutes and plasma proteins

Recall that blood is a connective tissue. Like all connective tissues, it is made up of cellular elements and an extracellular matrix. The cellular elements—referred to as the formed elements—include red blood cells (RBCs), white blood cells (WBCs), and cell fragments called platelets. The extracellular matrix, called plasma, makes blood unique among connective tissues because it is fluid. This fluid, which is mostly water, perpetually suspends the formed elements and enables them to circulate throughout the body within the cardiovascular system.

Functions of Blood

The primary function of blood is to deliver oxygen and nutrients to and remove wastes from body cells, but that is only the beginning of the story. The specific functions of blood also include defense, distribution of heat, and maintenance of homeostasis.

Transportation

Nutrients from the foods you eat are absorbed in the digestive tract. Most of these travel in the bloodstream directly to the liver, where they are processed and released back into the bloodstream for delivery to body cells. Oxygen from the air you breathe diffuses into the blood, which moves from the lungs to the heart, which then pumps it out to the rest of the body. Moreover, endocrine glands scattered throughout the body release their products, called hormones, into the bloodstream, which carries them to distant target cells. Blood also picks up cellular wastes and byproducts, and transports them to various organs for removal. For instance, blood moves carbon dioxide to the lungs for exhalation from the body, and various waste products are transported to the kidneys and liver for excretion from the body in the form of urine or bile.

Defense

Many types of WBCs protect the body from external threats, such as disease-causing bacteria that have entered the bloodstream in a wound. Other WBCs seek out and destroy internal threats, such as cells with mutated DNA that could multiply to become cancerous, or body cells infected with viruses.

When damage to the vessels results in bleeding, blood platelets and certain proteins dissolved in the plasma, the fluid portion of the blood, interact to block the ruptured areas of the blood vessels involved. This protects the body from further blood loss.

Maintenance of Homeostasis

Recall that body temperature is regulated via a classic negative-feedback loop. If you were exercising on a warm day, your rising core body temperature would trigger several homeostatic mechanisms, including increased transport of blood from your core to your body periphery, which is typically cooler. As blood passes through the vessels of the skin, heat would be dissipated to the environment, and the blood returning to your body core would be cooler. In contrast, on a cold day, blood is diverted away from the skin to maintain a warmer body core. In extreme cases, this may result in frostbite.

Blood also helps to maintain the chemical balance of the body. Proteins and other compounds in blood act as buffers, which thereby help to regulate the pH of body tissues. Blood also helps to regulate the water content of body cells.

Composition of Blood

You have probably had blood drawn from a superficial vein in your arm, which was then sent to a lab for analysis. Some of the most common blood tests—for instance, those measuring lipid or glucose levels in plasma—determine which substances are present within blood and in what quantities. Other blood tests check for the composition of the blood itself, including the quantities and types of formed elements.

One such test, called a hematocrit, measures the percentage of RBCs, clinically known as erythrocytes, in a blood sample. It is performed by spinning the blood sample in a specialized centrifuge, a process that causes the heavier elements suspended within the blood sample to separate from the lightweight, liquid plasma (Figure 1). Because the heaviest elements in blood are the erythrocytes, these settle at the very bottom of the hematocrit tube. Located above the erythrocytes is a pale, thin layer composed of the remaining formed elements of blood. These are the WBCs, clinically known as leukocytes, and the platelets, cell fragments also called thrombocytes. This layer is referred to as the buffy coat because of its color; it normally constitutes less than 1 percent of a blood sample. Above the buffy coat is the blood plasma, normally a pale, straw-colored fluid, which constitutes the remainder of the sample.

The volume of erythrocytes after centrifugation is also commonly referred to as packed cell volume (PCV). In normal blood, about 45 percent of a sample is erythrocytes. The hematocrit of any one sample can vary significantly, however, about 36–50 percent, according to gender and other factors. Normal hematocrit values for females range from 37 to 47, with a mean value of 41; for males, hematocrit ranges from 42 to 52, with a mean of 47. The percentage of other formed elements, the WBCs and platelets, is extremely small so it is not normally considered with the hematocrit. So the mean plasma percentage is the percent of blood that is not erythrocytes: for females, it is approximately 59 (or 100 minus 41), and for males, it is approximately 53 (or 100 minus 47).

[caption id="" align="aligncenter" width="380"]This figure shows three test tubes with a red and yellow liquid in them. The left panel shows normal blood, the center panel shows anemic blood and the right panel shows polycythemic blood. Figure 1. Composition of Blood. The cellular elements of blood include a vast number of erythrocytes and comparatively fewer leukocytes and platelets. Plasma is the fluid in which the formed elements are suspended. A sample of blood spun in a centrifuge reveals that plasma is the lightest component. It floats at the top of the tube separated from the heaviest elements, the erythrocytes, by a buffy coat of leukocytes and platelets. Hematocrit is the percentage of the total sample that is comprised of erythrocytes. Depressed and elevated hematocrit levels are shown for comparison.[/caption]

Characteristics of Blood

When you think about blood, the first characteristic that probably comes to mind is its color. Blood that has just taken up oxygen in the lungs is bright red, and blood that has released oxygen in the tissues is a more dusky red. This is because hemoglobin is a pigment that changes color, depending upon the degree of oxygen saturation.

Blood is viscous and somewhat sticky to the touch. It has a viscosity approximately five times greater than water. Viscosity is a measure of a fluid’s thickness or resistance to flow, and is influenced by the presence of the plasma proteins and formed elements within the blood. The viscosity of blood has a dramatic impact on blood pressure and flow. Consider the difference in flow between water and honey. The more viscous honey would demonstrate a greater resistance to flow than the less viscous water. The same principle applies to blood.

The normal temperature of blood is slightly higher than normal body temperature—about 38 °C (or 100.4 °F), compared to 37 °C (or 98.6 °F) for an internal body temperature reading, although daily variations of 0.5 °C are normal. Although the surface of blood vessels is relatively smooth, as blood flows through them, it experiences some friction and resistance, especially as vessels age and lose their elasticity, thereby producing heat. This accounts for its slightly higher temperature.

The pH of blood averages about 7.4; however, it can range from 7.35 to 7.45 in a healthy person. Blood is therefore somewhat more basic (alkaline) on a chemical scale than pure water, which has a pH of 7.0. Blood contains numerous buffers that actually help to regulate pH.

Blood constitutes approximately 8 percent of adult body weight. Adult males typically average about 5 to 6 liters of blood. Females average 4–5 liters.

Blood Plasma

Like other fluids in the body, plasma is composed primarily of water: In fact, it is about 92 percent water. Dissolved or suspended within this water is a mixture of substances, most of which are proteins. There are literally hundreds of substances dissolved or suspended in the plasma, although many of them are found only in very small quantities.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit this site for a list of normal levels established for many of the substances found in a sample of blood.[/caption]

Plasma Proteins

About 7 percent of the volume of plasma—nearly all that is not water—is made of proteins. These include several plasma proteins (proteins that are unique to the plasma), plus a much smaller number of regulatory proteins, including enzymes and some hormones. The major components of plasma are summarized in Figure 2.

The three major groups of plasma proteins are as follows:

  • Albumin is the most abundant of the plasma proteins. Manufactured by the liver, albumin molecules serve as binding proteins—transport vehicles for fatty acids and steroid hormones. Recall that lipids are hydrophobic; however, their binding to albumin enables their transport in the watery plasma. Albumin is also the most significant contributor to the osmotic pressure of blood; that is, its presence holds water inside the blood vessels and draws water from the tissues, across blood vessel walls, and into the bloodstream. This in turn helps to maintain both blood volume and blood pressure. Albumin normally accounts for approximately 54 percent of the total plasma protein content, in clinical levels of 3.5–5.0 g/dL blood.
  • The second most common plasma proteins are the globulins. A heterogeneous group, there are three main subgroups known as alpha, beta, and gamma globulins. The alpha and beta globulins transport iron, lipids, and the fat-soluble vitamins A, D, E, and K to the cells; like albumin, they also contribute to osmotic pressure. The gamma globulins are proteins involved in immunity and are better known as an antibodies or immunoglobulins. Although other plasma proteins are produced by the liver, immunoglobulins are produced by specialized leukocytes known as plasma cells. (Seek additional content for more information about immunoglobulins.) Globulins make up approximately 38 percent of the total plasma protein volume, in clinical levels of 1.0–1.5 g/dL blood.
  • The least abundant plasma protein is fibrinogen. Like albumin and the alpha and beta globulins, fibrinogen is produced by the liver. It is essential for blood clotting, a process described later in this chapter. Fibrinogen accounts for about 7 percent of the total plasma protein volume, in clinical levels of 0.2–0.45 g/dL blood.

Other Plasma Solutes

In addition to proteins, plasma contains a wide variety of other substances. These include various electrolytes, such as sodium, potassium, and calcium ions; dissolved gases, such as oxygen, carbon dioxide, and nitrogen; various organic nutrients, such as vitamins, lipids, glucose, and amino acids; and metabolic wastes. All of these nonprotein solutes combined contribute approximately 1 percent to the total volume of plasma.

[caption id="" align="aligncenter" width="600"]This table lists the components of blood, the percentage of each component, their site of production, and their major functions. Figure 2. Major Blood Components[/caption]
[caption id="attachment_2971" align="aligncenter" width="150"] Check out this CrashCourse video to learn more about the components of blood![/caption]
]]> 2200 2194 2 0 19.1 Heart Anatomy https://pressbooks.bccampus.ca/apdouglas2017/chapter/19-1-heart-anatomy/ Fri, 14 Jul 2017 22:57:26 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2237

Learning Objectives

By the end of this section, you will be able to:
  • Describe the location and position of the heart within the body cavity
  • Describe the internal and external anatomy of the heart
  • Identify the tissue layers of the heart
  • Relate the structure of the heart to its function as a pump
  • Compare systemic circulation to pulmonary circulation
  • Identify the veins and arteries of the coronary circulation system
  • Trace the pathway of oxygenated and deoxygenated blood thorough the chambers of the heart

The vital importance of the heart is obvious. If one assumes an average rate of contraction of 75 contractions per minute, a human heart would contract approximately 108,000 times in one day, more than 39 million times in one year, and nearly 3 billion times during a 75-year lifespan. Each of the major pumping chambers of the heart ejects approximately 70 mL blood per contraction in a resting adult. This would be equal to 5.25 liters of fluid per minute and approximately 14,000 liters per day. Over one year, that would equal 10,000,000 liters or 2.6 million gallons of blood sent through roughly 60,000 miles of vessels. In order to understand how that happens, it is necessary to understand the anatomy and physiology of the heart.

Location of the Heart

The human heart is located within the thoracic cavity, medially between the lungs in the space known as the mediastinum. Figure 1 shows the position of the heart within the thoracic cavity. Within the mediastinum, the heart is separated from the other mediastinal structures by a tough membrane known as the pericardium, or pericardial sac, and sits in its own space called the pericardial cavity. The dorsal surface of the heart lies near the bodies of the vertebrae, and its anterior surface sits deep to the sternum and costal cartilages. The great veins, the superior and inferior venae cavae, and the great arteries, the aorta and pulmonary trunk, are attached to the superior surface of the heart, called the base. The base of the heart is located at the level of the third costal cartilage, as seen in Figure 1. The inferior tip of the heart, the apex, lies just to the left of the sternum between the junction of the fourth and fifth ribs near their articulation with the costal cartilages. The right side of the heart is deflected anteriorly, and the left side is deflected posteriorly. It is important to remember the position and orientation of the heart when placing a stethoscope on the chest of a patient and listening for heart sounds, and also when looking at images taken from a midsagittal perspective. The slight deviation of the apex to the left is reflected in a depression in the medial surface of the inferior lobe of the left lung, called the cardiac notch.

[caption id="" align="aligncenter" width="500"]This diagram shows the location of the heart in the thorax. Figure 1. Position of the Heart in the Thorax. The heart is located within the thoracic cavity, medially between the lungs in the mediastinum. It is about the size of a fist, is broad at the top, and tapers toward the base.[/caption]

Shape and Size of the Heart

The shape of the heart is similar to a pinecone, rather broad at the superior surface and tapering to the apex (see Figure 1). A typical heart is approximately the size of your fist: 12 cm (5 in) in length, 8 cm (3.5 in) wide, and 6 cm (2.5 in) in thickness. Given the size difference between most members of the sexes, the weight of a female heart is approximately 250–300 grams (9 to 11 ounces), and the weight of a male heart is approximately 300–350 grams (11 to 12 ounces). The heart of a well-trained athlete, especially one specializing in aerobic sports, can be considerably larger than this. Cardiac muscle responds to exercise in a manner similar to that of skeletal muscle. That is, exercise results in the addition of protein myofilaments that increase the size of the individual cells without increasing their numbers, a concept called hypertrophy. Hearts of athletes can pump blood more effectively at lower rates than those of nonathletes. Enlarged hearts are not always a result of exercise; they can result from pathologies, such as hypertrophic cardiomyopathy. The cause of an abnormally enlarged heart muscle is unknown, but the condition is often undiagnosed and can cause sudden death in apparently otherwise healthy young people.

Chambers and Circulation through the Heart

The human heart consists of four chambers: The left side and the right side each have one atrium and one ventricle. Each of the upper chambers, the right atrium (plural = atria) and the left atrium, acts as a receiving chamber and contracts to push blood into the lower chambers, the right ventricle and the left ventricle. The ventricles serve as the primary pumping chambers of the heart, propelling blood to the lungs or to the rest of the body.

There are two distinct but linked circuits in the human circulation called the pulmonary and systemic circuits. Although both circuits transport blood and everything it carries, we can initially view the circuits from the point of view of gases. The pulmonary circuit transports blood to and from the lungs, where it picks up oxygen and delivers carbon dioxide for exhalation. The systemic circuit transports oxygenated blood to virtually all of the tissues of the body and returns relatively deoxygenated blood and carbon dioxide to the heart to be sent back to the pulmonary circulation.

The right ventricle pumps deoxygenated blood into the pulmonary trunk, which leads toward the lungs and bifurcates into the left and right pulmonary arteries. These vessels in turn branch many times before reaching the pulmonary capillaries, where gas exchange occurs: Carbon dioxide exits the blood and oxygen enters. The pulmonary trunk arteries and their branches are the only arteries in the post-natal body that carry relatively deoxygenated blood. Highly oxygenated blood returning from the pulmonary capillaries in the lungs passes through a series of vessels that join together to form the pulmonary veins—the only post-natal veins in the body that carry highly oxygenated blood. The pulmonary veins conduct blood into the left atrium, which pumps the blood into the left ventricle, which in turn pumps oxygenated blood into the aorta and on to the many branches of the systemic circuit. Eventually, these vessels will lead to the systemic capillaries, where exchange with the tissue fluid and cells of the body occurs. In this case, oxygen and nutrients exit the systemic capillaries to be used by the cells in their metabolic processes, and carbon dioxide and waste products will enter the blood.

The blood exiting the systemic capillaries is lower in oxygen concentration than when it entered. The capillaries will ultimately unite to form venules, joining to form ever-larger veins, eventually flowing into the two major systemic veins, the superior vena cava and the inferior vena cava, which return blood to the right atrium. The blood in the superior and inferior venae cavae flows into the right atrium, which pumps blood into the right ventricle. This process of blood circulation continues as long as the individual remains alive. Understanding the flow of blood through the pulmonary and systemic circuits is critical to all health professions (Figure 3).

[caption id="" align="aligncenter" width="520"]The top panel shows the human heart with the arteries and veins labeled. The bottom panel shows the human circulatory system. Figure 3. Dual System of the Human Blood Circulation. Blood flows from the right atrium to the right ventricle, where it is pumped into the pulmonary circuit. The blood in the pulmonary artery branches is low in oxygen but relatively high in carbon dioxide. Gas exchange occurs in the pulmonary capillaries (oxygen into the blood, carbon dioxide out), and blood high in oxygen and low in carbon dioxide is returned to the left atrium. From here, blood enters the left ventricle, which pumps it into the systemic circuit. Following exchange in the systemic capillaries (oxygen and nutrients out of the capillaries and carbon dioxide and wastes in), blood returns to the right atrium and the cycle is repeated.[/caption]

Membranes, Surface Features, and Layers

Our exploration of more in-depth heart structures begins by examining the membrane that surrounds the heart, the prominent surface features of the heart, and the layers that form the wall of the heart. Each of these components plays its own unique role in terms of function.

Membranes

The membrane that directly surrounds the heart and defines the pericardial cavity is called the pericardium or pericardial sac. It also surrounds the “roots” of the major vessels, or the areas of closest proximity to the heart. The pericardium, which literally translates as “around the heart,” consists of two distinct sublayers: the sturdy outer fibrous pericardium and the inner serous pericardium. The fibrous pericardium is made of tough, dense connective tissue that protects the heart and maintains its position in the thorax. The more delicate serous pericardium consists of two layers: the parietal pericardium, which is fused to the fibrous pericardium, and an inner visceral pericardium, or epicardium, which is fused to the heart and is part of the heart wall. The pericardial cavity, filled with lubricating serous fluid, lies between the epicardium and the pericardium.

In most organs within the body, visceral serous membranes such as the epicardium are microscopic. However, in the case of the heart, it is not a microscopic layer but rather a macroscopic layer, consisting of a simple squamous epithelium called a mesothelium, reinforced with loose, irregular, or areolar connective tissue that attaches to the pericardium. This mesothelium secretes the lubricating serous fluid that fills the pericardial cavity and reduces friction as the heart contracts. Figure 4 illustrates the pericardial membrane and the layers of the heart.

[caption id="" align="aligncenter" width="480"]This image shows a magnified view of the structure of the heart wall. Figure 4. Pericardial Membranes and Layers of the Heart Wall. The pericardial membrane that surrounds the heart consists of three layers and the pericardial cavity. The heart wall also consists of three layers. The pericardial membrane and the heart wall share the epicardium.[/caption]

Surface Features of the Heart

Inside the pericardium, the surface features of the heart are visible, including the four chambers. There is a superficial leaf-like extension of the atria near the superior surface of the heart, one on each side, called an auricle—a name that means “ear like”—because its shape resembles the external ear of a human (Figure 5). Auricles are relatively thin-walled structures that can fill with blood and empty into the atria or upper chambers of the heart. You may also hear them referred to as atrial appendages. Also prominent is a series of fat-filled grooves, each of which is known as a sulcus (plural = sulci), along the superior surfaces of the heart. Major coronary blood vessels are located in these sulci. The deep coronary sulcus is located between the atria and ventricles. Located between the left and right ventricles are two additional sulci that are not as deep as the coronary sulcus. The anterior interventricular sulcus is visible on the anterior surface of the heart, whereas the posterior interventricular sulcus is visible on the posterior surface of the heart. Figure 5 illustrates anterior and posterior views of the surface of the heart.

[caption id="" align="aligncenter" width="530"]The top panel shows the anterior view of the heart and the bottom panel shows the posterior view of the human heart. In both panels, the main parts of the heart are labeled. Figure 5. External Anatomy of the Heart. Inside the pericardium, the surface features of the heart are visible.[/caption]

Layers

The wall of the heart is composed of three layers of unequal thickness. From superficial to deep, these are the epicardium, the myocardium, and the endocardium (see Figure 4). The outermost layer of the wall of the heart is also the innermost layer of the pericardium, the epicardium, or the visceral pericardium discussed earlier.

The middle and thickest layer is the myocardium, made largely of cardiac muscle cells. It is built upon a framework of collagenous fibers, plus the blood vessels that supply the myocardium and the nerve fibers that help regulate the heart. It is the contraction of the myocardium that pumps blood through the heart and into the major arteries. The muscle pattern is elegant and complex, as the muscle cells swirl and spiral around the chambers of the heart. They form a figure 8 pattern around the atria and around the bases of the great vessels. Deeper ventricular muscles also form a figure 8 around the two ventricles and proceed toward the apex. More superficial layers of ventricular muscle wrap around both ventricles. This complex swirling pattern allows the heart to pump blood more effectively than a simple linear pattern would. Figure 6 illustrates the arrangement of muscle cells.

[caption id="" align="aligncenter" width="280"]This diagram shows the muscles in the heart. Figure 6. Heart Musculature. The swirling pattern of cardiac muscle tissue contributes significantly to the heart’s ability to pump blood effectively.[/caption]

Although the ventricles on the right and left sides pump the same amount of blood per contraction, the muscle of the left ventricle is much thicker and better developed than that of the right ventricle. In order to overcome the high resistance required to pump blood into the long systemic circuit, the left ventricle must generate a great amount of pressure. The right ventricle does not need to generate as much pressure, since the pulmonary circuit is shorter and provides less resistance. Figure 7 illustrates the differences in muscular thickness needed for each of the ventricles.

[caption id="" align="aligncenter" width="480"]In this figure the left panel shows the muscles of the heart in the relaxed position, and the right panel shows the muscles of the heart in contracted position. Figure 7. Differences in Ventricular Muscle Thickness. The myocardium in the left ventricle is significantly thicker than that of the right ventricle. Both ventricles pump the same amount of blood, but the left ventricle must generate a much greater pressure to overcome greater resistance in the systemic circuit. The ventricles are shown in both relaxed and contracting states. Note the differences in the relative size of the lumens, the region inside each ventricle where the blood is contained.[/caption]

The innermost layer of the heart wall, the endocardium, is joined to the myocardium with a thin layer of connective tissue. The endocardium lines the chambers where the blood circulates and covers the heart valves. It is made of simple squamous epithelium called endothelium, which is continuous with the endothelial lining of the blood vessels (see Figure 4).

Once regarded as a simple lining layer, recent evidence indicates that the endothelium of the endocardium and the coronary capillaries may play active roles in regulating the contraction of the muscle within the myocardium. The endothelium may also regulate the growth patterns of the cardiac muscle cells throughout life, and the endothelins it secretes create an environment in the surrounding tissue fluids that regulates ionic concentrations and states of contractility. Endothelins are potent vasoconstrictors and, in a normal individual, establish a homeostatic balance with other vasoconstrictors and vasodilators.

Internal Structure of the Heart

Recall that the heart’s contraction cycle follows a dual pattern of circulation—the pulmonary and systemic circuits—because of the pairs of chambers that pump blood into the circulation. In order to develop a more precise understanding of cardiac function, it is first necessary to explore the internal anatomical structures in more detail.

Septa of the Heart

The word septum is derived from the Latin for “something that encloses;” in this case, a septum (plural = septa) refers to a wall or partition that divides the heart into chambers. The septa are physical extensions of the myocardium lined with endocardium. Located between the two atria is the interatrial septum. Normally in an adult heart, the interatrial septum bears an oval-shaped depression known as the fossa ovalis, a remnant of an opening in the fetal heart known as the foramen ovale. The foramen ovale allowed blood in the fetal heart to pass directly from the right atrium to the left atrium, allowing some blood to bypass the pulmonary circuit. Within seconds after birth, a flap of tissue known as the septum primum that previously acted as a valve closes the foramen ovale and establishes the typical cardiac circulation pattern.

Between the two ventricles is a second septum known as the interventricular septum. Unlike the interatrial septum, the interventricular septum is normally intact after its formation during fetal development. It is substantially thicker than the interatrial septum, since the ventricles generate far greater pressure when they contract.

The septum between the atria and ventricles is known as the atrioventricular septum. It is marked by the presence of four openings that allow blood to move from the atria into the ventricles and from the ventricles into the pulmonary trunk and aorta. Located in each of these openings between the atria and ventricles is a valve, a specialized structure that ensures one-way flow of blood. The valves between the atria and ventricles are known generically as atrioventricular valves. The valves at the openings that lead to the pulmonary trunk and aorta are known generically as semilunar valves. The interventricular septum is visible in Figure 8. In this figure, the atrioventricular septum has been removed to better show the bicupid and tricuspid valves; the interatrial septum is not visible, since its location is covered by the aorta and pulmonary trunk. Since these openings and valves structurally weaken the atrioventricular septum, the remaining tissue is heavily reinforced with dense connective tissue called the cardiac skeleton, or skeleton of the heart. It includes four rings that surround the openings between the atria and ventricles, and the openings to the pulmonary trunk and aorta, and serve as the point of attachment for the heart valves. The cardiac skeleton also provides an important boundary in the heart electrical conduction system.

[caption id="" align="aligncenter" width="480"]In this figure the top panel shows the image of the heart with the major parts labeled. The bottom left panel shows a photo of the heart with the surface layer peeled off. The images on the bottom right show detailed musculature inside the heart. Figure 8. Internal Structures of the Heart. This anterior view of the heart shows the four chambers, the major vessels and their early branches, as well as the valves. The presence of the pulmonary trunk and aorta covers the interatrial septum, and the atrioventricular septum is cut away to show the atrioventricular valves.[/caption]

Right Atrium

The right atrium serves as the receiving chamber for blood returning to the heart from the systemic circulation. The two major systemic veins, the superior and inferior venae cavae, and the large coronary vein called the coronary sinus that drains the heart myocardium empty into the right atrium. The superior vena cava drains blood from regions superior to the diaphragm: the head, neck, upper limbs, and the thoracic region. It empties into the superior and posterior portions of the right atrium. The inferior vena cava drains blood from areas inferior to the diaphragm: the lower limbs and abdominopelvic region of the body. It, too, empties into the posterior portion of the atria, but inferior to the opening of the superior vena cava. Immediately superior and slightly medial to the opening of the inferior vena cava on the posterior surface of the atrium is the opening of the coronary sinus. This thin-walled vessel drains most of the coronary veins that return systemic blood from the heart. The majority of the internal heart structures discussed in this and subsequent sections are illustrated in Figure 8.

While the bulk of the internal surface of the right atrium is smooth, the depression of the fossa ovalis is medial, and the anterior surface demonstrates prominent ridges of muscle called the pectinate muscles. The right auricle also has pectinate muscles. The left atrium does not have pectinate muscles except in the auricle.

The atria receive venous blood on a nearly continuous basis, preventing venous flow from stopping while the ventricles are contracting. While most ventricular filling occurs while the atria are relaxed, they do demonstrate a contractile phase and actively pump blood into the ventricles just prior to ventricular contraction. The opening between the atrium and ventricle is guarded by the tricuspid valve.

Right Ventricle

The right ventricle receives blood from the right atrium through the tricuspid valve. Each flap of the valve is attached to strong strands of connective tissue, the chordae tendineae, literally “tendinous cords,” or sometimes more poetically referred to as “heart strings.” There are several chordae tendineae associated with each of the flaps. They are composed of approximately 80 percent collagenous fibers with the remainder consisting of elastic fibers and endothelium. They connect each of the flaps to a papillary muscle that extends from the inferior ventricular surface. There are three papillary muscles in the right ventricle, called the anterior, posterior, and septal muscles, which correspond to the three sections of the valves.

When the myocardium of the ventricle contracts, pressure within the ventricular chamber rises. Blood, like any fluid, flows from higher pressure to lower pressure areas, in this case, toward the pulmonary trunk and the atrium. To prevent any potential backflow, the papillary muscles also contract, generating tension on the chordae tendineae. This prevents the flaps of the valves from being forced into the atria and regurgitation of the blood back into the atria during ventricular contraction. Figure 10 shows papillary muscles and chordae tendineae attached to the tricuspid valve.

[caption id="" align="aligncenter" width="400"]This photo shows the inside of the heart with the main muscles labeled. Figure 10. Chordae Tendineae and Papillary Muscles. In this frontal section, you can see papillary muscles attached to the tricuspid valve on the right as well as the mitral valve on the left via chordae tendineae. (credit: modification of work by “PV KS”/flickr.com)[/caption]

The walls of the ventricle are lined with trabeculae carneae, ridges of cardiac muscle covered by endocardium. In addition to these muscular ridges, a band of cardiac muscle, also covered by endocardium, known as the moderator band (see Figure 8) reinforces the thin walls of the right ventricle and plays a crucial role in cardiac conduction. It arises from the inferior portion of the interventricular septum and crosses the interior space of the right ventricle to connect with the inferior papillary muscle.

When the right ventricle contracts, it ejects blood into the pulmonary trunk, which branches into the left and right pulmonary arteries that carry it to each lung. The superior surface of the right ventricle begins to taper as it approaches the pulmonary trunk. At the base of the pulmonary trunk is the pulmonary semilunar valve that prevents backflow from the pulmonary trunk.

Left Atrium

After exchange of gases in the pulmonary capillaries, blood returns to the left atrium high in oxygen via one of the four pulmonary veins. While the left atrium does not contain pectinate muscles, it does have an auricle that includes these pectinate ridges. Blood flows nearly continuously from the pulmonary veins back into the atrium, which acts as the receiving chamber, and from here through an opening into the left ventricle. Most blood flows passively into the heart while both the atria and ventricles are relaxed, but toward the end of the ventricular relaxation period, the left atrium will contract, pumping blood into the ventricle. This atrial contraction accounts for approximately 20 percent of ventricular filling. The opening between the left atrium and ventricle is guarded by the mitral valve.

Left Ventricle

Recall that, although both sides of the heart will pump the same amount of blood, the muscular layer is much thicker in the left ventricle compared to the right (see Figure 7). Like the right ventricle, the left also has trabeculae carneae, but there is no moderator band. The mitral valve is connected to papillary muscles via chordae tendineae. There are two papillary muscles on the left—the anterior and posterior—as opposed to three on the right.

The left ventricle is the major pumping chamber for the systemic circuit; it ejects blood into the aorta through the aortic semilunar valve.

Heart Valve Structure and Function

A transverse section through the heart slightly above the level of the atrioventricular septum reveals all four heart valves along the same plane (Figure 11). The valves ensure unidirectional blood flow through the heart. Between the right atrium and the right ventricle is the right atrioventricular valve, or tricuspid valve. It typically consists of three flaps, or leaflets, made of endocardium reinforced with additional connective tissue. The flaps are connected by chordae tendineae to the papillary muscles, which control the opening and closing of the valves.

[caption id="" align="aligncenter" width="420"]This diagram shows the anterior view of the heart with the different heart valves labeled. Figure 11. Heart Valves. With the atria and major vessels removed, all four valves are clearly visible, although it is difficult to distinguish the three separate cusps of the tricuspid valve.[/caption]

Emerging from the right ventricle at the base of the pulmonary trunk is the pulmonary semilunar valve, or the pulmonary valve; it is also known as the pulmonic valve or the right semilunar valve. The pulmonary valve is comprised of three small flaps of endothelium reinforced with connective tissue. When the ventricle relaxes, the pressure differential causes blood to flow back into the ventricle from the pulmonary trunk. This flow of blood fills the pocket-like flaps of the pulmonary valve, causing the valve to close and producing an audible sound. Unlike the atrioventricular valves, there are no papillary muscles or chordae tendineae associated with the pulmonary valve.

Located at the opening between the left atrium and left ventricle is the mitral valve, also called the bicuspid valve or the left atrioventricular valve. Structurally, this valve consists of two cusps, known as the anterior medial cusp and the posterior medial cusp, compared to the three cusps of the tricuspid valve. In a clinical setting, the valve is referred to as the mitral valve, rather than the bicuspid valve. The two cusps of the mitral valve are attached by chordae tendineae to two papillary muscles that project from the wall of the ventricle.

At the base of the aorta is the aortic semilunar valve, or the aortic valve, which prevents backflow from the aorta. It normally is composed of three flaps. When the ventricle relaxes and blood attempts to flow back into the ventricle from the aorta, blood will fill the cusps of the valve, causing it to close and producing an audible sound.

In Figure 12a, the two atrioventricular valves are open and the two semilunar valves are closed. This occurs when both atria and ventricles are relaxed and when the atria contract to pump blood into the ventricles. Figure 12b shows a frontal view. Although only the left side of the heart is illustrated, the process is virtually identical on the right.

[caption id="" align="aligncenter" width="480"]The left panel of this figure shows the anterior view of the heart with the different valves, and the right panel of this figure shows the location of the mitral valve in the open position in the heart. Figure 12. Blood Flow from the Left Atrium to the Left Ventricle. (a) A transverse section through the heart illustrates the four heart valves. The two atrioventricular valves are open; the two semilunar valves are closed. The atria and vessels have been removed. (b) A frontal section through the heart illustrates blood flow through the mitral valve. When the mitral valve is open, it allows blood to move from the left atrium to the left ventricle. The aortic semilunar valve is closed to prevent backflow of blood from the aorta to the left ventricle.[/caption]

Figure 13a shows the atrioventricular valves closed while the two semilunar valves are open. This occurs when the ventricles contract to eject blood into the pulmonary trunk and aorta. Closure of the two atrioventricular valves prevents blood from being forced back into the atria. This stage can be seen from a frontal view in Figure 13b.

[caption id="" align="aligncenter" width="480"]The left panel of this figure shows the anterior view of the heart with the different valves, and the right panel of this figure shows the location of the mitral valve in the closed position in the heart. Figure 13. Blood Flow from the Left Ventricle into the Great Vessels. (a) A transverse section through the heart illustrates the four heart valves during ventricular contraction. The two atrioventricular valves are closed, but the two semilunar valves are open. The atria and vessels have been removed. (b) A frontal view shows the closed mitral (bicuspid) valve that prevents backflow of blood into the left atrium. The aortic semilunar valve is open to allow blood to be ejected into the aorta.[/caption]

When the ventricles begin to contract, pressure within the ventricles rises and blood flows toward the area of lowest pressure, which is initially in the atria. This backflow causes the cusps of the tricuspid and mitral (bicuspid) valves to close. These valves are tied down to the papillary muscles by chordae tendineae. During the relaxation phase of the cardiac cycle, the papillary muscles are also relaxed and the tension on the chordae tendineae is slight (see Figure 12b). However, as the myocardium of the ventricle contracts, so do the papillary muscles. This creates tension on the chordae tendineae (see Figure 13b), helping to hold the cusps of the atrioventricular valves in place and preventing them from being blown back into the atria.

The aortic and pulmonary semilunar valves lack the chordae tendineae and papillary muscles associated with the atrioventricular valves. Instead, they consist of pocket-like folds of endocardium reinforced with additional connective tissue. When the ventricles relax and the change in pressure forces the blood toward the ventricles, the blood presses against these cusps and seals the openings.

Coronary Circulation

You will recall that the heart is a remarkable pump composed largely of cardiac muscle cells that are incredibly active throughout life. Like all other cells, a cardiomyocyte requires a reliable supply of oxygen and nutrients, and a way to remove wastes, so it needs a dedicated, complex, and extensive coronary circulation. And because of the critical and nearly ceaseless activity of the heart throughout life, this need for a blood supply is even greater than for a typical cell. However, coronary circulation is not continuous; rather, it cycles, reaching a peak when the heart muscle is relaxed and nearly ceasing while it is contracting.

Coronary Arteries

Coronary arteries supply blood to the myocardium and other components of the heart. The first portion of the aorta after it arises from the left ventricle gives rise to the coronary arteries. There are three dilations in the wall of the aorta just superior to the aortic semilunar valve. Two of these, the left posterior aortic sinus and anterior aortic sinus, give rise to the left and right coronary arteries, respectively. The third sinus, the right posterior aortic sinus, typically does not give rise to a vessel. Coronary vessel branches that remain on the surface of the artery and follow the sulci are called epicardial coronary arteries.

The left coronary artery distributes blood to the left side of the heart, the left atrium and ventricle, and the interventricular septum. The circumflex artery arises from the left coronary artery and follows the coronary sulcus to the left. Eventually, it will fuse with the small branches of the right coronary artery. The larger anterior interventricular artery, also known as the left anterior descending artery (LAD), is the second major branch arising from the left coronary artery. It follows the anterior interventricular sulcus around the pulmonary trunk. Along the way it gives rise to numerous smaller branches that interconnect with the branches of the posterior interventricular artery, forming anastomoses. An anastomosis is an area where vessels unite to form interconnections that normally allow blood to circulate to a region even if there may be partial blockage in another branch. The anastomoses in the heart are very small. Therefore, this ability is somewhat restricted in the heart so a coronary artery blockage often results in death of the cells (myocardial infarction) supplied by the particular vessel.

The right coronary artery proceeds along the coronary sulcus and distributes blood to the right atrium, portions of both ventricles, and the heart conduction system. Normally, one or more marginal arteries arise from the right coronary artery inferior to the right atrium. The marginal arteries supply blood to the superficial portions of the right ventricle. On the posterior surface of the heart, the right coronary artery gives rise to the posterior interventricular artery, also known as the posterior descending artery. It runs along the posterior portion of the interventricular sulcus toward the apex of the heart, giving rise to branches that supply the interventricular septum and portions of both ventricles. Figure 14 presents views of the coronary circulation from both the anterior and posterior views.

[caption id="" align="aligncenter" width="550"]The top panel of this figure shows the anterior view of the heart while the bottom panel shows the posterior view of the heart. The different blood vessels are labeled. Figure 14. Coronary Circulation. The anterior view of the heart shows the prominent coronary surface vessels. The posterior view of the heart shows the prominent coronary surface vessels.[/caption]

Coronary Veins

Coronary veins drain the heart and generally parallel the large surface arteries (see Figure 14). The great cardiac vein can be seen initially on the surface of the heart following the interventricular sulcus, but it eventually flows along the coronary sulcus into the coronary sinus on the posterior surface. The great cardiac vein initially parallels the anterior interventricular artery and drains the areas supplied by this vessel. It receives several major branches, including the posterior cardiac vein, the middle cardiac vein, and the small cardiac vein. The posterior cardiac vein parallels and drains the areas supplied by the marginal artery branch of the circumflex artery. The middle cardiac vein parallels and drains the areas supplied by the posterior interventricular artery. The small cardiac vein parallels the right coronary artery and drains the blood from the posterior surfaces of the right atrium and ventricle. The coronary sinus is a large, thin-walled vein on the posterior surface of the heart lying within the atrioventricular sulcus and emptying directly into the right atrium. The anterior cardiac veins parallel the small cardiac arteries and drain the anterior surface of the right ventricle. Unlike these other cardiac veins, it bypasses the coronary sinus and drains directly into the right atrium.

[caption id="attachment_2973" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of the heart.[/caption]
]]> 2237 2222 2 0 20.5 Circulatory Pathways https://pressbooks.bccampus.ca/apdouglas2017/chapter/20-5-circulatory-pathways/ Fri, 14 Jul 2017 22:59:02 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2309

Learning Objectives

By the end of this section, you will be able to:
  • Identify the vessels through which blood travels within the pulmonary circuit, beginning from the right ventricle of the heart and ending at the left atrium
  • Create a flow chart showing the major systemic arteries through which blood travels from the aorta and its major branches, to the most significant arteries feeding into the right and left upper and lower limbs
  • Create a flow chart showing the major systemic veins through which blood travels from the feet to the right atrium of the heart

Virtually every cell, tissue, organ, and system in the body is impacted by the circulatory system. This includes the generalized and more specialized functions of transport of materials, capillary exchange, maintaining health by transporting white blood cells and various immunoglobulins (antibodies), hemostasis, regulation of body temperature, and helping to maintain acid-base balance. In addition to these shared functions, many systems enjoy a unique relationship with the circulatory system. Figure 1 summarizes these relationships.

[caption id="" align="aligncenter" width="520"]This table outlines the role of the circulatory system in the other organ systems in the body. Figure 1. Interaction of the Circulatory System with Other Body Systems[/caption]

As you learn about the vessels of the systemic and pulmonary circuits, notice that many arteries and veins share the same names, parallel one another throughout the body, and are very similar on the right and left sides of the body. These pairs of vessels will be traced through only one side of the body. Where differences occur in branching patterns or when vessels are singular, this will be indicated. For example, you will find a pair of femoral arteries and a pair of femoral veins, with one vessel on each side of the body. In contrast, some vessels closer to the midline of the body, such as the aorta, are unique. Moreover, some superficial veins, such as the great saphenous vein in the femoral region, have no arterial counterpart. Another phenomenon that can make the study of vessels challenging is that names of vessels can change with location. Like a street that changes name as it passes through an intersection, an artery or vein can change names as it passes an anatomical landmark. For example, the left subclavian artery becomes the axillary artery as it passes through the body wall and into the axillary region, and then becomes the brachial artery as it flows from the axillary region into the upper arm (or brachium). You will also find examples of anastomoses where two blood vessels that previously branched reconnect. Anastomoses are especially common in veins, where they help maintain blood flow even when one vessel is blocked or narrowed, although there are some important ones in the arteries supplying the brain.

As you read about circular pathways, notice that there is an occasional, very large artery referred to as a trunk, a term indicating that the vessel gives rise to several smaller arteries. For example, the celiac trunk gives rise to the left gastric, common hepatic, and splenic arteries.

As you study this section, imagine you are on a “Voyage of Discovery” similar to Lewis and Clark’s expedition in 1804–1806, which followed rivers and streams through unfamiliar territory, seeking a water route from the Atlantic to the Pacific Ocean. You might envision being inside a miniature boat, exploring the various branches of the circulatory system. This simple approach has proven effective for many students in mastering these major circulatory patterns. Another approach that works well for many students is to create simple line drawings similar to the ones provided, labeling each of the major vessels. It is beyond the scope of this text to name every vessel in the body. However, we will attempt to discuss the major pathways for blood and acquaint you with the major named arteries and veins in the body. Also, please keep in mind that individual variations in circulation patterns are not uncommon.

[caption id="attachment_2976" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about the blood vessels.[/caption]

Pulmonary Circulation

Recall that blood returning from the systemic circuit enters the right atrium (Figure 2) via the superior and inferior venae cavae and the coronary sinus, which drains the blood supply of the heart muscle. These vessels will be described more fully later in this section. This blood is relatively low in oxygen and relatively high in carbon dioxide, since much of the oxygen has been extracted for use by the tissues and the waste gas carbon dioxide was picked up to be transported to the lungs for elimination. From the right atrium, blood moves into the right ventricle, which pumps it to the lungs for gas exchange. This system of vessels is referred to as the pulmonary circuit.

The single vessel exiting the right ventricle is the pulmonary trunk. At the base of the pulmonary trunk is the pulmonary semilunar valve, which prevents backflow of blood into the right ventricle during ventricular diastole. As the pulmonary trunk reaches the superior surface of the heart, it curves posteriorly and rapidly bifurcates (divides) into two branches, a left and a right pulmonary artery. To prevent confusion between these vessels, it is important to refer to the vessel exiting the heart as the pulmonary trunk, rather than also calling it a pulmonary artery. The pulmonary arteries in turn branch many times within the lung, forming a series of smaller arteries and arterioles that eventually lead to the pulmonary capillaries. The pulmonary capillaries surround lung structures known as alveoli that are the sites of oxygen and carbon dioxide exchange.

Once gas exchange is completed, oxygenated blood flows from the pulmonary capillaries into a series of pulmonary venules that eventually lead to a series of larger pulmonary veins. Four pulmonary veins, two on the left and two on the right, return blood to the left atrium. At this point, the pulmonary circuit is complete. Table 4 defines the major arteries and veins of the pulmonary circuit discussed in the text.

[caption id="" align="aligncenter" width="420"]This diagram shows the network of blood vessels in the lungs. Figure 2. Pulmonary Circuit. Blood exiting from the right ventricle flows into the pulmonary trunk, which bifurcates into the two pulmonary arteries. These vessels branch to supply blood to the pulmonary capillaries, where gas exchange occurs within the lung alveoli. Blood returns via the pulmonary veins to the left atrium.[/caption]
Pulmonary Arteries and Veins (Table 4)
Vessel Description
Pulmonary trunk Single large vessel exiting the right ventricle that divides to form the right and left pulmonary arteries
Pulmonary arteries Left and right vessels that form from the pulmonary trunk and lead to smaller arterioles and eventually to the pulmonary capillaries
Pulmonary veins Two sets of paired vessels—one pair on each side—that are formed from the small venules, leading away from the pulmonary capillaries to flow into the left atrium

Overview of Systemic Arteries

Blood relatively high in oxygen concentration is returned from the pulmonary circuit to the left atrium via the four pulmonary veins. From the left atrium, blood moves into the left ventricle, which pumps blood into the aorta. The aorta and its branches—the systemic arteries—send blood to virtually every organ of the body (Figure 3).

[caption id="" align="aligncenter" width="400"]This diagrams shows the major arteries in the human body. Figure 3. Systemic Arteries. The major systemic arteries shown here deliver oxygenated blood throughout the body.[/caption]

The Aorta

The aorta is the largest artery in the body (Figure 4). It arises from the left ventricle and eventually descends to the abdominal region, where it bifurcates at the level of the fourth lumbar vertebra into the two common iliac arteries. The aorta consists of the ascending aorta, the aortic arch, and the descending aorta, which passes through the diaphragm and a landmark that divides into the superior thoracic and inferior abdominal components. Arteries originating from the aorta ultimately distribute blood to virtually all tissues of the body. At the base of the aorta is the aortic semilunar valve that prevents backflow of blood into the left ventricle while the heart is relaxing. After exiting the heart, the ascending aorta moves in a superior direction for approximately 5 cm and ends at the sternal angle. Following this ascent, it reverses direction, forming a graceful arc to the left, called the aortic arch. The aortic arch descends toward the inferior portions of the body and ends at the level of the intervertebral disk between the fourth and fifth thoracic vertebrae. Beyond this point, the descending aorta continues close to the bodies of the vertebrae and passes through an opening in the diaphragm known as the aortic hiatus. Superior to the diaphragm, the aorta is called the thoracic aorta, and inferior to the diaphragm, it is called the abdominal aorta. The abdominal aorta terminates when it bifurcates into the two common iliac arteries at the level of the fourth lumbar vertebra. See Figure 4 for an illustration of the ascending aorta, the aortic arch, and the initial segment of the descending aorta plus major branches; Table 5 summarizes the structures of the aorta.

[caption id="" align="aligncenter" width="380"]This diagram shows the aorta and the major parts are labeled. Figure 4. Aorta. The aorta has distinct regions, including the ascending aorta, aortic arch, and the descending aorta, which includes the thoracic and abdominal regions.[/caption]
Components of the Aorta (Table 5)
Vessel Description
Aorta Largest artery in the body, originating from the left ventricle and descending to the abdominal region, where it bifurcates into the common iliac arteries at the level of the fourth lumbar vertebra; arteries originating from the aorta distribute blood to virtually all tissues of the body
Ascending aorta Initial portion of the aorta, rising superiorly from the left ventricle for a distance of approximately 5 cm
Aortic arch Graceful arc to the left that connects the ascending aorta to the descending aorta; ends at the intervertebral disk between the fourth and fifth thoracic vertebrae
Descending aorta Portion of the aorta that continues inferiorly past the end of the aortic arch; subdivided into the thoracic aorta and the abdominal aorta
Thoracic aorta Portion of the descending aorta superior to the aortic hiatus
Abdominal aorta Portion of the aorta inferior to the aortic hiatus and superior to the common iliac arteries

Coronary Circulation

The first vessels that branch from the ascending aorta are the paired coronary arteries (see Figure 4), which arise from two of the three sinuses in the ascending aorta just superior to the aortic semilunar valve. These sinuses contain the aortic baroreceptors and chemoreceptors critical to maintain cardiac function. The left coronary artery arises from the left posterior aortic sinus. The right coronary artery arises from the anterior aortic sinus. Normally, the right posterior aortic sinus does not give rise to a vessel.

The coronary arteries encircle the heart, forming a ring-like structure that divides into the next level of branches that supplies blood to the heart tissues. (Seek additional content for more detail on cardiac circulation.)

Aortic Arch Branches

There are three major branches of the aortic arch: the brachiocephalic artery, the left common carotid artery, and the left subclavian (literally “under the clavicle”) artery. As you would expect based upon proximity to the heart, each of these vessels is classified as an elastic artery.

The brachiocephalic artery is located only on the right side of the body; there is no corresponding artery on the left. The brachiocephalic artery branches into the right subclavian artery and the right common carotid artery. The left subclavian and left common carotid arteries arise independently from the aortic arch but otherwise follow a similar pattern and distribution to the corresponding arteries on the right side (see Figure 2).

Each subclavian artery supplies blood to the arms, chest, shoulders, back, and central nervous system. It then gives rise to three major branches: the internal thoracic artery, the vertebral artery, and the thyrocervical artery. The internal thoracic artery, or mammary artery, supplies blood to the thymus, the pericardium of the heart, and the anterior chest wall. The vertebral artery passes through the vertebral foramen in the cervical vertebrae and then through the foramen magnum into the cranial cavity to supply blood to the brain and spinal cord. The paired vertebral arteries join together to form the large basilar artery at the base of the medulla oblongata. This is an example of an anastomosis. The subclavian artery also gives rise to the thyrocervical artery that provides blood to the thyroid, the cervical region of the neck, and the upper back and shoulder.

The common carotid artery divides into internal and external carotid arteries. The right common carotid artery arises from the brachiocephalic artery and the left common carotid artery arises directly from the aortic arch. The external carotid artery supplies blood to numerous structures within the face, lower jaw, neck, esophagus, and larynx. These branches include the lingual, facial, occipital, maxillary, and superficial temporal arteries. The internal carotid artery initially forms an expansion known as the carotid sinus, containing the carotid baroreceptors and chemoreceptors. Like their counterparts in the aortic sinuses, the information provided by these receptors is critical to maintaining cardiovascular homeostasis (see Figure 2).

The internal carotid arteries along with the vertebral arteries are the two primary suppliers of blood to the human brain. Given the central role and vital importance of the brain to life, it is critical that blood supply to this organ remains uninterrupted. Recall that blood flow to the brain is remarkably constant, with approximately 20 percent of blood flow directed to this organ at any given time. When blood flow is interrupted, even for just a few seconds, a transient ischemic attack (TIA), or mini-stroke, may occur, resulting in loss of consciousness or temporary loss of neurological function. In some cases, the damage may be permanent. Loss of blood flow for longer periods, typically between 3 and 4 minutes, will likely produce irreversible brain damage or a stroke, also called a cerebrovascular accident (CVA). The locations of the arteries in the brain not only provide blood flow to the brain tissue but also prevent interruption in the flow of blood. Both the carotid and vertebral arteries branch once they enter the cranial cavity, and some of these branches form a structure known as the arterial circle (or circle of Willis), an anastomosis that is remarkably like a traffic circle that sends off branches (in this case, arterial branches to the brain). As a rule, branches to the anterior portion of the cerebrum are normally fed by the internal carotid arteries; the remainder of the brain receives blood flow from branches associated with the vertebral arteries.

The internal carotid artery continues through the carotid canal of the temporal bone and enters the base of the brain through the carotid foramen where it gives rise to several branches (Figure 5 and Figure 6). One of these branches is the anterior cerebral artery that supplies blood to the frontal lobe of the cerebrum. Another branch, the middle cerebral artery, supplies blood to the temporal and parietal lobes, which are the most common sites of CVAs. The ophthalmic artery, the third major branch, provides blood to the eyes.

The right and left anterior cerebral arteries join together to form an anastomosis called the anterior communicating artery. The initial segments of the anterior cerebral arteries and the anterior communicating artery form the anterior portion of the arterial circle. The posterior portion of the arterial circle is formed by a left and a right posterior communicating artery that branches from the posterior cerebral artery, which arises from the basilar artery. It provides blood to the posterior portion of the cerebrum and brain stem. The basilar artery is an anastomosis that begins at the junction of the two vertebral arteries and sends branches to the cerebellum and brain stem. It flows into the posterior cerebral arteries. Table 6 summarizes the aortic arch branches, including the major branches supplying the brain.

[caption id="" align="aligncenter" width="420"]This diagram shows the blood vessels in the head and brain. Figure 5. Arteries Supplying the Head and Neck. The common carotid artery gives rise to the external and internal carotid arteries. The external carotid artery remains superficial and gives rise to many arteries of the head. The internal carotid artery first forms the carotid sinus and then reaches the brain via the carotid canal and carotid foramen, emerging into the cranium via the foramen lacerum. The vertebral artery branches from the subclavian artery and passes through the transverse foramen in the cervical vertebrae, entering the base of the skull at the vertebral foramen. The subclavian artery continues toward the arm as the axillary artery.[/caption]
[caption id="" align="aligncenter" width="450"]This diagram shows the arteries of the brain. Figure 6. Arteries Serving the Brain. This inferior view shows the network of arteries serving the brain. The structure is referred to as the arterial circle or circle of Willis.[/caption]
Aortic Arch Branches and Brain Circulation (Table 6)
Vessel Description
Brachiocephalic artery Single vessel located on the right side of the body; the first vessel branching from the aortic arch; gives rise to the right subclavian artery and the right common carotid artery; supplies blood to the head, neck, upper limb, and wall of the thoracic region
Subclavian artery The right subclavian artery arises from the brachiocephalic artery while the left subclavian artery arises from the aortic arch; gives rise to the internal thoracic, vertebral, and thyrocervical arteries; supplies blood to the arms, chest, shoulders, back, and central nervous system
Internal thoracic artery Also called the mammary artery; arises from the subclavian artery; supplies blood to the thymus, pericardium of the heart, and anterior chest wall
Vertebral artery Arises from the subclavian artery and passes through the vertebral foramen through the foramen magnum to the brain; joins with the internal carotid artery to form the arterial circle; supplies blood to the brain and spinal cord
Thyrocervical artery Arises from the subclavian artery; supplies blood to the thyroid, the cervical region, the upper back, and shoulder
Common carotid artery The right common carotid artery arises from the brachiocephalic artery and the left common carotid artery arises from the aortic arch; each gives rise to the external and internal carotid arteries; supplies the respective sides of the head and neck
External carotid artery Arises from the common carotid artery; supplies blood to numerous structures within the face, lower jaw, neck, esophagus, and larynx
Internal carotid artery Arises from the common carotid artery and begins with the carotid sinus; goes through the carotid canal of the temporal bone to the base of the brain; combines with the branches of the vertebral artery, forming the arterial circle; supplies blood to the brain
Arterial circle or circle of Willis An anastomosis located at the base of the brain that ensures continual blood supply; formed from the branches of the internal carotid and vertebral arteries; supplies blood to the brain
Anterior cerebral artery Arises from the internal carotid artery; supplies blood to the frontal lobe of the cerebrum
Middle cerebral artery Another branch of the internal carotid artery; supplies blood to the temporal and parietal lobes of the cerebrum
Ophthalmic artery Branch of the internal carotid artery; supplies blood to the eyes
Anterior communicating artery An anastomosis of the right and left internal carotid arteries; supplies blood to the brain
Posterior communicating artery Branches of the posterior cerebral artery that form part of the posterior portion of the arterial circle; supplies blood to the brain
Posterior cerebral artery Branch of the basilar artery that forms a portion of the posterior segment of the arterial circle of Willis; supplies blood to the posterior portion of the cerebrum and brain stem
Basilar artery Formed from the fusion of the two vertebral arteries; sends branches to the cerebellum, brain stem, and the posterior cerebral arteries; the main blood supply to the brain stem

Thoracic Aorta and Major Branches

The thoracic aorta begins at the level of vertebra T5 and continues through to the diaphragm at the level of T12, initially traveling within the mediastinum to the left of the vertebral column. As it passes through the thoracic region, the thoracic aorta gives rise to several branches, which are collectively referred to as visceral branches and parietal branches (Figure 7). Those branches that supply blood primarily to visceral organs are known as the visceral branches and include the bronchial arteries, pericardial arteries, esophageal arteries, and the mediastinal arteries, each named after the tissues it supplies. Each bronchial artery (typically two on the left and one on the right) supplies systemic blood to the lungs and visceral pleura, in addition to the blood pumped to the lungs for oxygenation via the pulmonary circuit. The bronchial arteries follow the same path as the respiratory branches, beginning with the bronchi and ending with the bronchioles. There is considerable, but not total, intermingling of the systemic and pulmonary blood at anastomoses in the smaller branches of the lungs. This may sound incongruous—that is, the mixing of systemic arterial blood high in oxygen with the pulmonary arterial blood lower in oxygen—but the systemic vessels also deliver nutrients to the lung tissue just as they do elsewhere in the body. The mixed blood drains into typical pulmonary veins, whereas the bronchial artery branches remain separate and drain into bronchial veins described later. Each pericardial artery supplies blood to the pericardium, the esophageal artery provides blood to the esophagus, and the mediastinal artery provides blood to the mediastinum. The remaining thoracic aorta branches are collectively referred to as parietal branches or somatic branches, and include the intercostal and superior phrenic arteries. Each intercostal artery provides blood to the muscles of the thoracic cavity and vertebral column. The superior phrenic artery provides blood to the superior surface of the diaphragm. Table 7 lists the arteries of the thoracic region.

[caption id="" align="aligncenter" width="480"]This diagram shows the arteries in the thoracic and abdominal cavity. Figure 7. Arteries of the Thoracic and Abdominal Regions. The thoracic aorta gives rise to the arteries of the visceral and parietal branches.[/caption]
Arteries of the Thoracic Region (Table 7)
Vessel Description
Visceral branches A group of arterial branches of the thoracic aorta; supplies blood to the viscera (i.e., organs) of the thorax
Bronchial artery Systemic branch from the aorta that provides oxygenated blood to the lungs; this blood supply is in addition to the pulmonary circuit that brings blood for oxygenation
Pericardial artery Branch of the thoracic aorta; supplies blood to the pericardium
Esophageal artery Branch of the thoracic aorta; supplies blood to the esophagus
Mediastinal artery Branch of the thoracic aorta; supplies blood to the mediastinum
Parietal branches Also called somatic branches, a group of arterial branches of the thoracic aorta; include those that supply blood to the thoracic wall, vertebral column, and the superior surface of the diaphragm
Intercostal artery Branch of the thoracic aorta; supplies blood to the muscles of the thoracic cavity and vertebral column
Superior phrenic artery Branch of the thoracic aorta; supplies blood to the superior surface of the diaphragm

Abdominal Aorta and Major Branches

After crossing through the diaphragm at the aortic hiatus, the thoracic aorta is called the abdominal aorta (see Figure 7). This vessel remains to the left of the vertebral column and is embedded in adipose tissue behind the peritoneal cavity. It formally ends at approximately the level of vertebra L4, where it bifurcates to form the common iliac arteries. Before this division, the abdominal aorta gives rise to several important branches. A single celiac trunk (artery) emerges and divides into the left gastric artery to supply blood to the stomach and esophagus, the splenic artery to supply blood to the spleen, and the common hepatic artery, which in turn gives rise to the hepatic artery proper to supply blood to the liver, the right gastric artery to supply blood to the stomach, the cystic artery to supply blood to the gall bladder, and several branches, one to supply blood to the duodenum and another to supply blood to the pancreas. Two additional single vessels arise from the abdominal aorta. These are the superior and inferior mesenteric arteries. The superior mesenteric artery arises approximately 2.5 cm after the celiac trunk and branches into several major vessels that supply blood to the small intestine (duodenum, jejunum, and ileum), the pancreas, and a majority of the large intestine. The inferior mesenteric artery supplies blood to the distal segment of the large intestine, including the rectum. It arises approximately 5 cm superior to the common iliac arteries.

In addition to these single branches, the abdominal aorta gives rise to several significant paired arteries along the way. These include the inferior phrenic arteries, the adrenal arteries, the renal arteries, the gonadal arteries, and the lumbar arteries. Each inferior phrenic artery is a counterpart of a superior phrenic artery and supplies blood to the inferior surface of the diaphragm. The adrenal artery supplies blood to the adrenal (suprarenal) glands and arises near the superior mesenteric artery. Each renal artery branches approximately 2.5 cm inferior to the superior mesenteric arteries and supplies a kidney. The right renal artery is longer than the left since the aorta lies to the left of the vertebral column and the vessel must travel a greater distance to reach its target. Renal arteries branch repeatedly to supply blood to the kidneys. Each gonadal artery supplies blood to the gonads, or reproductive organs, and is also described as either an ovarian artery or a testicular artery (internal spermatic), depending upon the sex of the individual. An ovarian artery supplies blood to an ovary, uterine (Fallopian) tube, and the uterus, and is located within the suspensory ligament of the uterus. It is considerably shorter than a testicular artery, which ultimately travels outside the body cavity to the testes, forming one component of the spermatic cord. The gonadal arteries arise inferior to the renal arteries and are generally retroperitoneal. The ovarian artery continues to the uterus where it forms an anastomosis with the uterine artery that supplies blood to the uterus. Both the uterine arteries and vaginal arteries, which distribute blood to the vagina, are branches of the internal iliac artery. The four paired lumbar arteries are the counterparts of the intercostal arteries and supply blood to the lumbar region, the abdominal wall, and the spinal cord. In some instances, a fifth pair of lumbar arteries emerges from the median sacral artery.

The aorta divides at approximately the level of vertebra L4 into a left and a right common iliac artery but continues as a small vessel, the median sacral artery, into the sacrum. The common iliac arteries provide blood to the pelvic region and ultimately to the lower limbs. They split into external and internal iliac arteries approximately at the level of the lumbar-sacral articulation. Each internal iliac artery sends branches to the urinary bladder, the walls of the pelvis, the external genitalia, and the medial portion of the femoral region. In females, they also provide blood to the uterus and vagina. The much larger external iliac artery supplies blood to each of the lower limbs. Figure 8 shows the distribution of the major branches of the aorta into the thoracic and abdominal regions. Figure 9 shows the distribution of the major branches of the common iliac arteries. Table 8 summarizes the major branches of the abdominal aorta.

[caption id="" align="aligncenter" width="480"]This table shows the different arteries in the thoracic and abdominal cavity. The list on the left shows unpaired arteries, and the list on the right shows paired cavities. Figure 8. Major Branches of the Aorta. The flow chart summarizes the distribution of the major branches of the aorta into the thoracic and abdominal regions.[/caption]
Major Branches of the Iliac Arteries
The flow chart summarizes the distribution of the major branches of the common iliac arteries into the pelvis and lower limbs. The left side follows a similar pattern to the right.
[caption id="" align="aligncenter" width="480"]This flowchart shows the different branches into which that the abdominal aorta is divided. Figure 9. Major Branches of the Iliac Arteries. The flow chart summarizes the distribution of the major branches of the common iliac arteries into the pelvis and lower limbs. The left side follows a similar pattern to the right.[/caption]
Vessels of the Abdominal Aorta
Vessel Description
Celiac trunk Also called the celiac artery; a major branch of the abdominal aorta; gives rise to the left gastric artery, the splenic artery, and the common hepatic artery that forms the hepatic artery to the liver, the right gastric artery to the stomach, and the cystic artery to the gall bladder
Left gastric artery Branch of the celiac trunk; supplies blood to the stomach
Splenic artery Branch of the celiac trunk; supplies blood to the spleen
Common hepatic artery Branch of the celiac trunk that forms the hepatic artery, the right gastric artery, and the cystic artery
Hepatic artery proper Branch of the common hepatic artery; supplies systemic blood to the liver
Right gastric artery Branch of the common hepatic artery; supplies blood to the stomach
Cystic artery Branch of the common hepatic artery; supplies blood to the gall bladder
Superior mesenteric artery Branch of the abdominal aorta; supplies blood to the small intestine (duodenum, jejunum, and ileum), the pancreas, and a majority of the large intestine
Inferior mesenteric artery Branch of the abdominal aorta; supplies blood to the distal segment of the large intestine and rectum
Inferior phrenic arteries Branches of the abdominal aorta; supply blood to the inferior surface of the diaphragm
Adrenal artery Branch of the abdominal aorta; supplies blood to the adrenal (suprarenal) glands
Renal artery Branch of the abdominal aorta; supplies each kidney
Gonadal artery Branch of the abdominal aorta; supplies blood to the gonads or reproductive organs; also described as ovarian arteries or testicular arteries, depending upon the sex of the individual
Ovarian artery Branch of the abdominal aorta; supplies blood to ovary, uterine (Fallopian) tube, and uterus
Testicular artery Branch of the abdominal aorta; ultimately travels outside the body cavity to the testes and forms one component of the spermatic cord
Lumbar arteries Branches of the abdominal aorta; supply blood to the lumbar region, the abdominal wall, and spinal cord
Common iliac artery Branch of the aorta that leads to the internal and external iliac arteries
Median sacral artery Continuation of the aorta into the sacrum
Internal iliac artery Branch from the common iliac arteries; supplies blood to the urinary bladder, walls of the pelvis, external genitalia, and the medial portion of the femoral region; in females, also provides blood to the uterus and vagina
External iliac artery Branch of the common iliac artery that leaves the body cavity and becomes a femoral artery; supplies blood to the lower limbs

Arteries Serving the Upper Limbs

As the subclavian artery exits the thorax into the axillary region, it is renamed the axillary artery. Although it does branch and supply blood to the region near the head of the humerus (via the humeral circumflex arteries), the majority of the vessel continues into the upper arm, or brachium, and becomes the brachial artery (Figure 10). The brachial artery supplies blood to much of the brachial region and divides at the elbow into several smaller branches, including the deep brachial arteries, which provide blood to the posterior surface of the arm, and the ulnar collateral arteries, which supply blood to the region of the elbow. As the brachial artery approaches the coronoid fossa, it bifurcates into the radial and ulnar arteries, which continue into the forearm, or antebrachium. The radial artery and ulnar artery parallel their namesake bones, giving off smaller branches until they reach the wrist, or carpal region. At this level, they fuse to form the superficial and deep palmar arches that supply blood to the hand, as well as the digital arteries that supply blood to the digits. Figure 11 shows the distribution of systemic arteries from the heart into the upper limb. Table 9 summarizes the arteries serving the upper limbs.

[caption id="" align="aligncenter" width="280"]This diagram shows the arteries in the arm. Figure 10. Major Arteries Serving the Thorax and Upper Limb. The arteries that supply blood to the arms and hands are extensions of the subclavian arteries.[/caption]
[caption id="" align="aligncenter" width="520"]This chart shows the arteries present in the thoracic upper limb. Figure 11. Major Arteries of the Upper Limb. The flow chart summarizes the distribution of the major arteries from the heart into the upper limb.[/caption]
Arteries Serving the Upper Limbs (Table 9)
Vessel Description
Axillary artery Continuation of the subclavian artery as it penetrates the body wall and enters the axillary region; supplies blood to the region near the head of the humerus (humeral circumflex arteries); the majority of the vessel continues into the brachium and becomes the brachial artery
Brachial artery Continuation of the axillary artery in the brachium; supplies blood to much of the brachial region; gives off several smaller branches that provide blood to the posterior surface of the arm in the region of the elbow; bifurcates into the radial and ulnar arteries at the coronoid fossa
Radial artery Formed at the bifurcation of the brachial artery; parallels the radius; gives off smaller branches until it reaches the carpal region where it fuses with the ulnar artery to form the superficial and deep palmar arches; supplies blood to the lower arm and carpal region
Ulnar artery Formed at the bifurcation of the brachial artery; parallels the ulna; gives off smaller branches until it reaches the carpal region where it fuses with the radial artery to form the superficial and deep palmar arches; supplies blood to the lower arm and carpal region
Palmar arches (superficial and deep) Formed from anastomosis of the radial and ulnar arteries; supply blood to the hand and digital arteries
Digital arteries Formed from the superficial and deep palmar arches; supply blood to the digits

Arteries Serving the Lower Limbs

The external iliac artery exits the body cavity and enters the femoral region of the lower leg (Figure 12). As it passes through the body wall, it is renamed the femoral artery. It gives off several smaller branches as well as the lateral deep femoral artery that in turn gives rise to a lateral circumflex artery. These arteries supply blood to the deep muscles of the thigh as well as ventral and lateral regions of the integument. The femoral artery also gives rise to the genicular artery, which provides blood to the region of the knee. As the femoral artery passes posterior to the knee near the popliteal fossa, it is called the popliteal artery. The popliteal artery branches into the anterior and posterior tibial arteries.

The anterior tibial artery is located between the tibia and fibula, and supplies blood to the muscles and integument of the anterior tibial region. Upon reaching the tarsal region, it becomes the dorsalis pedis artery, which branches repeatedly and provides blood to the tarsal and dorsal regions of the foot. The posterior tibial artery provides blood to the muscles and integument on the posterior surface of the tibial region. The fibular or peroneal artery branches from the posterior tibial artery. It bifurcates and becomes the medial plantar artery and lateral plantar artery, providing blood to the plantar surfaces. There is an anastomosis with the dorsalis pedis artery, and the medial and lateral plantar arteries form two arches called the dorsal arch (also called the arcuate arch) and the plantar arch, which provide blood to the remainder of the foot and toes. Figure 13 shows the distribution of the major systemic arteries in the lower limb. Table 10 summarizes the major systemic arteries discussed in the text.

[caption id="" align="aligncenter" width="500"]The left panel shows the anterior view of arteries in the legs, and the right panel shows the posterior view. Figure 12. Major Arteries Serving the Lower Limb. Major arteries serving the lower limb are shown in anterior and posterior views.[/caption]
[caption id="" align="aligncenter" width="520"]This chart shows the major arteries present in the lower limbs. Figure 13. Systemic Arteries of the Lower Limb. The flow chart summarizes the distribution of the systemic arteries from the external iliac artery into the lower limb.[/caption]
Arteries Serving the Lower Limbs (Table 10)
Vessel Description
Femoral artery Continuation of the external iliac artery after it passes through the body cavity; divides into several smaller branches, the lateral deep femoral artery, and the genicular artery; becomes the popliteal artery as it passes posterior to the knee
Deep femoral artery Branch of the femoral artery; gives rise to the lateral circumflex arteries
Lateral circumflex artery Branch of the deep femoral artery; supplies blood to the deep muscles of the thigh and the ventral and lateral regions of the integument
Genicular artery Branch of the femoral artery; supplies blood to the region of the knee
Popliteal artery Continuation of the femoral artery posterior to the knee; branches into the anterior and posterior tibial arteries
Anterior tibial artery Branches from the popliteal artery; supplies blood to the anterior tibial region; becomes the dorsalis pedis artery
Dorsalis pedis artery Forms from the anterior tibial artery; branches repeatedly to supply blood to the tarsal and dorsal regions of the foot
Posterior tibial artery Branches from the popliteal artery and gives rise to the fibular or peroneal artery; supplies blood to the posterior tibial region
Medial plantar artery Arises from the bifurcation of the posterior tibial arteries; supplies blood to the medial plantar surfaces of the foot
Lateral plantar artery Arises from the bifurcation of the posterior tibial arteries; supplies blood to the lateral plantar surfaces of the foot
Dorsal or arcuate arch Formed from the anastomosis of the dorsalis pedis artery and the medial and plantar arteries; branches supply the distal portions of the foot and digits
Plantar arch Formed from the anastomosis of the dorsalis pedis artery and the medial and plantar arteries; branches supply the distal portions of the foot and digits

Overview of Systemic Veins

Systemic veins return blood to the right atrium. Since the blood has already passed through the systemic capillaries, it will be relatively low in oxygen concentration. In many cases, there will be veins draining organs and regions of the body with the same name as the arteries that supplied these regions and the two often parallel one another. This is often described as a “complementary” pattern. However, there is a great deal more variability in the venous circulation than normally occurs in the arteries. For the sake of brevity and clarity, this text will discuss only the most commonly encountered patterns. However, keep this variation in mind when you move from the classroom to clinical practice.

In both the neck and limb regions, there are often both superficial and deeper levels of veins. The deeper veins generally correspond to the complementary arteries. The superficial veins do not normally have direct arterial counterparts, but in addition to returning blood, they also make contributions to the maintenance of body temperature. When the ambient temperature is warm, more blood is diverted to the superficial veins where heat can be more easily dissipated to the environment. In colder weather, there is more constriction of the superficial veins and blood is diverted deeper where the body can retain more of the heat.

The “Voyage of Discovery” analogy and stick drawings mentioned earlier remain valid techniques for the study of systemic veins, but veins present a more difficult challenge because there are numerous anastomoses and multiple branches. It is like following a river with many tributaries and channels, several of which interconnect. Tracing blood flow through arteries follows the current in the direction of blood flow, so that we move from the heart through the large arteries and into the smaller arteries to the capillaries. From the capillaries, we move into the smallest veins and follow the direction of blood flow into larger veins and back to the heart. Figure 14 outlines the path of the major systemic veins.
[caption id="" align="aligncenter" width="460"]This diagram shows the major veins in the human body. Figure 14. Major Systemic Veins of the Body. The major systemic veins of the body are shown here in an anterior view.[/caption]

The right atrium receives all of the systemic venous return. Most of the blood flows into either the superior vena cava or inferior vena cava. If you draw an imaginary line at the level of the diaphragm, systemic venous circulation from above that line will generally flow into the superior vena cava; this includes blood from the head, neck, chest, shoulders, and upper limbs. The exception to this is that most venous blood flow from the coronary veins flows directly into the coronary sinus and from there directly into the right atrium. Beneath the diaphragm, systemic venous flow enters the inferior vena cava, that is, blood from the abdominal and pelvic regions and the lower limbs.

The Superior Vena Cava

The superior vena cava drains most of the body superior to the diaphragm (Figure 15). On both the left and right sides, the subclavian vein forms when the axillary vein passes through the body wall from the axillary region. It fuses with the external and internal jugular veins from the head and neck to form the brachiocephalic vein. Each vertebral vein also flows into the brachiocephalic vein close to this fusion. These veins arise from the base of the brain and the cervical region of the spinal cord, and flow largely through the intervertebral foramina in the cervical vertebrae. They are the counterparts of the vertebral arteries. Each internal thoracic vein, also known as an internal mammary vein, drains the anterior surface of the chest wall and flows into the brachiocephalic vein.

The remainder of the blood supply from the thorax drains into the azygos vein. Each intercostal vein drains muscles of the thoracic wall, each esophageal vein delivers blood from the inferior portions of the esophagus, each bronchial vein drains the systemic circulation from the lungs, and several smaller veins drain the mediastinal region. Bronchial veins carry approximately 13 percent of the blood that flows into the bronchial arteries; the remainder intermingles with the pulmonary circulation and returns to the heart via the pulmonary veins. These veins flow into the azygos vein, and with the smaller hemiazygos vein (hemi- = “half”) on the left of the vertebral column, drain blood from the thoracic region. The hemiazygos vein does not drain directly into the superior vena cava but enters the brachiocephalic vein via the superior intercostal vein.

The azygos vein passes through the diaphragm from the thoracic cavity on the right side of the vertebral column and begins in the lumbar region of the thoracic cavity. It flows into the superior vena cava at approximately the level of T2, making a significant contribution to the flow of blood. It combines with the two large left and right brachiocephalic veins to form the superior vena cava.

Table 11 summarizes the veins of the thoracic region that flow into the superior vena cava.

[caption id="" align="aligncenter" width="480"]This diagram shows the veins present in the thoracic abdominal cavity. Figure 15. Veins of the Thoracic and Abdominal Regions. Veins of the thoracic and abdominal regions drain blood from the area above the diaphragm, returning it to the right atrium via the superior vena cava.[/caption]
Veins of the Thoracic Region (Table 11)
Vessel Description
Superior vena cava Large systemic vein; drains blood from most areas superior to the diaphragm; empties into the right atrium
Subclavian vein Located deep in the thoracic cavity; formed by the axillary vein as it enters the thoracic cavity from the axillary region; drains the axillary and smaller local veins near the scapular region and leads to the brachiocephalic vein
Brachiocephalic veins Pair of veins that form from a fusion of the external and internal jugular veins and the subclavian vein; subclavian, external and internal jugulars, vertebral, and internal thoracic veins flow into it; drain the upper thoracic region and lead to the superior vena cava
Vertebral vein Arises from the base of the brain and the cervical region of the spinal cord; passes through the intervertebral foramina in the cervical vertebrae; drains smaller veins from the cranium, spinal cord, and vertebrae, and leads to the brachiocephalic vein; counterpart of the vertebral artery
Internal thoracic veins Also called internal mammary veins; drain the anterior surface of the chest wall and lead to the brachiocephalic vein
Intercostal vein Drains the muscles of the thoracic wall and leads to the azygos vein
Esophageal vein Drains the inferior portions of the esophagus and leads to the azygos vein
Bronchial vein Drains the systemic circulation from the lungs and leads to the azygos vein
Azygos vein Originates in the lumbar region and passes through the diaphragm into the thoracic cavity on the right side of the vertebral column; drains blood from the intercostal veins, esophageal veins, bronchial veins, and other veins draining the mediastinal region, and leads to the superior vena cava
Hemiazygos vein Smaller vein complementary to the azygos vein; drains the esophageal veins from the esophagus and the left intercostal veins, and leads to the brachiocephalic vein via the superior intercostal vein

Veins of the Head and Neck

Blood from the brain and the superficial facial vein flow into each internal jugular vein (Figure 16). Blood from the more superficial portions of the head, scalp, and cranial regions, including the temporal vein and maxillary vein, flow into each external jugular vein. Although the external and internal jugular veins are separate vessels, there are anastomoses between them close to the thoracic region. Blood from the external jugular vein empties into the subclavian vein. Table 12 summarizes the major veins of the head and neck.

Major Veins of the Head and Neck (Table 12)
Vessel Description
Internal jugular vein Parallel to the common carotid artery, which is more or less its counterpart, and passes through the jugular foramen and canal; primarily drains blood from the brain, receives the superficial facial vein, and empties into the subclavian vein
Temporal vein Drains blood from the temporal region and flows into the external jugular vein
Maxillary vein Drains blood from the maxillary region and flows into the external jugular vein
External jugular vein Drains blood from the more superficial portions of the head, scalp, and cranial regions, and leads to the subclavian vein

Venous Drainage of the Brain

Circulation to the brain is both critical and complex (see Table 16). Many smaller veins of the brain stem and the superficial veins of the cerebrum lead to larger vessels referred to as intracranial sinuses. These include the superior and inferior sagittal sinuses, straight sinus, cavernous sinuses, left and right sinuses, the petrosal sinuses, and the occipital sinuses. Ultimately, sinuses will lead back to either the inferior jugular vein or vertebral vein.

Most of the veins on the superior surface of the cerebrum flow into the largest of the sinuses, the superior sagittal sinus. It is located midsagittally between the meningeal and periosteal layers of the dura mater within the falx cerebri and, at first glance in images or models, can be mistaken for the subarachnoid space. Most reabsorption of cerebrospinal fluid occurs via the chorionic villi (arachnoid granulations) into the superior sagittal sinus. Blood from most of the smaller vessels originating from the inferior cerebral veins flows into the great cerebral vein and into the straight sinus. Other cerebral veins and those from the eye socket flow into the cavernous sinus, which flows into the petrosal sinus and then into the internal jugular vein. The occipital sinus, sagittal sinus, and straight sinuses all flow into the left and right transverse sinuses near the lambdoid suture. The transverse sinuses in turn flow into the sigmoid sinuses that pass through the jugular foramen and into the internal jugular vein. The internal jugular vein flows parallel to the common carotid artery and is more or less its counterpart. It empties into the brachiocephalic vein. The veins draining the cervical vertebrae and the posterior surface of the skull, including some blood from the occipital sinus, flow into the vertebral veins. These parallel the vertebral arteries and travel through the transverse foramina of the cervical vertebrae. The vertebral veins also flow into the brachiocephalic veins. Table 13 summarizes the major veins of the brain.

[caption id="" align="aligncenter" width="460"]This diagram shows the veins present in the head and neck. Figure 16. Veins of the Head and Neck. This left lateral view shows the veins of the head and neck, including the intercranial sinuses.[/caption]
Major Veins of the Brain (Table 13)
Vessel Description
Superior sagittal sinus Enlarged vein located midsagittally between the meningeal and periosteal layers of the dura mater within the falx cerebri; receives most of the blood drained from the superior surface of the cerebrum and leads to the inferior jugular vein and the vertebral vein
Great cerebral vein Receives most of the smaller vessels from the inferior cerebral veins and leads to the straight sinus
Straight sinus Enlarged vein that drains blood from the brain; receives most of the blood from the great cerebral vein and leads to the left or right transverse sinus
Cavernous sinus Enlarged vein that receives blood from most of the other cerebral veins and the eye socket, and leads to the petrosal sinus
Petrosal sinus Enlarged vein that receives blood from the cavernous sinus and leads into the internal jugular veins
Occipital sinus Enlarged vein that drains the occipital region near the falx cerebelli and leads to the left and right transverse sinuses, and also the vertebral veins
Transverse sinuses Pair of enlarged veins near the lambdoid suture that drains the occipital, sagittal, and straight sinuses, and leads to the sigmoid sinuses
Sigmoid sinuses Enlarged vein that receives blood from the transverse sinuses and leads through the jugular foramen to the internal jugular vein

Veins Draining the Upper Limbs

The digital veins in the fingers come together in the hand to form the palmar venous arches (Figure 17). From here, the veins come together to form the radial vein, the ulnar vein, and the median antebrachial vein. The radial vein and the ulnar vein parallel the bones of the forearm and join together at the antebrachium to form the brachial vein, a deep vein that flows into the axillary vein in the brachium.

The median antebrachial vein parallels the ulnar vein, is more medial in location, and joins the basilic vein in the forearm. As the basilic vein reaches the antecubital region, it gives off a branch called the median cubital vein that crosses at an angle to join the cephalic vein. The median cubital vein is the most common site for drawing venous blood in humans. The basilic vein continues through the arm medially and superficially to the axillary vein.

The cephalic vein begins in the antebrachium and drains blood from the superficial surface of the arm into the axillary vein. It is extremely superficial and easily seen along the surface of the biceps brachii muscle in individuals with good muscle tone and in those without excessive subcutaneous adipose tissue in the arms.

The subscapular vein drains blood from the subscapular region and joins the cephalic vein to form the axillary vein. As it passes through the body wall and enters the thorax, the axillary vein becomes the subclavian vein.

Many of the larger veins of the thoracic and abdominal region and upper limb are further represented in the flow chart in Figure 18. Table 14 summarizes the veins of the upper limbs.

[caption id="" align="aligncenter" width="385"]This diagram shows the veins present in the upper limb. Figure 17. Veins of the Upper Limb. This anterior view shows the veins that drain the upper limb.[/caption]
[caption id="" align="aligncenter" width="530"]This flowchart shows the different veins in the body, and how they are connected to the superior vena cava. Figure 18. Veins Flowing into the Superior Vena Cava. The flow chart summarizes the distribution of the veins flowing into the superior vena cava.[/caption]
Veins of the Upper Limbs (Table 14)
Vessel Description
Digital veins Drain the digits and lead to the palmar arches of the hand and dorsal venous arch of the foot
Palmar venous arches Drain the hand and digits, and lead to the radial vein, ulnar veins, and the median antebrachial vein
Radial vein Vein that parallels the radius and radial artery; arises from the palmar venous arches and leads to the brachial vein
Ulnar vein Vein that parallels the ulna and ulnar artery; arises from the palmar venous arches and leads to the brachial vein
Brachial vein Deeper vein of the arm that forms from the radial and ulnar veins in the lower arm; leads to the axillary vein
Median antebrachial vein Vein that parallels the ulnar vein but is more medial in location; intertwines with the palmar venous arches; leads to the basilic vein
Basilic vein Superficial vein of the arm that arises from the median antebrachial vein, intersects with the median cubital vein, parallels the ulnar vein, and continues into the upper arm; along with the brachial vein, it leads to the axillary vein
Median cubital vein Superficial vessel located in the antecubital region that links the cephalic vein to the basilic vein in the form of a v; a frequent site from which to draw blood
Cephalic vein Superficial vessel in the upper arm; leads to the axillary vein
Subscapular vein Drains blood from the subscapular region and leads to the axillary vein
Axillary vein The major vein in the axillary region; drains the upper limb and becomes the subclavian vein

The Inferior Vena Cava

Other than the small amount of blood drained by the azygos and hemiazygos veins, most of the blood inferior to the diaphragm drains into the inferior vena cava before it is returned to the heart (see Figure 15). Lying just beneath the parietal peritoneum in the abdominal cavity, the inferior vena cava parallels the abdominal aorta, where it can receive blood from abdominal veins. The lumbar portions of the abdominal wall and spinal cord are drained by a series of lumbar veins, usually four on each side. The ascending lumbar veins drain into either the azygos vein on the right or the hemiazygos vein on the left, and return to the superior vena cava. The remaining lumbar veins drain directly into the inferior vena cava.

Blood supply from the kidneys flows into each renal vein, normally the largest veins entering the inferior vena cava. A number of other, smaller veins empty into the left renal vein. Each adrenal vein drains the adrenal or suprarenal glands located immediately superior to the kidneys. The right adrenal vein enters the inferior vena cava directly, whereas the left adrenal vein enters the left renal vein.

From the male reproductive organs, each testicular vein flows from the scrotum, forming a portion of the spermatic cord. Each ovarian vein drains an ovary in females. Each of these veins is generically called a gonadal vein. The right gonadal vein empties directly into the inferior vena cava, and the left gonadal vein empties into the left renal vein.

Each side of the diaphragm drains into a phrenic vein; the right phrenic vein empties directly into the inferior vena cava, whereas the left phrenic vein empties into the left renal vein. Blood supply from the liver drains into each hepatic vein and directly into the inferior vena cava. Since the inferior vena cava lies primarily to the right of the vertebral column and aorta, the left renal vein is longer, as are the left phrenic, adrenal, and gonadal veins. The longer length of the left renal vein makes the left kidney the primary target of surgeons removing this organ for donation. Figure 19 provides a flow chart of the veins flowing into the inferior vena cava. Table 15 summarizes the major veins of the abdominal region.

[caption id="" align="aligncenter" width="540"]This chart shows the connection between the different veins and the inferior vena cava. Figure 19. Venous Flow into Inferior Vena Cava. The flow chart summarizes veins that deliver blood to the inferior vena cava.[/caption]
Major Veins of the Abdominal Region (Table 15)
Vessel Description
Inferior vena cava Large systemic vein that drains blood from areas largely inferior to the diaphragm; empties into the right atrium
Lumbar veins Series of veins that drain the lumbar portion of the abdominal wall and spinal cord; the ascending lumbar veins drain into the azygos vein on the right or the hemiazygos vein on the left; the remaining lumbar veins drain directly into the inferior vena cava
Renal vein Largest vein entering the inferior vena cava; drains the kidneys and flows into the inferior vena cava
Adrenal vein Drains the adrenal or suprarenal; the right adrenal vein enters the inferior vena cava directly and the left adrenal vein enters the left renal vein
Testicular vein Drains the testes and forms part of the spermatic cord; the right testicular vein empties directly into the inferior vena cava and the left testicular vein empties into the left renal vein
Ovarian vein Drains the ovary; the right ovarian vein empties directly into the inferior vena cava and the left ovarian vein empties into the left renal vein
Gonadal vein Generic term for a vein draining a reproductive organ; may be either an ovarian vein or a testicular vein, depending on the sex of the individual
Phrenic vein Drains the diaphragm; the right phrenic vein flows into the inferior vena cava and the left phrenic vein empties into the left renal vein
Hepatic vein Drains systemic blood from the liver and flows into the inferior vena cava

Veins Draining the Lower Limbs

The superior surface of the foot drains into the digital veins, and the inferior surface drains into the plantar veins, which flow into a complex series of anastomoses in the feet and ankles, including the dorsal venous arch and the plantar venous arch (Figure 20). From the dorsal venous arch, blood supply drains into the anterior and posterior tibial veins. The anterior tibial vein drains the area near the tibialis anterior muscle and combines with the posterior tibial vein and the fibular vein to form the popliteal vein. The posterior tibial vein drains the posterior surface of the tibia and joins the popliteal vein. The fibular vein drains the muscles and integument in proximity to the fibula and also joins the popliteal vein. The small saphenous vein located on the lateral surface of the leg drains blood from the superficial regions of the lower leg and foot, and flows into to the popliteal vein. As the popliteal vein passes behind the knee in the popliteal region, it becomes the femoral vein. It is palpable in patients without excessive adipose tissue.

Close to the body wall, the great saphenous vein, the deep femoral vein, and the femoral circumflex vein drain into the femoral vein. The great saphenous vein is a prominent surface vessel located on the medial surface of the leg and thigh that collects blood from the superficial portions of these areas. The deep femoral vein, as the name suggests, drains blood from the deeper portions of the thigh. The femoral circumflex vein forms a loop around the femur just inferior to the trochanters and drains blood from the areas in proximity to the head and neck of the femur.

As the femoral vein penetrates the body wall from the femoral portion of the upper limb, it becomes the external iliac vein, a large vein that drains blood from the leg to the common iliac vein. The pelvic organs and integument drain into the internal iliac vein, which forms from several smaller veins in the region, including the umbilical veins that run on either side of the bladder. The external and internal iliac veins combine near the inferior portion of the sacroiliac joint to form the common iliac vein. In addition to blood supply from the external and internal iliac veins, the middle sacral vein drains the sacral region into the common iliac vein. Similar to the common iliac arteries, the common iliac veins come together at the level of L5 to form the inferior vena cava.

Figure 21 is a flow chart of veins flowing into the lower limb. Table 16 summarizes the major veins of the lower limbs.

[caption id="" align="aligncenter" width="480"]The left panel shows the anterior view of veins in the legs, and the right panel shows the posterior view. Figure 20. Major Veins Serving the Lower Limbs Anterior and posterior views show the major veins that drain the lower limb into the inferior vena cava.[/caption]
[caption id="" align="aligncenter" width="480"]This charts shows the veins in the lower limbs, and how they are connected. Figure 21. Major Veins of the Lower Limb. The flow chart summarizes venous flow from the lower limb.[/caption]
Veins of the Lower Limbs (Table 16)
Vessel Description
Plantar veins Drain the foot and flow into the plantar venous arch
Dorsal venous arch Drains blood from digital veins and vessels on the superior surface of the foot
Plantar venous arch Formed from the plantar veins; flows into the anterior and posterior tibial veins through anastomoses
Anterior tibial vein Formed from the dorsal venous arch; drains the area near the tibialis anterior muscle and flows into the popliteal vein
Posterior tibial vein Formed from the dorsal venous arch; drains the area near the posterior surface of the tibia and flows into the popliteal vein
Fibular vein Drains the muscles and integument near the fibula and flows into the popliteal vein
Small saphenous vein Located on the lateral surface of the leg; drains blood from the superficial regions of the lower leg and foot, and flows into the popliteal vein
Popliteal vein Drains the region behind the knee and forms from the fusion of the fibular, anterior, and posterior tibial veins; flows into the femoral vein
Great saphenous vein Prominent surface vessel located on the medial surface of the leg and thigh; drains the superficial portions of these areas and flows into the femoral vein
Deep femoral vein Drains blood from the deeper portions of the thigh and flows into the femoral vein
Femoral circumflex vein Forms a loop around the femur just inferior to the trochanters; drains blood from the areas around the head and neck of the femur; flows into the femoral vein
Femoral vein Drains the upper leg; receives blood from the great saphenous vein, the deep femoral vein, and the femoral circumflex vein; becomes the external iliac vein when it crosses the body wall
External iliac vein Formed when the femoral vein passes into the body cavity; drains the legs and flows into the common iliac vein
Internal iliac vein Drains the pelvic organs and integument; formed from several smaller veins in the region; flows into the common iliac vein
Middle sacral vein Drains the sacral region and flows into the left common iliac vein
Common iliac vein Flows into the inferior vena cava at the level of L5; the left common iliac vein drains the sacral region; formed from the union of the external and internal iliac veins near the inferior portion of the sacroiliac joint

Hepatic Portal System

The liver is a complex biochemical processing plant. It packages nutrients absorbed by the digestive system; produces plasma proteins, clotting factors, and bile; and disposes of worn-out cell components and waste products. Instead of entering the circulation directly, absorbed nutrients and certain wastes (for example, materials produced by the spleen) travel to the liver for processing. They do so via the hepatic portal system (Figure 22). Portal systems begin and end in capillaries. In this case, the initial capillaries from the stomach, small intestine, large intestine, and spleen lead to the hepatic portal vein and end in specialized capillaries within the liver, the hepatic sinusoids. You saw the only other portal system with the hypothalamic-hypophyseal portal vessel in the endocrine chapter.

The hepatic portal system consists of the hepatic portal vein and the veins that drain into it. The hepatic portal vein itself is relatively short, beginning at the level of L2 with the confluence of the superior mesenteric and splenic veins. It also receives branches from the inferior mesenteric vein, plus the splenic veins and all their tributaries. The superior mesenteric vein receives blood from the small intestine, two-thirds of the large intestine, and the stomach. The inferior mesenteric vein drains the distal third of the large intestine, including the descending colon, the sigmoid colon, and the rectum. The splenic vein is formed from branches from the spleen, pancreas, and portions of the stomach, and the inferior mesenteric vein. After its formation, the hepatic portal vein also receives branches from the gastric veins of the stomach and cystic veins from the gall bladder. The hepatic portal vein delivers materials from these digestive and circulatory organs directly to the liver for processing.

Because of the hepatic portal system, the liver receives its blood supply from two different sources: from normal systemic circulation via the hepatic artery and from the hepatic portal vein. The liver processes the blood from the portal system to remove certain wastes and excess nutrients, which are stored for later use. This processed blood, as well as the systemic blood that came from the hepatic artery, exits the liver via the right, left, and middle hepatic veins, and flows into the inferior vena cava. Overall systemic blood composition remains relatively stable, since the liver is able to metabolize the absorbed digestive components.

[caption id="" align="aligncenter" width="480"]This diagram shows the veins in the digestive system. Figure 22. Hepatic Portal System. The liver receives blood from the normal systemic circulation via the hepatic artery. It also receives and processes blood from other organs, delivered via the veins of the hepatic portal system. All blood exits the liver via the hepatic vein, which delivers the blood to the inferior vena cava. (Different colors are used to help distinguish among the different vessels in the system.)[/caption]

]]> 2309 2261 6 0 21.1 Anatomy of the Lymphatic and Immune Systems https://pressbooks.bccampus.ca/apdouglas2017/chapter/21-1-anatomy-of-the-lymphatic-and-immune-systems/ Fri, 14 Jul 2017 23:01:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2325

Learning Objectives

By the end of this section, you will be able to:
  • Describe the structure and function of the lymphatic tissue (lymph fluid, vessels, ducts, and organs)
  • Describe the structure and function of the primary and secondary lymphatic organs
  • Discuss the cells of the immune system, how they function, and their relationship with the lymphatic system

The immune system is the complex collection of cells and organs that destroys or neutralizes pathogens that would otherwise cause disease or death. The lymphatic system, for most people, is associated with the immune system to such a degree that the two systems are virtually indistinguishable. The lymphatic system is the system of vessels, cells, and organs that carries excess fluids to the bloodstream and filters pathogens from the blood. The swelling of lymph nodes during an infection and the transport of lymphocytes via the lymphatic vessels are but two examples of the many connections between these critical organ systems.

Functions of the Lymphatic System

A major function of the lymphatic system is to drain body fluids and return them to the bloodstream. Blood pressure causes leakage of fluid from the capillaries, resulting in the accumulation of fluid in the interstitial space—that is, spaces between individual cells in the tissues. In humans, 20 liters of plasma is released into the interstitial space of the tissues each day due to capillary filtration. Once this filtrate is out of the bloodstream and in the tissue spaces, it is referred to as interstitial fluid. Of this, 17 liters is reabsorbed directly by the blood vessels. But what happens to the remaining three liters? This is where the lymphatic system comes into play. It drains the excess fluid and empties it back into the bloodstream via a series of vessels, trunks, and ducts. Lymph is the term used to describe interstitial fluid once it has entered the lymphatic system. When the lymphatic system is damaged in some way, such as by being blocked by cancer cells or destroyed by injury, protein-rich interstitial fluid accumulates (sometimes “backs up” from the lymph vessels) in the tissue spaces. This inappropriate accumulation of fluid referred to as lymphedema may lead to serious medical consequences.

As the vertebrate immune system evolved, the network of lymphatic vessels became convenient avenues for transporting the cells of the immune system. Additionally, the transport of dietary lipids and fat-soluble vitamins absorbed in the gut uses this system.

Cells of the immune system not only use lymphatic vessels to make their way from interstitial spaces back into the circulation, but they also use lymph nodes as major staging areas for the development of critical immune responses. A lymph node is one of the small, bean-shaped organs located throughout the lymphatic system.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit this website for an overview of the lymphatic system.[/caption]

Structure of the Lymphatic System

The lymphatic vessels begin as open-ended capillaries, which feed into larger and larger lymphatic vessels, and eventually empty into the bloodstream by a series of ducts. Along the way, the lymph travels through the lymph nodes, which are commonly found near the groin, armpits, neck, chest, and abdomen. Humans have about 500–600 lymph nodes throughout the body (Figure 1).

[caption id="" align="aligncenter" width="480"]The left panel shows a female human body, and the entire lymphatic system is shown. The right panel shows magnified images of the thymus and the lymph node. All the major parts in the lymphatic system are labeled. Figure 1. Anatomy of the Lymphatic System. Lymphatic vessels in the arms and legs convey lymph to the larger lymphatic vessels in the torso.[/caption]

A major distinction between the lymphatic and cardiovascular systems in humans is that lymph is not actively pumped by the heart, but is forced through the vessels by the movements of the body, the contraction of skeletal muscles during body movements, and breathing. One-way valves (semi-lunar valves) in lymphatic vessels keep the lymph moving toward the heart. Lymph flows from the lymphatic capillaries, through lymphatic vessels, and then is dumped into the circulatory system via the lymphatic ducts located at the junction of the jugular and subclavian veins in the neck.

Lymphatic Capillaries

Lymphatic capillaries, also called the terminal lymphatics, are vessels where interstitial fluid enters the lymphatic system to become lymph fluid. Located in almost every tissue in the body, these vessels are interlaced among the arterioles and venules of the circulatory system in the soft connective tissues of the body (Figure 2). Exceptions are the central nervous system, bone marrow, bones, teeth, and the cornea of the eye, which do not contain lymph vessels.

[caption id="" align="aligncenter" width="500"]This image shows the lymph capillaries in the tissue spaces, and a magnified image shows the interstitial fluid and the lymph vessels. The major parts are labeled. Figure 2. Lymphatic Capillaries. Lymphatic capillaries are interlaced with the arterioles and venules of the cardiovascular system. Collagen fibers anchor a lymphatic capillary in the tissue (inset). Interstitial fluid slips through spaces between the overlapping endothelial cells that compose the lymphatic capillary.[/caption]

Lymphatic capillaries are formed by a one cell-thick layer of endothelial cells and represent the open end of the system, allowing interstitial fluid to flow into them via overlapping cells (see Figure 2). When interstitial pressure is low, the endothelial flaps close to prevent “backflow.” As interstitial pressure increases, the spaces between the cells open up, allowing the fluid to enter. Entry of fluid into lymphatic capillaries is also enabled by the collagen filaments that anchor the capillaries to surrounding structures. As interstitial pressure increases, the filaments pull on the endothelial cell flaps, opening up them even further to allow easy entry of fluid.

In the small intestine, lymphatic capillaries called lacteals are critical for the transport of dietary lipids and lipid-soluble vitamins to the bloodstream. In the small intestine, dietary triglycerides combine with other lipids and proteins, and enter the lacteals to form a milky fluid called chyle. The chyle then travels through the lymphatic system, eventually entering the liver and then the bloodstream.

Larger Lymphatic Vessels, Trunks, and Ducts

The lymphatic capillaries empty into larger lymphatic vessels, which are similar to veins in terms of their three-tunic structure and the presence of valves. These one-way valves are located fairly close to one another, and each one causes a bulge in the lymphatic vessel, giving the vessels a beaded appearance (see Figure 2).

The superficial and deep lymphatics eventually merge to form larger lymphatic vessels known as lymphatic trunks. On the right side of the body, the right sides of the head, thorax, and right upper limb drain lymph fluid into the right subclavian vein via the right lymphatic duct (Figure 3). On the left side of the body, the remaining portions of the body drain into the larger thoracic duct, which drains into the left subclavian vein. The thoracic duct itself begins just beneath the diaphragm in the cisterna chyli, a sac-like chamber that receives lymph from the lower abdomen, pelvis, and lower limbs by way of the left and right lumbar trunks and the intestinal trunk.

[caption id="" align="aligncenter" width="480"]This figure shows the lymphatic trunks and the duct system in the human body. Callouts to the left and right show the magnified views of the left and right jugular vein respectively. Figure 3. Major Trunks and Ducts of the Lymphatic System. The thoracic duct drains a much larger portion of the body than does the right lymphatic duct.[/caption]

The overall drainage system of the body is asymmetrical (see Figure 3). The right lymphatic duct receives lymph from only the upper right side of the body. The lymph from the rest of the body enters the bloodstream through the thoracic duct via all the remaining lymphatic trunks. In general, lymphatic vessels of the subcutaneous tissues of the skin, that is, the superficial lymphatics, follow the same routes as veins, whereas the deep lymphatic vessels of the viscera generally follow the paths of arteries.

The Organization of Immune Function

The immune system is a collection of barriers, cells, and soluble proteins that interact and communicate with each other in extraordinarily complex ways. The modern model of immune function is organized into three phases based on the timing of their effects. The three temporal phases consist of the following:

  • Barrier defenses such as the skin and mucous membranes, which act instantaneously to prevent pathogenic invasion into the body tissues
  • The rapid but nonspecific innate immune response, which consists of a variety of specialized cells and soluble factors
  • The slower but more specific and effective adaptive immune response, which involves many cell types and soluble factors, but is primarily controlled by white blood cells (leukocytes) known as lymphocytes, which help control immune responses
The cells of the blood, including all those involved in the immune response, arise in the bone marrow via various differentiation pathways from hematopoietic stem cells (Figure 4). In contrast with embryonic stem cells, hematopoietic stem cells are present throughout adulthood and allow for the continuous differentiation of blood cells to replace those lost to age or function. These cells can be divided into three classes based on function:
  • Phagocytic cells, which ingest pathogens to destroy them
  • Lymphocytes, which specifically coordinate the activities of adaptive immunity
  • Cells containing cytoplasmic granules, which help mediate immune responses against parasites and intracellular pathogens such as viruses
[caption id="" align="aligncenter" width="600"]This flowchart shows the steps in which a multipotential hematopoietic stem cell differentiates into the different cell types in blood. Figure 4. Hematopoietic System of the Bone Marrow. All the cells of the immune response as well as of the blood arise by differentiation from hematopoietic stem cells. Platelets are cell fragments involved in the clotting of blood.[/caption]

Lymphocytes: B Cells, T Cells, Plasma Cells, and Natural Killer Cells

As stated above, lymphocytes are the primary cells of adaptive immune responses (Table 1). The two basic types of lymphocytes, B cells and T cells, are identical morphologically with a large central nucleus surrounded by a thin layer of cytoplasm. They are distinguished from each other by their surface protein markers as well as by the molecules they secrete. While B cells mature in red bone marrow and T cells mature in the thymus, they both initially develop from bone marrow. T cells migrate from bone marrow to the thymus gland where they further mature. B cells and T cells are found in many parts of the body, circulating in the bloodstream and lymph, and residing in secondary lymphoid organs, including the spleen and lymph nodes, which will be described later in this section. The human body contains approximately 1012 lymphocytes.

B Cells

B cells are immune cells that function primarily by producing antibodies. An antibody is any of the group of proteins that binds specifically to pathogen-associated molecules known as antigens. An antigen is a chemical structure on the surface of a pathogen that binds to T or B lymphocyte antigen receptors. Once activated by binding to antigen, B cells differentiate into cells that secrete a soluble form of their surface antibodies. These activated B cells are known as plasma cells.

T Cells

The T cell, on the other hand, does not secrete antibody but performs a variety of functions in the adaptive immune response. Different T cell types have the ability to either secrete soluble factors that communicate with other cells of the adaptive immune response or destroy cells infected with intracellular pathogens. The roles of T and B lymphocytes in the adaptive immune response will be discussed further in this chapter.

Plasma Cells

Another type of lymphocyte of importance is the plasma cell. A plasma cell is a B cell that has differentiated in response to antigen binding, and has thereby gained the ability to secrete soluble antibodies. These cells differ in morphology from standard B and T cells in that they contain a large amount of cytoplasm packed with the protein-synthesizing machinery known as rough endoplasmic reticulum.

Natural Killer Cells

A fourth important lymphocyte is the natural killer cell, a participant in the innate immune response. A natural killer cell (NK) is a circulating blood cell that contains cytotoxic (cell-killing) granules in its extensive cytoplasm. It shares this mechanism with the cytotoxic T cells of the adaptive immune response. NK cells are among the body’s first lines of defense against viruses and certain types of cancer.

Lymphocytes
Type of lymphocyte Primary function
B lymphocyte Generates diverse antibodies
T lymphocyte Secretes chemical messengers
Plasma cell Secretes antibodies
NK cell Destroys virally infected cells
[caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit this website to learn about the many different cell types in the immune system and their very specialized jobs.[/caption]

Primary Lymphoid Organs and Lymphocyte Development

Understanding the differentiation and development of B and T cells is critical to the understanding of the adaptive immune response. It is through this process that the body (ideally) learns to destroy only pathogens and leaves the body’s own cells relatively intact. The primary lymphoid organs are the bone marrow and thymus gland. The lymphoid organs are where lymphocytes mature, proliferate, and are selected, which enables them to attack pathogens without harming the cells of the body.

Bone Marrow

In the embryo, blood cells are made in the yolk sac. As development proceeds, this function is taken over by the spleen, lymph nodes, and liver. Later, the bone marrow takes over most hematopoietic functions, although the final stages of the differentiation of some cells may take place in other organs. The red bone marrow is a loose collection of cells where hematopoiesis occurs, and the yellow bone marrow is a site of energy storage, which consists largely of fat cells (Figure 5). The B cell undergoes nearly all of its development in the red bone marrow, whereas the immature T cell, called a thymocyte, leaves the bone marrow and matures largely in the thymus gland.

[caption id="" align="aligncenter" width="300"]This photograph shows the bone marrow. Figure 5. Bone Marrow. Red bone marrow fills the head of the femur, and a spot of yellow bone marrow is visible in the center. The white reference bar is 1 cm.[/caption]

Thymus

The thymus gland is a bilobed organ found in the space between the sternum and the aorta of the heart (Figure 6). Connective tissue holds the lobes closely together but also separates them and forms a capsule.

[caption id="" align="aligncenter" width="480"]The left panel of this figure shows the head and chest of a woman and the location of the thymus is marked. The top right panel shows a micrograph of the thymus and the bottom right panel shows a magnified view of the structure of the thymus. Figure 6. Location, Structure, and Histology of the Thymus. The thymus lies above the heart. The trabeculae and lobules, including the darkly staining cortex and the lighter staining medulla of each lobule, are clearly visible in the light micrograph of the thymus of a newborn. LM × 100. (Micrograph provided by the Regents of the University of Michigan Medical School © 2012)[/caption]

The connective tissue capsule further divides the thymus into lobules via extensions called trabeculae. The outer region of the organ is known as the cortex and contains large numbers of thymocytes with some epithelial cells, macrophages, and dendritic cells (two types of phagocytic cells that are derived from monocytes). The cortex is densely packed so it stains more intensely than the rest of the thymus (see Figure 6). The medulla, where thymocytes migrate before leaving the thymus, contains a less dense collection of thymocytes, epithelial cells, and dendritic cells.

Secondary Lymphoid Organs and their Roles in Active Immune Responses

Lymphocytes develop and mature in the primary lymphoid organs, but they mount immune responses from the secondary lymphoid organs. A naïve lymphocyte is one that has left the primary organ and entered a secondary lymphoid organ. Naïve lymphocytes are fully functional immunologically, but have yet to encounter an antigen to respond to. In addition to circulating in the blood and lymph, lymphocytes concentrate in secondary lymphoid organs, which include the lymph nodes, spleen, and lymphoid nodules. All of these tissues have many features in common, including the following:

  • The presence of lymphoid follicles, the sites of the formation of lymphocytes, with specific B cell-rich and T cell-rich areas
  • An internal structure of reticular fibers with associated fixed macrophages
  • Germinal centers, which are the sites of rapidly dividing B lymphocytes and plasma cells, with the exception of the spleen
  • Specialized post-capillary vessels known as high endothelial venules; the cells lining these venules are thicker and more columnar than normal endothelial cells, which allow cells from the blood to directly enter these tissues

Lymph Nodes

Lymph nodes function to remove debris and pathogens from the lymph, and are thus sometimes referred to as the “filters of the lymph” (Figure 7). Any bacteria that infect the interstitial fluid are taken up by the lymphatic capillaries and transported to a regional lymph node. Dendritic cells and macrophages within this organ internalize and kill many of the pathogens that pass through, thereby removing them from the body. The lymph node is also the site of adaptive immune responses mediated by T cells, B cells, and accessory cells of the adaptive immune system. Like the thymus, the bean-shaped lymph nodes are surrounded by a tough capsule of connective tissue and are separated into compartments by trabeculae, the extensions of the capsule. In addition to the structure provided by the capsule and trabeculae, the structural support of the lymph node is provided by a series of reticular fibers laid down by fibroblasts.

[caption id="" align="aligncenter" width="500"]The left panel of this figure shows a micrograph of the cross section of a lymph node. The right panel shows the structure of a lymph node. Figure 7. Structure and Histology of a Lymph Node. Lymph nodes are masses of lymphatic tissue located along the larger lymph vessels. The micrograph of the lymph nodes shows a germinal center, which consists of rapidly dividing B cells surrounded by a layer of T cells and other accessory cells. LM × 128. (Micrograph provided by the Regents of the University of Michigan Medical School © 2012)[/caption]

The major routes into the lymph node are via afferent lymphatic vessels (see [link]). Cells and lymph fluid that leave the lymph node may do so by another set of vessels known as the efferent lymphatic vessels. Lymph enters the lymph node via the subcapsular sinus, which is occupied by dendritic cells, macrophages, and reticular fibers. Within the cortex of the lymph node are lymphoid follicles, which consist of germinal centers of rapidly dividing B cells surrounded by a layer of T cells and other accessory cells. As the lymph continues to flow through the node, it enters the medulla, which consists of medullary cords of B cells and plasma cells, and the medullary sinuses where the lymph collects before leaving the node via the efferent lymphatic vessels.

Spleen

In addition to the lymph nodes, the spleen is a major secondary lymphoid organ (Figure 8). It is about 12 cm (5 in) long and is attached to the lateral border of the stomach via the gastrosplenic ligament. The spleen is a fragile organ without a strong capsule, and is dark red due to its extensive vascularization. The spleen is sometimes called the “filter of the blood” because of its extensive vascularization and the presence of macrophages and dendritic cells that remove microbes and other materials from the blood, including dying red blood cells. The spleen also functions as the location of immune responses to blood-borne pathogens.

[caption id="" align="aligncenter" width="500"]The top left panel shows the location of the spleen in the human body. The top center panel shows a close up view of the location of the spleen. The top right panel shows the blood vessels and spleen tissue. The bottom panel shows a histological micrograph. Figure 8. Spleen. (a) The spleen is attached to the stomach. (b) A micrograph of spleen tissue shows the germinal center. The marginal zone is the region between the red pulp and white pulp, which sequesters particulate antigens from the circulation and presents these antigens to lymphocytes in the white pulp. EM × 660. (Micrograph provided by the Regents of the University of Michigan Medical School © 2012)[/caption]

The spleen is also divided by trabeculae of connective tissue, and within each splenic nodule is an area of red pulp, consisting of mostly red blood cells, and white pulp, which resembles the lymphoid follicles of the lymph nodes. Upon entering the spleen, the splenic artery splits into several arterioles (surrounded by white pulp) and eventually into sinusoids. Blood from the capillaries subsequently collects in the venous sinuses and leaves via the splenic vein. The red pulp consists of reticular fibers with fixed macrophages attached, free macrophages, and all of the other cells typical of the blood, including some lymphocytes. The white pulp surrounds a central arteriole and consists of germinal centers of dividing B cells surrounded by T cells and accessory cells, including macrophages and dendritic cells. Thus, the red pulp primarily functions as a filtration system of the blood, using cells of the relatively nonspecific immune response, and white pulp is where adaptive T and B cell responses are mounted.

Lymphoid Nodules

The other lymphoid tissues, the lymphoid nodules, have a simpler architecture than the spleen and lymph nodes in that they consist of a dense cluster of lymphocytes without a surrounding fibrous capsule. These nodules are located in the respiratory and digestive tracts, areas routinely exposed to environmental pathogens.

Tonsils are lymphoid nodules located along the inner surface of the pharynx and are important in developing immunity to oral pathogens (Figure 9). The tonsil located at the back of the throat, the pharyngeal tonsil, is sometimes referred to as the adenoid when swollen. Such swelling is an indication of an active immune response to infection. Histologically, tonsils do not contain a complete capsule, and the epithelial layer invaginates deeply into the interior of the tonsil to form tonsillar crypts. These structures, which accumulate all sorts of materials taken into the body through eating and breathing, actually “encourage” pathogens to penetrate deep into the tonsillar tissues where they are acted upon by numerous lymphoid follicles and eliminated. This seems to be the major function of tonsils—to help children’s bodies recognize, destroy, and develop immunity to common environmental pathogens so that they will be protected in their later lives. Tonsils are often removed in those children who have recurring throat infections, especially those involving the palatine tonsils on either side of the throat, whose swelling may interfere with their breathing and/or swallowing.

[caption id="" align="aligncenter" width="480"]The top panel of this image shows the location of the tonsils. All the major parts are labeled. The bottom panel shows the histological micrograph of the tonsils. Figure 9. Locations and Histology of the Tonsils. (a) The pharyngeal tonsil is located on the roof of the posterior superior wall of the nasopharynx. The palatine tonsils lay on each side of the pharynx. (b) A micrograph shows the palatine tonsil tissue. LM × 40. (Micrograph provided by the Regents of the University of Michigan Medical School © 2012)[/caption]

Mucosa-associated lymphoid tissue (MALT) consists of an aggregate of lymphoid follicles directly associated with the mucous membrane epithelia. MALT makes up dome-shaped structures found underlying the mucosa of the gastrointestinal tract, breast tissue, lungs, and eyes. Peyer’s patches, a type of MALT in the small intestine, are especially important for immune responses against ingested substances (Figure 10). Peyer’s patches contain specialized endothelial cells called M (or microfold) cells that sample material from the intestinal lumen and transport it to nearby follicles so that adaptive immune responses to potential pathogens can be mounted.

[caption id="" align="aligncenter" width="480"]This figure shows a micrograph of a mucosa associated lymphoid tissue nodule. Figure 10. Mucosa-associated Lymphoid Tissue (MALT) Nodule. LM × 40. (Micrograph provided by the Regents of the University of Michigan Medical School © 2012)[/caption]

Bronchus-associated lymphoid tissue (BALT) consists of lymphoid follicular structures with an overlying epithelial layer found along the bifurcations of the bronchi, and between bronchi and arteries. They also have the typically less-organized structure of other lymphoid nodules. These tissues, in addition to the tonsils, are effective against inhaled pathogens.

[caption id="attachment_2978" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about the lymphatic system.[/caption]
]]> 2325 2310 2 0 21.2 Barrier Defenses and the Innate Immune Response https://pressbooks.bccampus.ca/apdouglas2017/chapter/21-2-barrier-defenses-and-the-innate-immune-response/ Fri, 14 Jul 2017 23:02:10 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2329

Learning Objectives

By the end of this section, you will be able to:
  • Describe the barrier defenses of the body
  • Show how the innate immune response is important and how it helps guide and prepare the body for adaptive immune responses
  • Describe various soluble factors that are part of the innate immune response
  • Explain the steps of inflammation and how they lead to destruction of a pathogen
  • Discuss early induced immune responses and their level of effectiveness

The immune system can be divided into two overlapping mechanisms to destroy pathogens: the innate immune response, which is relatively rapid but nonspecific and thus not always effective, and the adaptive immune response, which is slower in its development during an initial infection with a pathogen, but is highly specific and effective at attacking a wide variety of pathogens (Figure 1).

[caption id="" align="aligncenter" width="550"]This figure shows a lateral view of a human face in the top left. A magnified callout shows the germinal center of the palatine tonsil. Another magnified view shows how the innate immune system works. Figure 1. Cooperation between Innate and Adaptive Immune Responses. The innate immune system enhances adaptive immune responses so they can be more effective[/caption]

Any discussion of the innate immune response usually begins with the physical barriers that prevent pathogens from entering the body, destroy them after they enter, or flush them out before they can establish themselves in the hospitable environment of the body’s soft tissues. Barrier defenses are part of the body’s most basic defense mechanisms. The barrier defenses are not a response to infections, but they are continuously working to protect against a broad range of pathogens.

The different modes of barrier defenses are associated with the external surfaces of the body, where pathogens may try to enter (Table 2). The primary barrier to the entrance of microorganisms into the body is the skin. Not only is the skin covered with a layer of dead, keratinized epithelium that is too dry for bacteria in which to grow, but as these cells are continuously sloughed off from the skin, they carry bacteria and other pathogens with them. Additionally, sweat and other skin secretions may lower pH, contain toxic lipids, and physically wash microbes away.
Barrier Defenses (Table 2)
Site Specific defense Protective aspect
Skin Epidermal surface Keratinized cells of surface, Langerhans cells
Skin (sweat/secretions) Sweat glands, sebaceous glands Low pH, washing action
Oral cavity Salivary glands Lysozyme
Stomach Gastrointestinal tract Low pH
Mucosal surfaces Mucosal epithelium Nonkeratinized epithelial cells
Normal flora (nonpathogenic bacteria) Mucosal tissues Prevent pathogens from growing on mucosal surfaces

Another barrier is the saliva in the mouth, which is rich in lysozyme—an enzyme that destroys bacteria by digesting their cell walls. The acidic environment of the stomach, which is fatal to many pathogens, is also a barrier. Additionally, the mucus layer of the gastrointestinal tract, respiratory tract, reproductive tract, eyes, ears, and nose traps both microbes and debris, and facilitates their removal. In the case of the upper respiratory tract, ciliated epithelial cells move potentially contaminated mucus upwards to the mouth, where it is then swallowed into the digestive tract, ending up in the harsh acidic environment of the stomach. Considering how often you breathe compared to how often you eat or perform other activities that expose you to pathogens, it is not surprising that multiple barrier mechanisms have evolved to work in concert to protect this vital area.

Cells of the Innate Immune Response

A phagocyte is a cell that is able to surround and engulf a particle or cell, a process called phagocytosis. The phagocytes of the immune system engulf other particles or cells, either to clean an area of debris, old cells, or to kill pathogenic organisms such as bacteria. The phagocytes are the body’s fast acting, first line of immunological defense against organisms that have breached barrier defenses and have entered the vulnerable tissues of the body.

Phagocytes: Macrophages and Neutrophils

Many of the cells of the immune system have a phagocytic ability, at least at some point during their life cycles. Phagocytosis is an important and effective mechanism of destroying pathogens during innate immune responses. The phagocyte takes the organism inside itself as a phagosome, which subsequently fuses with a lysosome and its digestive enzymes, effectively killing many pathogens. On the other hand, some bacteria including Mycobacteria tuberculosis, the cause of tuberculosis, may be resistant to these enzymes and are therefore much more difficult to clear from the body. Macrophages, neutrophils, and dendritic cells are the major phagocytes of the immune system.

A macrophage is an irregularly shaped phagocyte that is amoeboid in nature and is the most versatile of the phagocytes in the body. Macrophages move through tissues and squeeze through capillary walls using pseudopodia. They not only participate in innate immune responses but have also evolved to cooperate with lymphocytes as part of the adaptive immune response. Macrophages exist in many tissues of the body, either freely roaming through connective tissues or fixed to reticular fibers within specific tissues such as lymph nodes. When pathogens breach the body’s barrier defenses, macrophages are the first line of defense (Table 3). They are called different names, depending on the tissue: Kupffer cells in the liver, histiocytes in connective tissue, and alveolar macrophages in the lungs.

A neutrophil is a phagocytic cell that is attracted via chemotaxis from the bloodstream to infected tissues. These spherical cells are granulocytes. A granulocyte contains cytoplasmic granules, which in turn contain a variety of vasoactive mediators such as histamine. In contrast, macrophages are agranulocytes. An agranulocyte has few or no cytoplasmic granules. Whereas macrophages act like sentries, always on guard against infection, neutrophils can be thought of as military reinforcements that are called into a battle to hasten the destruction of the enemy. Although, usually thought of as the primary pathogen-killing cell of the inflammatory process of the innate immune response, new research has suggested that neutrophils play a role in the adaptive immune response as well, just as macrophages do.

A monocyte is a circulating precursor cell that differentiates into either a macrophage or dendritic cell, which can be rapidly attracted to areas of infection by signal molecules of inflammation.

Phagocytic Cells of the Innate Immune System (Table 3)
Cell Cell type Primary location Function in the innate immune response
Macrophage Agranulocyte Body cavities/organs Phagocytosis
Neutrophil Granulocyte Blood Phagocytosis
Monocyte Agranulocyte Blood Precursor of macrophage/dendritic cell

Natural Killer Cells

NK cells are a type of lymphocyte that have the ability to induce apoptosis, that is, programmed cell death, in cells infected with intracellular pathogens such as obligate intracellular bacteria and viruses. NK cells recognize these cells by mechanisms that are still not well understood, but that presumably involve their surface receptors. NK cells can induce apoptosis, in which a cascade of events inside the cell causes its own death by either of two mechanisms:

1) NK cells are able to respond to chemical signals and express the fas ligand. The fas ligand is a surface molecule that binds to the fas molecule on the surface of the infected cell, sending it apoptotic signals, thus killing the cell and the pathogen within it; or

2) The granules of the NK cells release perforins and granzymes. A perforin is a protein that forms pores in the membranes of infected cells. A granzyme is a protein-digesting enzyme that enters the cell via the perforin pores and triggers apoptosis intracellularly.

Both mechanisms are especially effective against virally infected cells. If apoptosis is induced before the virus has the ability to synthesize and assemble all its components, no infectious virus will be released from the cell, thus preventing further infection.

Recognition of Pathogens

Cells of the innate immune response, the phagocytic cells, and the cytotoxic NK cells recognize patterns of pathogen-specific molecules, such as bacterial cell wall components or bacterial flagellar proteins, using pattern recognition receptors. A pattern recognition receptor (PRR) is a membrane-bound receptor that recognizes characteristic features of a pathogen and molecules released by stressed or damaged cells.

These receptors, which are thought to have evolved prior to the adaptive immune response, are present on the cell surface whether they are needed or not. Their variety, however, is limited by two factors. First, the fact that each receptor type must be encoded by a specific gene requires the cell to allocate most or all of its DNA to make receptors able to recognize all pathogens. Secondly, the variety of receptors is limited by the finite surface area of the cell membrane. Thus, the innate immune system must “get by” using only a limited number of receptors that are active against as wide a variety of pathogens as possible. This strategy is in stark contrast to the approach used by the adaptive immune system, which uses large numbers of different receptors, each highly specific to a particular pathogen.

Should the cells of the innate immune system come into contact with a species of pathogen they recognize, the cell will bind to the pathogen and initiate phagocytosis (or cellular apoptosis in the case of an intracellular pathogen) in an effort to destroy the offending microbe. Receptors vary somewhat according to cell type, but they usually include receptors for bacterial components and for complement, discussed below.

Soluble Mediators of the Innate Immune Response

The previous discussions have alluded to chemical signals that can induce cells to change various physiological characteristics, such as the expression of a particular receptor. These soluble factors are secreted during innate or early induced responses, and later during adaptive immune responses.

Cytokines and Chemokines

A cytokine is signaling molecule that allows cells to communicate with each other over short distances. Cytokines are secreted into the intercellular space, and the action of the cytokine induces the receiving cell to change its physiology. A chemokine is a soluble chemical mediator similar to cytokines except that its function is to attract cells (chemotaxis) from longer distances.

Early induced Proteins

Early induced proteins are those that are not constitutively present in the body, but are made as they are needed early during the innate immune response. Interferons are an example of early induced proteins. Cells infected with viruses secrete interferons that travel to adjacent cells and induce them to make antiviral proteins. Thus, even though the initial cell is sacrificed, the surrounding cells are protected. Other early induced proteins specific for bacterial cell wall components are mannose-binding protein and C-reactive protein, made in the liver, which bind specifically to polysaccharide components of the bacterial cell wall. Phagocytes such as macrophages have receptors for these proteins, and they are thus able to recognize them as they are bound to the bacteria. This brings the phagocyte and bacterium into close proximity and enhances the phagocytosis of the bacterium by the process known as opsonization. Opsonization is the tagging of a pathogen for phagocytosis by the binding of an antibody or an antimicrobial protein.

Complement System

The complement system is a series of proteins constitutively found in the blood plasma. As such, these proteins are not considered part of the early induced immune response, even though they share features with some of the antibacterial proteins of this class. Made in the liver, they have a variety of functions in the innate immune response, using what is known as the “alternate pathway” of complement activation. Additionally, complement functions in the adaptive immune response as well, in what is called the classical pathway. The complement system consists of several proteins that enzymatically alter and fragment later proteins in a series, which is why it is termed cascade. Once activated, the series of reactions is irreversible, and releases fragments that have the following actions:

  • Bind to the cell membrane of the pathogen that activates it, labeling it for phagocytosis (opsonization)
  • Diffuse away from the pathogen and act as chemotactic agents to attract phagocytic cells to the site of inflammation
  • Form damaging pores in the plasma membrane of the pathogen

Figure 2 shows the classical pathway, which requires antibodies of the adaptive immune response. The alternate pathway does not require an antibody to become activated.

[caption id="" align="aligncenter" width="500"]This flow chart shows an invading pathogen and the series of events that results in the complement cascade and function. Figure 2. Complement Cascade and Function. The classical pathway, used during adaptive immune responses, occurs when C1 reacts with antibodies that have bound an antigen.[/caption]

The splitting of the C3 protein is the common step to both pathways. In the alternate pathway, C3 is activated spontaneously and, after reacting with the molecules factor P, factor B, and factor D, splits apart. The larger fragment, C3b, binds to the surface of the pathogen and C3a, the smaller fragment, diffuses outward from the site of activation and attracts phagocytes to the site of infection. Surface-bound C3b then activates the rest of the cascade, with the last five proteins, C5–C9, forming the membrane-attack complex (MAC). The MAC can kill certain pathogens by disrupting their osmotic balance. The MAC is especially effective against a broad range of bacteria. The classical pathway is similar, except the early stages of activation require the presence of antibody bound to antigen, and thus is dependent on the adaptive immune response. The earlier fragments of the cascade also have important functions. Phagocytic cells such as macrophages and neutrophils are attracted to an infection site by chemotactic attraction to smaller complement fragments. Additionally, once they arrive, their receptors for surface-bound C3b opsonize the pathogen for phagocytosis and destruction.

Inflammatory Response

The hallmark of the innate immune response is inflammation. Inflammation is something everyone has experienced. Stub a toe, cut a finger, or do any activity that causes tissue damage and inflammation will result, with its four characteristics: heat, redness, pain, and swelling (“loss of function” is sometimes mentioned as a fifth characteristic). It is important to note that inflammation does not have to be initiated by an infection, but can also be caused by tissue injuries. The release of damaged cellular contents into the site of injury is enough to stimulate the response, even in the absence of breaks in physical barriers that would allow pathogens to enter (by hitting your thumb with a hammer, for example). The inflammatory reaction brings in phagocytic cells to the damaged area to clear cellular debris and to set the stage for wound repair (Figure 3).

[caption id="" align="aligncenter" width="350"]The top panel of this figure shows the mast cells detecting an injury and initiating an inflammatory response. The bottom panel shows the increase in blood flow in response to histamine. Figure 3. Inflammatory Response.[/caption]

This reaction also brings in the cells of the innate immune system, allowing them to get rid of the sources of a possible infection. Inflammation is part of a very basic form of immune response. The process not only brings fluid and cells into the site to destroy the pathogen and remove it and debris from the site, but also helps to isolate the site, limiting the spread of the pathogen. Acute inflammation is a short-term inflammatory response to an insult to the body. If the cause of the inflammation is not resolved, however, it can lead to chronic inflammation, which is associated with major tissue destruction and fibrosis. Chronic inflammation is ongoing inflammation. It can be caused by foreign bodies, persistent pathogens, and autoimmune diseases such as rheumatoid arthritis.

There are four important parts to the inflammatory response:

  • Tissue Injury. The released contents of injured cells stimulate the release of mast cell granules and their potent inflammatory mediators such as histamine, leukotrienes, and prostaglandins. Histamine increases the diameter of local blood vessels (vasodilation), causing an increase in blood flow. Histamine also increases the permeability of local capillaries, causing plasma to leak out and form interstitial fluid. This causes the swelling associated with inflammation.
    Additionally, injured cells, phagocytes, and basophils are sources of inflammatory mediators, including prostaglandins and leukotrienes. Leukotrienes attract neutrophils from the blood by chemotaxis and increase vascular permeability. Prostaglandins cause vasodilation by relaxing vascular smooth muscle and are a major cause of the pain associated with inflammation. Nonsteroidal anti-inflammatory drugs such as aspirin and ibuprofen relieve pain by inhibiting prostaglandin production.
  • Vasodilation. Many inflammatory mediators such as histamine are vasodilators that increase the diameters of local capillaries. This causes increased blood flow and is responsible for the heat and redness of inflamed tissue. It allows greater access of the blood to the site of inflammation.
  • Increased Vascular Permeability. At the same time, inflammatory mediators increase the permeability of the local vasculature, causing leakage of fluid into the interstitial space, resulting in the swelling, or edema, associated with inflammation.
  • Recruitment of Phagocytes. Leukotrienes are particularly good at attracting neutrophils from the blood to the site of infection by chemotaxis. Following an early neutrophil infiltrate stimulated by macrophage cytokines, more macrophages are recruited to clean up the debris left over at the site. When local infections are severe, neutrophils are attracted to the sites of infections in large numbers, and as they phagocytose the pathogens and subsequently die, their accumulated cellular remains are visible as pus at the infection site.

Overall, inflammation is valuable for many reasons. Not only are the pathogens killed and debris removed, but the increase in vascular permeability encourages the entry of clotting factors, the first step towards wound repair. Inflammation also facilitates the transport of antigen to lymph nodes by dendritic cells for the development of the adaptive immune response.

[caption id="attachment_2980" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about the immune system.[/caption]
]]> 2329 2310 3 0 21.4 The Adaptive Immune Response: B-lymphocytes and Antibodies https://pressbooks.bccampus.ca/apdouglas2017/chapter/21-4-the-adaptive-immune-response-b-lymphocytes-and-antibodies/ Fri, 14 Jul 2017 23:03:15 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2342

Learning Objectives

By the end of this section, you will be able to:
  • Explain how B cells mature and how B cell tolerance develops
  • Discuss how B cells are activated and differentiate into plasma cells
  • Describe the structure of the antibody classes and their functions

Antibodies were the first component of the adaptive immune response to be characterized by scientists working on the immune system. It was already known that individuals who survived a bacterial infection were immune to re-infection with the same pathogen. Early microbiologists took serum from an immune patient and mixed it with a fresh culture of the same type of bacteria, then observed the bacteria under a microscope. The bacteria became clumped in a process called agglutination. When a different bacterial species was used, the agglutination did not happen. Thus, there was something in the serum of immune individuals that could specifically bind to and agglutinate bacteria.

Scientists now know the cause of the agglutination is an antibody molecule, also called an immunoglobulin. What is an antibody? An antibody protein is essentially a secreted form of a B cell receptor. (In fact, surface immunoglobulin is another name for the B cell receptor.) Not surprisingly, the same genes encode both the secreted antibodies and the surface immunoglobulins. One minor difference in the way these proteins are synthesized distinguishes a naïve B cell with antibody on its surface from an antibody-secreting plasma cell with no antibodies on its surface. The antibodies of the plasma cell have the exact same antigen-binding site and specificity as their B cell precursors.

There are five different classes of antibody found in humans: IgM, IgD, IgG, IgA, and IgE. Each of these has specific functions in the immune response, so by learning about them, researchers can learn about the great variety of antibody functions critical to many adaptive immune responses.

B cells do not recognize antigen in the complex fashion of T cells. B cells can recognize native, unprocessed antigen and do not require the participation of MHC molecules and antigen-presenting cells.

B Cell Differentiation and Activation

B cells differentiate in the bone marrow. During the process of maturation, up to 100 trillion different clones of B cells are generated, which is similar to the diversity of antigen receptors seen in T cells.

B cell differentiation and the development of tolerance are not quite as well understood as it is in T cells. Central tolerance is the destruction or inactivation of B cells that recognize self-antigens in the bone marrow, and its role is critical and well established. In the process of clonal deletion, immature B cells that bind strongly to self-antigens expressed on tissues are signaled to commit suicide by apoptosis, removing them from the population. In the process of clonal anergy, however, B cells exposed to soluble antigen in the bone marrow are not physically deleted, but become unable to function.

Another mechanism called peripheral tolerance is a direct result of T cell tolerance. In peripheral tolerance, functional, mature B cells leave the bone marrow but have yet to be exposed to self-antigen. Most protein antigens require signals from helper T cells (Th2) to proceed to make antibody. When a B cell binds to a self-antigen but receives no signals from a nearby Th2 cell to produce antibody, the cell is signaled to undergo apoptosis and is destroyed. This is yet another example of the control that T cells have over the adaptive immune response.

After B cells are activated by their binding to antigen, they differentiate into plasma cells. Plasma cells often leave the secondary lymphoid organs, where the response is generated, and migrate back to the bone marrow, where the whole differentiation process started. After secreting antibodies for a specific period, they die, as most of their energy is devoted to making antibodies and not to maintaining themselves. Thus, plasma cells are said to be terminally differentiated.

The final B cell of interest is the memory B cell, which results from the clonal expansion of an activated B cell. Memory B cells function in a way similar to memory T cells. They lead to a stronger and faster secondary response when compared to the primary response, as illustrated below.

Antibody Structure

Antibodies are glycoproteins consisting of two types of polypeptide chains with attached carbohydrates. The heavy chain and the light chain are the two polypeptides that form the antibody. The main differences between the classes of antibodies are in the differences between their heavy chains, but as you shall see, the light chains have an important role, forming part of the antigen-binding site on the antibody molecules.

Four-chain Models of Antibody Structures

All antibody molecules have two identical heavy chains and two identical light chains. (Some antibodies contain multiple units of this four-chain structure.) The Fc region of the antibody is formed by the two heavy chains coming together, usually linked by disulfide bonds (Figure 1). The Fc portion of the antibody is important in that many effector cells of the immune system have Fc receptors. Cells having these receptors can then bind to antibody-coated pathogens, greatly increasing the specificity of the effector cells. At the other end of the molecule are two identical antigen-binding sites.

[caption id="attachment_1771" align="aligncenter" width="550"]This diagram shows the four chain structure of a generic antibody. Figure 1. Antibody and IgG2 Structures. The typical four chain structure of a generic antibody (a) and the corresponding three-dimensional structure of the antibody IgG2 (b). (credit b: modification of work by Tim Vickers)[/caption]

Five Classes of Antibodies and their Functions

In general, antibodies have two basic functions. They can act as the B cell antigen receptor or they can be secreted, circulate, and bind to a pathogen, often labeling it for identification by other forms of the immune response. Of the five antibody classes, notice that only two can function as the antigen receptor for naïve B cells: IgM and IgD (Figure 2). Mature B cells that leave the bone marrow express both IgM and IgD, but both antibodies have the same antigen specificity. Only IgM is secreted, however, and no other nonreceptor function for IgD has been discovered.

[caption id="" align="aligncenter" width="480"]This table shows the five classes of the immunoglobulins. The table shows the molecular weight, number of antigen binding sites, and their function. Figure 2. Five Classes of Antibodies.[/caption]

IgM consists of five four-chain structures (20 total chains with 10 identical antigen-binding sites) and is thus the largest of the antibody molecules. IgM is usually the first antibody made during a primary response. Its 10 antigen-binding sites and large shape allow it to bind well to many bacterial surfaces. It is excellent at binding complement proteins and activating the complement cascade, consistent with its role in promoting chemotaxis, opsonization, and cell lysis. Thus, it is a very effective antibody against bacteria at early stages of a primary antibody response. As the primary response proceeds, the antibody produced in a B cell can change to IgG, IgA, or IgE by the process known as class switching. Class switching is the change of one antibody class to another. While the class of antibody changes, the specificity and the antigen-binding sites do not. Thus, the antibodies made are still specific to the pathogen that stimulated the initial IgM response.

IgG is a major antibody of late primary responses and the main antibody of secondary responses in the blood. This is because class switching occurs during primary responses. IgG is a monomeric antibody that clears pathogens from the blood and can activate complement proteins (although not as well as IgM), taking advantage of its antibacterial activities. Furthermore, this class of antibody is the one that crosses the placenta to protect the developing fetus from disease exits the blood to the interstitial fluid to fight extracellular pathogens.

IgA exists in two forms, a four-chain monomer in the blood and an eight-chain structure, or dimer, in exocrine gland secretions of the mucous membranes, including mucus, saliva, and tears. Thus, dimeric IgA is the only antibody to leave the interior of the body to protect body surfaces. IgA is also of importance to newborns, because this antibody is present in mother’s breast milk (colostrum), which serves to protect the infant from disease.

IgE is usually associated with allergies and anaphylaxis. It is present in the lowest concentration in the blood, because its Fc region binds strongly to an IgE-specific Fc receptor on the surfaces of mast cells. IgE makes mast cell degranulation very specific, such that if a person is allergic to peanuts, there will be peanut-specific IgE bound to his or her mast cells. In this person, eating peanuts will cause the mast cells to degranulate, sometimes causing severe allergic reactions, including anaphylaxis, a severe, systemic allergic response that can cause death.

Clonal Selection of B Cells

Clonal selection and expansion work much the same way in B cells as in T cells. Only B cells with appropriate antigen specificity are selected for and expanded (Figure 3). Eventually, the plasma cells secrete antibodies with antigenic specificity identical to those that were on the surfaces of the selected B cells. Notice in the figure that both plasma cells and memory B cells are generated simultaneously.

[caption id="" align="aligncenter" width="480"]This flow chart shows how the clonal selection of B cells takes place. The left panel shows the primary response and the right panel shows the secondary response. Figure 3. Clonal Selection of B Cells. During a primary B cell immune response, both antibody-secreting plasma cells and memory B cells are produced. These memory cells lead to the differentiation of more plasma cells and memory B cells during secondary responses.[/caption]

Primary versus Secondary B Cell Responses

Primary and secondary responses as they relate to T cells were discussed earlier. This section will look at these responses with B cells and antibody production. Because antibodies are easily obtained from blood samples, they are easy to follow and graph (Figure 4). As you will see from the figure, the primary response to an antigen (representing a pathogen) is delayed by several days. This is the time it takes for the B cell clones to expand and differentiate into plasma cells. The level of antibody produced is low, but it is sufficient for immune protection. The second time a person encounters the same antigen, there is no time delay, and the amount of antibody made is much higher. Thus, the secondary antibody response overwhelms the pathogens quickly and, in most situations, no symptoms are felt. When a different antigen is used, another primary response is made with its low antibody levels and time delay.

[caption id="" align="aligncenter" width="380"]This graph shows the antibody concentration as a function of time in primary and secondary response. Figure 4. Primary and Secondary Antibody Responses. Antigen A is given once to generate a primary response and later to generate a secondary response. When a different antigen is given for the first time, a new primary response is made.[/caption]

Active versus Passive Immunity

Immunity to pathogens, and the ability to control pathogen growth so that damage to the tissues of the body is limited, can be acquired by (1) the active development of an immune response in the infected individual or (2) the passive transfer of immune components from an immune individual to a nonimmune one. Both active and passive immunity have examples in the natural world and as part of medicine.

Active immunity is the resistance to pathogens acquired during an adaptive immune response within an individual (Table 6). Naturally acquired active immunity, the response to a pathogen, is the focus of this chapter. Artificially acquired active immunity involves the use of vaccines. A vaccine is a killed or weakened pathogen or its components that, when administered to a healthy individual, leads to the development of immunological memory (a weakened primary immune response) without causing much in the way of symptoms. Thus, with the use of vaccines, one can avoid the damage from disease that results from the first exposure to the pathogen, yet reap the benefits of protection from immunological memory. The advent of vaccines was one of the major medical advances of the twentieth century and led to the eradication of smallpox and the control of many infectious diseases, including polio, measles, and whooping cough.

Active versus Passive Immunity (Table 6)
Natural Artificial
Active Adaptive immune response Vaccine response
Passive Trans-placental antibodies/breastfeeding Immune globulin injections
Passive immunity arises from the transfer of antibodies to an individual without requiring them to mount their own active immune response. Naturally acquired passive immunity is seen during fetal development. IgG is transferred from the maternal circulation to the fetus via the placenta, protecting the fetus from infection and protecting the newborn for the first few months of its life. As already stated, a newborn benefits from the IgA antibodies it obtains from milk during breastfeeding. The fetus and newborn thus benefit from the immunological memory of the mother to the pathogens to which she has been exposed. In medicine, artificially acquired passive immunity usually involves injections of immunoglobulins, taken from animals previously exposed to a specific pathogen. This treatment is a fast-acting method of temporarily protecting an individual who was possibly exposed to a pathogen. The downside to both types of passive immunity is the lack of the development of immunological memory. Once the antibodies are transferred, they are effective for only a limited time before they degrade.

T cell-dependent versus T cell-independent Antigens

As discussed previously, Th2 cells secrete cytokines that drive the production of antibodies in a B cell, responding to complex antigens such as those made by proteins. On the other hand, some antigens are T cell independent. A T cell-independent antigen usually is in the form of repeated carbohydrate moieties found on the cell walls of bacteria. Each antibody on the B cell surface has two binding sites, and the repeated nature of T cell-independent antigen leads to crosslinking of the surface antibodies on the B cell. The crosslinking is enough to activate it in the absence of T cell cytokines.

A T cell-dependent antigen, on the other hand, usually is not repeated to the same degree on the pathogen and thus does not crosslink surface antibody with the same efficiency. To elicit a response to such antigens, the B and T cells must come close together (Figure 5). The B cell must receive two signals to become activated. Its surface immunoglobulin must recognize native antigen. Some of this antigen is internalized, processed, and presented to the Th2 cells on a class II MHC molecule. The T cell then binds using its antigen receptor and is activated to secrete cytokines that diffuse to the B cell, finally activating it completely. Thus, the B cell receives signals from both its surface antibody and the T cell via its cytokines, and acts as a professional antigen-presenting cell in the process.

[caption id="" align="aligncenter" width="480"]This diagram shows the binding of a B cell and a T cell. Figure 5. T and B Cell Binding. To elicit a response to a T cell-dependent antigen, the B and T cells must come close together. To become fully activated, the B cell must receive two signals from the native antigen and the T cell’s cytokines.[/caption] [caption id="attachment_2982" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of the adaptive immune response.[/caption]
]]> 2342 2310 5 0 1.5 Homeostasis https://pressbooks.bccampus.ca/apdouglas2017/chapter/1-5-homeostasis-2/ Mon, 17 Jul 2017 18:52:22 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2428

Learning Objectives

By the end of this section, you will be able to:
  • Discuss the role of homeostasis in healthy functioning
  • Contrast negative and positive feedback, giving one physiologic example of each mechanism

Maintaining homeostasis requires that the body continuously monitor its internal conditions. From body temperature to blood pressure to levels of certain nutrients, each physiological condition has a particular set point. A set point is the physiological value around which the normal range fluctuates. A normal range is the restricted set of values that is optimally healthful and stable. For example, the set point for normal human body temperature is approximately 37°C (98.6°F) Physiological parameters, such as body temperature and blood pressure, tend to fluctuate within a normal range a few degrees above and below that point. Control centers in the brain and other parts of the body monitor and react to deviations from homeostasis using negative feedback. Negative feedback is a mechanism that reverses a deviation from the set point. Therefore, negative feedback maintains body parameters within their normal range. The maintenance of homeostasis by negative feedback goes on throughout the body at all times, and an understanding of negative feedback is thus fundamental to an understanding of human physiology.

Negative Feedback

A negative feedback system has three basic components (Figure 1a). A sensor, also referred to a receptor, is a component of a feedback system that monitors a physiological value. This value is reported to the control center. The control center is the component in a feedback system that compares the value to the normal range. If the value deviates too much from the set point, then the control center activates an effector. An effector is the component in a feedback system that causes a change to reverse the situation and return the value to the normal range.

[caption id="" align="aligncenter" width="450"]This figure shows three flow charts labeled A, B, and C. Chart A shows a general negative feedback loop. The loop starts with a stimulus. Information about the stimulus is perceived by a sensor which sends that information to a control center. The control center sends a signal to an effector, which then feeds back to the top of the flow chart by inhibiting the stimulus. Part B shows body temperature regulation as an example of negative feedback system. Here, the stimulus is body temperature exceeding 37 degrees Celsius. The sensor is a set of nerve cells in the skin and brain and the control center is the temperature regulatory center of the brain. The effectors are sweat glands throughout the body which inhibit the rising body temperature. Figure 1. Negative Feedback Loop. In a negative feedback loop, a stimulus—a deviation from a set point—is resisted through a physiological process that returns the body to homeostasis. (a) A negative feedback loop has four basic parts. (b) Body temperature is regulated by negative feedback.[/caption]

In order to set the system in motion, a stimulus must drive a physiological parameter beyond its normal range (that is, beyond homeostasis). This stimulus is “heard” by a specific sensor. For example, in the control of blood glucose, specific endocrine cells in the pancreas detect excess glucose (the stimulus) in the bloodstream. These pancreatic beta cells respond to the increased level of blood glucose by releasing the hormone insulin into the bloodstream. The insulin signals skeletal muscle fibers, fat cells (adipocytes), and liver cells to take up the excess glucose, removing it from the bloodstream. As glucose concentration in the bloodstream drops, the decrease in concentration—the actual negative feedback—is detected by pancreatic alpha cells, and insulin release stops. This prevents blood sugar levels from continuing to drop below the normal range.

Humans have a similar temperature regulation feedback system that works by promoting either heat loss or heat gain (Figure 1b). When the brain’s temperature regulation center receives data from the sensors indicating that the body’s temperature exceeds its normal range, it stimulates a cluster of brain cells referred to as the “heat-loss center.” This stimulation has three major effects:

  • Blood vessels in the skin begin to dilate allowing more blood from the body core to flow to the surface of the skin allowing the heat to radiate into the environment.
  • As blood flow to the skin increases, sweat glands are activated to increase their output. As the sweat evaporates from the skin surface into the surrounding air, it takes heat with it.
  • The depth of respiration increases, and a person may breathe through an open mouth instead of through the nasal passageways. This further increases heat loss from the lungs.

In contrast, activation of the brain’s heat-gain center by exposure to cold reduces blood flow to the skin, and blood returning from the limbs is diverted into a network of deep veins. This arrangement traps heat closer to the body core and restricts heat loss. If heat loss is severe, the brain triggers an increase in random signals to skeletal muscles, causing them to contract and producing shivering. The muscle contractions of shivering release heat while using up ATP. The brain triggers the thyroid gland in the endocrine system to release thyroid hormone, which increases metabolic activity and heat production in cells throughout the body. The brain also signals the adrenal glands to release epinephrine (adrenaline), a hormone that causes the breakdown of glycogen into glucose, which can be used as an energy source. The breakdown of glycogen into glucose also results in increased metabolism and heat production.

[caption id="attachment_2988" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about homeostasis.[/caption]

Positive Feedback

Positive feedback intensifies a change in the body’s physiological condition rather than reversing it. A deviation from the normal range results in more change, and the system moves farther away from the normal range. Positive feedback in the body is normal only when there is a definite end point. Childbirth and the body’s response to blood loss are two examples of positive feedback loops that are normal but are activated only when needed.

Childbirth at full term is an example of a situation in which the maintenance of the existing body state is not desired. Enormous changes in the mother’s body are required to expel the baby at the end of pregnancy. And the events of childbirth, once begun, must progress rapidly to a conclusion or the life of the mother and the baby are at risk. The extreme muscular work of labor and delivery are the result of a positive feedback system (Figure 2).

[caption id="" align="aligncenter" width="380"]This diagram shows the steps of a positive feedback loop as a series of stepwise arrows looping around a diagram of an infant within the uterus of a pregnant woman. Initially the head of the baby pushes against the cervix, transmitting nerve impulses from the cervix to the brain. Next the brain stimulates the pituitary gland to secrete oxytocin which is carried in the bloodstream to the uterus. Finally, the oxytocin simulates uterine contractions and pushes the baby harder into the cervix. As the head of the baby pushes against the cervix with greater and greater force, the uterine contractions grow stronger and more frequent. This mechanism is a positive feedback loop. Figure 2. Positive Feedback Loop. Normal childbirth is driven by a positive feedback loop. A positive feedback loop results in a change in the body’s status, rather than a return to homeostasis.[/caption]
The first contractions of labor (the stimulus) push the baby toward the cervix (the lowest part of the uterus). The cervix contains stretch-sensitive nerve cells that monitor the degree of stretching (the sensors). These nerve cells send messages to the brain, which in turn causes the pituitary gland at the base of the brain to release the hormone oxytocin into the bloodstream. Oxytocin causes stronger contractions of the smooth muscles in of the uterus (the effectors), pushing the baby further down the birth canal. This causes even greater stretching of the cervix. The cycle of stretching, oxytocin release, and increasingly more forceful contractions stops only when the baby is born. At this point, the stretching of the cervix halts, stopping the release of oxytocin.

A second example of positive feedback centers on reversing extreme damage to the body. Following a penetrating wound, the most immediate threat is excessive blood loss. Less blood circulating means reduced blood pressure and reduced perfusion (penetration of blood) to the brain and other vital organs. If perfusion is severely reduced, vital organs will shut down and the person will die. The body responds to this potential catastrophe by releasing substances in the injured blood vessel wall that begin the process of blood clotting. As each step of clotting occurs, it stimulates the release of more clotting substances. This accelerates the processes of clotting and sealing off the damaged area. Clotting is contained in a local area based on the tightly controlled availability of clotting proteins. This is an adaptive, life-saving cascade of events.

]]> 2428 2420 3 0 2.2 Chemical Bonds https://pressbooks.bccampus.ca/apdouglas2017/chapter/2-2-chemical-bonds-2/ Mon, 17 Jul 2017 18:54:22 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2442

Learning Objectives

By the end of this section, you will be able to:
  • Explain the relationship between molecules and compounds
  • Distinguish between ions, cations, and anions
  • Identify the key difference between ionic and covalent bonds
  • Distinguish between nonpolar and polar covalent bonds
  • Explain how water molecules link via hydrogen bonds

Atoms separated by a great distance cannot link; rather, they must come close enough for the electrons in their valence shells to interact. But do atoms ever actually touch one another? Most physicists would say no, because the negatively charged electrons in their valence shells repel one another. No force within the human body—or anywhere in the natural world—is strong enough to overcome this electrical repulsion. So when you read about atoms linking together or colliding, bear in mind that the atoms are not merging in a physical sense.

Instead, atoms link by forming a chemical bond. A bond is a weak or strong electrical attraction that holds atoms in the same vicinity. The new grouping is typically more stable—less likely to react again—than its component atoms were when they were separate. A more or less stable grouping of two or more atoms held together by chemical bonds is called a molecule. The bonded atoms may be of the same element, as in the case of H2, which is called molecular hydrogen or hydrogen gas. When a molecule is made up of two or more atoms of different elements, it is called a chemical compound. Thus, a unit of water, or H2O, is a compound, as is a single molecule of the gas methane, or CH4.

Three types of chemical bonds are important in human physiology, because they hold together substances that are used by the body for critical aspects of homeostasis, signaling, and energy production, to name just a few important processes. These are ionic bonds, covalent bonds, and hydrogen bonds.

Ions and Ionic Bonds

Recall that an atom typically has the same number of positively charged protons and negatively charged electrons. As long as this situation remains, the atom is electrically neutral. But when an atom participates in a chemical reaction that results in the donation or acceptance of one or more electrons, the atom will then become positively or negatively charged. This happens frequently for most atoms in order to have a full valence shell, as described previously. This can happen either by gaining electrons to fill a shell that is more than half-full, or by giving away electrons to empty a shell than is less than half-full, thereby leaving the next smaller electron shell as the new, full, valence shell. An atom that has an electrical charge—whether positive or negative—is an ion.

[caption id="attachment_2990" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about electric charges.[/caption]

Potassium (K), for instance, is an important element in all body cells. Its atomic number is 19. It has just one electron in its valence shell. This characteristic makes potassium highly likely to participate in chemical reactions in which it donates one electron. (It is easier for potassium to donate one electron than to gain seven electrons.) The loss will cause the positive charge of potassium’s protons to be more influential than the negative charge of potassium’s electrons. In other words, the resulting potassium ion will be slightly positive. A potassium ion is written K+, indicating that it has lost a single electron. A positively charged ion is known as a cation.

Now consider fluorine (F), a component of bones and teeth. Its atomic number is nine, and it has seven electrons in its valence shell. Thus, it is highly likely to bond with other atoms in such a way that fluorine accepts one electron (it is easier for fluorine to gain one electron than to donate seven electrons). When it does, its electrons will outnumber its protons by one, and it will have an overall negative charge. The ionized form of fluorine is called fluoride, and is written as F–. A negatively charged ion is known as an anion.

Atoms that have more than one electron to donate or accept will end up with stronger positive or negative charges. A cation that has donated two electrons has a net charge of +2. Using magnesium (Mg) as an example, this can be written Mg++ or Mg2+. An anion that has accepted two electrons has a net charge of –2. The ionic form of selenium (Se), for example, is typically written Se2–.

The opposite charges of cations and anions exert a moderately strong mutual attraction that keeps the atoms in close proximity forming an ionic bond. An ionic bond is an ongoing, close association between ions of opposite charge. The table salt you sprinkle on your food owes its existence to ionic bonding. As shown in Figure 1, sodium commonly donates an electron to chlorine, becoming the cation Na+. When chlorine accepts the electron, it becomes the chloride anion, Cl–. With their opposing charges, these two ions strongly attract each other.

[caption id="" align="aligncenter" width="420"]The top panel of this figure shows the orbit model of a sodium atom and a chlorine atom and arrows pointing towards the transfer of electrons from sodium to chlorine to form sodium and chlorine ions. The bottom panel shows sodium and chloride ions in a crystal structure. Figure 1. Ionic Bonding. (a) Sodium readily donates the solitary electron in its valence shell to chlorine, which needs only one electron to have a full valence shell. (b) The opposite electrical charges of the resulting sodium cation and chloride anion result in the formation of a bond of attraction called an ionic bond. (c) The attraction of many sodium and chloride ions results in the formation of large groupings called crystals.[/caption]

Water is an essential component of life because it is able to break the ionic bonds in salts to free the ions. In fact, in biological fluids, most individual atoms exist as ions. These dissolved ions produce electrical charges within the body. The behavior of these ions produces the tracings of heart and brain function observed as waves on an electrocardiogram (EKG or ECG) or an electroencephalogram (EEG). The electrical activity that derives from the interactions of the charged ions is why they are also called electrolytes.

Covalent Bonds

Unlike ionic bonds formed by the attraction between a cation’s positive charge and an anion’s negative charge, molecules formed by a covalent bond share electrons in a mutually stabilizing relationship. Like next-door neighbors whose kids hang out first at one home and then at the other, the atoms do not lose or gain electrons permanently. Instead, the electrons move back and forth between the elements. Because of the close sharing of pairs of electrons (one electron from each of two atoms), covalent bonds are stronger than ionic bonds.

Nonpolar Covalent Bonds

Figure 2 shows several common types of covalent bonds. Notice that the two covalently bonded atoms typically share just one or two electron pairs, though larger sharings are possible. The important concept to take from this is that in covalent bonds, electrons in the outermost valence shell are shared to fill the valence shells of both atoms, ultimately stabilizing both of the atoms involved. In a single covalent bond, a single electron is shared between two atoms, while in a double covalent bond, two pairs of electrons are shared between two atoms. There even are triple covalent bonds, where three atoms are shared.
[caption id="" align="aligncenter" width="550"]The top panel in this figure shows two hydrogen atoms sharing two electrons. The middle panel shows two oxygen atoms sharing four electrons, and the bottom panel shows two oxygen atoms and one carbon atom sharing 2 pairs of electrons each. Figure 2. Covalent Bonding.[/caption]
You can see that the covalent bonds shown in Figure 2 are balanced. The sharing of the negative electrons is relatively equal, as is the electrical pull of the positive protons in the nucleus of the atoms involved. This is why covalently bonded molecules that are electrically balanced in this way are described as nonpolar; that is, no region of the molecule is either more positive or more negative than any other.

Polar Covalent Bonds

Groups of legislators with completely opposite views on a particular issue are often described as “polarized” by news writers. In chemistry, a polar molecule is a molecule that contains regions that have opposite electrical charges. Polar molecules occur when atoms share electrons unequally, in polar covalent bonds.

The most familiar example of a polar molecule is water (Figure 3). The molecule has three parts: one atom of oxygen, the nucleus of which contains eight protons, and two hydrogen atoms, whose nuclei each contain only one proton. Because every proton exerts an identical positive charge, a nucleus that contains eight protons exerts a charge eight times greater than a nucleus that contains one proton. This means that the negatively charged electrons present in the water molecule are more strongly attracted to the oxygen nucleus than to the hydrogen nuclei. Each hydrogen atom’s single negative electron therefore migrates toward the oxygen atom, making the oxygen end of their bond slightly more negative than the hydrogen end of their bond.

[caption id="" align="aligncenter" width="380"]This figure shows the structure of a water molecule. The top panel shows two oxygen atoms and one hydrogen atom with electrons in orbit and the shared electrons. The middle panel shows a three-dimensional model of a water molecule and the bottom panel shows the structural formula for water. Figure 3. Polar Covalent Bonds in a Water Molecule.[/caption]

What is true for the bonds is true for the water molecule as a whole; that is, the oxygen region has a slightly negative charge and the regions of the hydrogen atoms have a slightly positive charge. These charges are often referred to as “partial charges” because the strength of the charge is less than one full electron, as would occur in an ionic bond. As shown in Figure 3, regions of weak polarity are indicated with the Greek letter delta (∂) and a plus (+) or minus (–) sign.

Even though a single water molecule is unimaginably tiny, it has mass, and the opposing electrical charges on the molecule pull that mass in such a way that it creates a shape somewhat like a triangular tent (see Figure 3b). This dipole, with the positive charges at one end formed by the hydrogen atoms at the “bottom” of the tent and the negative charge at the opposite end (the oxygen atom at the “top” of the tent) makes the charged regions highly likely to interact with charged regions of other polar molecules. For human physiology, the resulting bond is one of the most important formed by water—the hydrogen bond.

Hydrogen Bonds

A hydrogen bond is formed when a weakly positive hydrogen atom already bonded to one electronegative atom (for example, the oxygen in the water molecule) is attracted to another electronegative atom from another molecule. In other words, hydrogen bonds always include hydrogen that is already part of a polar molecule.

The most common example of hydrogen bonding in the natural world occurs between molecules of water. It happens before your eyes whenever two raindrops merge into a larger bead, or a creek spills into a river. Hydrogen bonding occurs because the weakly negative oxygen atom in one water molecule is attracted to the weakly positive hydrogen atoms of two other water molecules (Figure 4).

[caption id="" align="aligncenter" width="280"]This figure shows three water molecules and the hydrogen bonds between them. Figure 4. Hydrogen Bonds between Water Molecules. Notice that the bonds occur between the weakly positive charge on the hydrogen atoms and the weakly negative charge on the oxygen atoms. Hydrogen bonds are relatively weak, and therefore are indicated with a dotted (rather than a solid) line.[/caption]

Water molecules also strongly attract other types of charged molecules as well as ions. This explains why “table salt,” for example, actually is a molecule called a “salt” in chemistry, which consists of equal numbers of positively-charged sodium (Na+) and negatively-charged chloride (Cl–), dissolves so readily in water, in this case forming dipole-ion bonds between the water and the electrically-charged ions (electrolytes). Water molecules also repel molecules with nonpolar covalent bonds, like fats, lipids, and oils. You can demonstrate this with a simple kitchen experiment: pour a teaspoon of vegetable oil, a compound formed by nonpolar covalent bonds, into a glass of water. Instead of instantly dissolving in the water, the oil forms a distinct bead because the polar water molecules repel the nonpolar oil.

]]> 2442 2429 2 0 3.5 Cell Growth and Division https://pressbooks.bccampus.ca/apdouglas2017/chapter/3-5-cell-growth-and-division-2/ Mon, 17 Jul 2017 18:56:47 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2466

Learning Objectives

By the end of this section, you will be able to:
  • Describe the stages of the cell cycle
  • Discuss how the cell cycle is regulated
  • Describe the implications of losing control over the cell cycle
  • Describe the stages of mitosis and cytokinesis, in order

So far in this chapter, you have read numerous times of the importance and prevalence of cell division. While there are a few cells in the body that do not undergo cell division (such as gametes, red blood cells, most neurons, and some muscle cells), most somatic cells divide regularly. A somatic cell is a general term for a body cell, and all human cells, except for the cells that produce eggs and sperm (which are referred to as germ cells), are somatic cells. Somatic cells contain two copies of each of their chromosomes (one copy received from each parent). A homologous pair of chromosomes is the two copies of a single chromosome found in each somatic cell. The human is a diploid organism, having 23 homologous pairs of chromosomes in each of the somatic cells. The condition of having pairs of chromosomes is known as diploidy.

Cells in the body replace themselves over the lifetime of a person. For example, the cells lining the gastrointestinal tract must be frequently replaced when constantly “worn off” by the movement of food through the gut. But what triggers a cell to divide, and how does it prepare for and complete cell division? The cell cycle is the sequence of events in the life of the cell from the moment it is created at the end of a previous cycle of cell division until it then divides itself, generating two new cells.

The Cell Cycle

One “turn” or cycle of the cell cycle consists of two general phases: interphase, followed by mitosis and cytokinesis. Interphase is the period of the cell cycle during which the cell is not dividing. The majority of cells are in interphase most of the time. Mitosis is the division of genetic material, during which the cell nucleus breaks down and two new, fully functional, nuclei are formed. Cytokinesis divides the cytoplasm into two distinctive cells.

Interphase

A cell grows and carries out all normal metabolic functions and processes in a period called G1 (Figure 1). G1 phase (gap 1 phase) is the first gap, or growth phase in the cell cycle. For cells that will divide again, G1 is followed by replication of the DNA, during the S phase. The S phase (synthesis phase) is period during which a cell replicates its DNA.

[caption id="" align="aligncenter" width="280"]This figure shows the different stages of the cell cycle. The G0 phase where the cells are not actively dividing is also labeled. Figure 1. Cell Cycle. The two major phases of the cell cycle include mitosis (cell division), and interphase, when the cell grows and performs all of its normal functions. Interphase is further subdivided into G1, S, and G2 phases.[/caption]

After the synthesis phase, the cell proceeds through the G2 phase. The G2 phase is a second gap phase, during which the cell continues to grow and makes the necessary preparations for mitosis. Between G1, S, and G2 phases, cells will vary the most in their duration of the G1 phase. It is here that a cell might spend a couple of hours, or many days. The S phase typically lasts between 8-10 hours and the G2 phase approximately 5 hours. In contrast to these phases, the G0 phase is a resting phase of the cell cycle. Cells that have temporarily stopped dividing and are resting (a common condition) and cells that have permanently ceased dividing (like nerve cells) are said to be in G0.

The Structure of Chromosomes

Billions of cells in the human body divide every day. During the synthesis phase (S, for DNA synthesis) of interphase, the amount of DNA within the cell precisely doubles. Therefore, after DNA replication but before cell division, each cell actually contains two copies of each chromosome. Each copy of the chromosome is referred to as a sister chromatid and is physically bound to the other copy. The centromere is the structure that attaches one sister chromatid to another. Because a human cell has 46 chromosomes, during this phase, there are 92 chromatids (46 × 2) in the cell. Make sure not to confuse the concept of a pair of chromatids (one chromosome and its exact copy attached during mitosis) and a homologous pair of chromosomes (two paired chromosomes which were inherited separately, one from each parent) (Figure 2).

[caption id="" align="aligncenter" width="250"]This image shows a pair of chromosomes. The major parts such as the homologous chromosomes, kinetochore and the sister chromatids are labeled. Figure 2. A Homologous Pair of Chromosomes with their Attached Sister Chromatids. The red and blue colors correspond to a homologous pair of chromosomes. Each member of the pair was separately inherited from one parent. Each chromosome in the homologous pair is also bound to an identical sister chromatid, which is produced by DNA replication, and results in the familiar “X” shape.[/caption]

Mitosis and Cytokinesis

The mitotic phase of the cell typically takes between 1 and 2 hours. During this phase, a cell undergoes two major processes. First, it completes mitosis, during which the contents of the nucleus are equitably pulled apart and distributed between its two halves. Cytokinesis then occurs, dividing the cytoplasm and cell body into two new cells. Mitosis is divided into four major stages that take place after interphase (Figure 3) and in the following order: prophase, metaphase, anaphase, and telophase. The process is then followed by cytokinesis.

[caption id="attachment_1512" align="aligncenter" width="600"]This tabular image shows the different stages of mitosis and cytokinesis using both drawings and text. The top panel is a series of schematics for each step, followed by text listing the important aspects of that step. The bottom panel shows fluorescent micrographs for the corresponding stage. Figure 3. Cell Division: Mitosis Followed by Cytokinesis. The stages of cell division oversee the separation of identical genetic material into two new nuclei, followed by the division of the cytoplasm.[/caption]
Prophase is the first phase of mitosis, during which the loosely packed chromatin coils and condenses into visible chromosomes. During prophase, each chromosome becomes visible with its identical partner attached, forming the familiar X-shape of sister chromatids. The nucleolus disappears early during this phase, and the nuclear envelope also disintegrates.A major occurrence during prophase concerns a very important structure that contains the origin site for microtubule growth. Recall the cellular structures called centrioles that serve as origin points from which microtubules extend. These tiny structures also play a very important role during mitosis. A centrosome is a pair of centrioles together. The cell contains two centrosomes side-by-side, which begin to move apart during prophase. As the centrosomes migrate to two different sides of the cell, microtubules begin to extend from each like long fingers from two hands extending toward each other. The mitotic spindle is the structure composed of the centrosomes and their emerging microtubules.

Near the end of prophase there is an invasion of the nuclear area by microtubules from the mitotic spindle. The nuclear membrane has disintegrated, and the microtubules attach themselves to the centromeres that adjoin pairs of sister chromatids. The kinetochore is a protein structure on the centromere that is the point of attachment between the mitotic spindle and the sister chromatids. This stage is referred to as late prophase or “prometaphase” to indicate the transition between prophase and metaphase.

Metaphase is the second stage of mitosis. During this stage, the sister chromatids, with their attached microtubules, line up along a linear plane in the middle of the cell. A metaphase plate forms between the centrosomes that are now located at either end of the cell. The metaphase plate is the name for the plane through the center of the spindle on which the sister chromatids are positioned. The microtubules are now poised to pull apart the sister chromatids and bring one from each pair to each side of the cell.

Anaphase is the third stage of mitosis. Anaphase takes place over a few minutes, when the pairs of sister chromatids are separated from one another, forming individual chromosomes once again. These chromosomes are pulled to opposite ends of the cell by their kinetochores, as the microtubules shorten. Each end of the cell receives one partner from each pair of sister chromatids, ensuring that the two new daughter cells will contain identical genetic material.

Telophase is the final stage of mitosis. Telophase is characterized by the formation of two new daughter nuclei at either end of the dividing cell. These newly formed nuclei surround the genetic material, which uncoils such that the chromosomes return to loosely packed chromatin. Nucleoli also reappear within the new nuclei, and the mitotic spindle breaks apart, each new cell receiving its own complement of DNA, organelles, membranes, and centrioles. At this point, the cell is already beginning to split in half as cytokinesis begins.

The cleavage furrow is a contractile band made up of microfilaments that forms around the midline of the cell during cytokinesis. (Recall that microfilaments consist of actin.) This contractile band squeezes the two cells apart until they finally separate. Two new cells are now formed. One of these cells (the “stem cell”) enters its own cell cycle; able to grow and divide again at some future time. The other cell transforms into the functional cell of the tissue, typically replacing an “old” cell there.

Imagine a cell that completed mitosis but never underwent cytokinesis. In some cases, a cell may divide its genetic material and grow in size, but fail to undergo cytokinesis. This results in larger cells with more than one nucleus. Usually this is an unwanted aberration and can be a sign of cancerous cells.

Cell Cycle Control

A very elaborate and precise system of regulation controls direct the way cells proceed from one phase to the next in the cell cycle and begin mitosis. The control system involves molecules within the cell as well as external triggers. These internal and external control triggers provide “stop” and “advance” signals for the cell. Precise regulation of the cell cycle is critical for maintaining the health of an organism, and loss of cell cycle control can lead to cancer.

Mechanisms of Cell Cycle Control

As the cell proceeds through its cycle, each phase involves certain processes that must be completed before the cell should advance to the next phase. A checkpoint is a point in the cell cycle at which the cycle can be signaled to move forward or stopped. At each of these checkpoints, different varieties of molecules provide the stop or go signals, depending on certain conditions within the cell. A cyclin is one of the primary classes of cell cycle control molecules (Figure 4). A cyclin-dependent kinase (CDK) is one of a group of molecules that work together with cyclins to determine progression past cell checkpoints. By interacting with many additional molecules, these triggers push the cell cycle forward unless prevented from doing so by “stop” signals, if for some reason the cell is not ready. At the G1 checkpoint, the cell must be ready for DNA synthesis to occur. At the G2 checkpoint the cell must be fully prepared for mitosis. Even during mitosis, a crucial stop and go checkpoint in metaphase ensures that the cell is fully prepared to complete cell division. The metaphase checkpoint ensures that all sister chromatids are properly attached to their respective microtubules and lined up at the metaphase plate before the signal is given to separate them during anaphase.

[caption id="" align="aligncenter" width="350"]This image shows the different stages of the cell cycle along with the checkpoints between them and the cyclins responsible for the checkpoint at each stage. Figure 4. Control of the Cell Cycle. Cells proceed through the cell cycle under the control of a variety of molecules, such as cyclins and cyclin-dependent kinases. These control molecules determine whether or not the cell is prepared to move into the following stage.[/caption]

The Cell Cycle Out of Control: Implications

Most people understand that cancer or tumors are caused by abnormal cells that multiply continuously. If the abnormal cells continue to divide unstopped, they can damage the tissues around them, spread to other parts of the body, and eventually result in death. In healthy cells, the tight regulation mechanisms of the cell cycle prevent this from happening, while failures of cell cycle control can cause unwanted and excessive cell division. Failures of control may be caused by inherited genetic abnormalities that compromise the function of certain “stop” and “go” signals. Environmental insult that damages DNA can also cause dysfunction in those signals. Often, a combination of both genetic predisposition and environmental factors lead to cancer.

The process of a cell escaping its normal control system and becoming cancerous may actually happen throughout the body quite frequently. Fortunately, certain cells of the immune system are capable of recognizing cells that have become cancerous and destroying them. However, in certain cases the cancerous cells remain undetected and continue to proliferate. If the resulting tumor does not pose a threat to surrounding tissues, it is said to be benign and can usually be easily removed. If capable of damage, the tumor is considered malignant and the patient is diagnosed with cancer.
Homeostatic Imbalances

Cancer Arises from Homeostatic Imbalances Cancer is an extremely complex condition, capable of arising from a wide variety of genetic and environmental causes. Typically, mutations or aberrations in a cell’s DNA that compromise normal cell cycle control systems lead to cancerous tumors. Cell cycle control is an example of a homeostatic mechanism that maintains proper cell function and health. While progressing through the phases of the cell cycle, a large variety of intracellular molecules provide stop and go signals to regulate movement forward to the next phase. These signals are maintained in an intricate balance so that the cell only proceeds to the next phase when it is ready. This homeostatic control of the cell cycle can be thought of like a car’s cruise control. Cruise control will continually apply just the right amount of acceleration to maintain a desired speed, unless the driver hits the brakes, in which case the car will slow down. Similarly, the cell includes molecular messengers, such as cyclins, that push the cell forward in its cycle.

In addition to cyclins, a class of proteins that are encoded by genes called proto-oncogenes provide important signals that regulate the cell cycle and move it forward. Examples of proto-oncogene products include cell-surface receptors for growth factors, or cell-signaling molecules, two classes of molecules that can promote DNA replication and cell division. In contrast, a second class of genes known as tumor suppressor genes sends stop signals during a cell cycle. For example, certain protein products of tumor suppressor genes signal potential problems with the DNA and thus stop the cell from dividing, while other proteins signal the cell to die if it is damaged beyond repair. Some tumor suppressor proteins also signal a sufficient surrounding cellular density, which indicates that the cell need not presently divide. The latter function is uniquely important in preventing tumor growth: normal cells exhibit a phenomenon called “contact inhibition;” thus, extensive cellular contact with neighboring cells causes a signal that stops further cell division.

These two contrasting classes of genes, proto-oncogenes and tumor suppressor genes, are like the accelerator and brake pedal of the cell’s own “cruise control system,” respectively. Under normal conditions, these stop and go signals are maintained in a homeostatic balance. Generally speaking, there are two ways that the cell’s cruise control can lose control: a malfunctioning (overactive) accelerator, or a malfunctioning (underactive) brake. When compromised through a mutation, or otherwise altered, proto-oncogenes can be converted to oncogenes, which produce oncoproteins that push a cell forward in its cycle and stimulate cell division even when it is undesirable to do so. For example, a cell that should be programmed to self-destruct (a process called apoptosis) due to extensive DNA damage might instead be triggered to proliferate by an oncoprotein. On the other hand, a dysfunctional tumor suppressor gene may fail to provide the cell with a necessary stop signal, also resulting in unwanted cell division and proliferation.

A delicate homeostatic balance between the many proto-oncogenes and tumor suppressor genes delicately controls the cell cycle and ensures that only healthy cells replicate. Therefore, a disruption of this homeostatic balance can cause aberrant cell division and cancerous growths.

[caption id="attachment_2992" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about the process of mitosis![/caption] [caption id="attachment_3039" align="aligncenter" width="150"] Watch this amoeba sisters video to learn more about mitosis![/caption] [caption id="attachment_3040" align="aligncenter" width="150"] Watch this amoeba sisters video to learn about the process of meiosis![/caption]
]]> 2466 2462 1 0 4.2 Epithelial Tissue https://pressbooks.bccampus.ca/apdouglas2017/chapter/4-2-epithelial-tissue-2/ Mon, 17 Jul 2017 18:57:53 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2476

Learning Objectives

By the end of this section, you will be able to:
  • Explain the structure and function of epithelial tissue
  • Distinguish between tight junctions, anchoring junctions, and gap junctions
  • Distinguish between simple epithelia and stratified epithelia, as well as between squamous, cuboidal, and columnar epithelia
  • Describe the structure and function of endocrine and exocrine glands and their respective secretions

Most epithelial tissues are essentially large sheets of cells covering all the surfaces of the body exposed to the outside world and lining the outside of organs. Epithelium also forms much of the glandular tissue of the body. Skin is not the only area of the body exposed to the outside. Other areas include the airways, the digestive tract, as well as the urinary and reproductive systems, all of which are lined by an epithelium. Hollow organs and body cavities that do not connect to the exterior of the body, which includes, blood vessels and serous membranes, are lined by endothelium (plural = endothelia), which is a type of epithelium.

Epithelial cells derive from all three major embryonic layers. The epithelia lining the skin, parts of the mouth and nose, and the anus develop from the ectoderm. Cells lining the airways and most of the digestive system originate in the endoderm. The epithelium that lines vessels in the lymphatic and cardiovascular system derives from the mesoderm and is called an endothelium.

All epithelia share some important structural and functional features. This tissue is highly cellular, with little or no extracellular material present between cells. Adjoining cells form a specialized intercellular connection between their cell membranes called a cell junction. The epithelial cells exhibit polarity with differences in structure and function between the exposed or apical facing surface of the cell and the basal surface close to the underlying body structures. The basal lamina, a mixture of glycoproteins and collagen, provides an attachment site for the epithelium, separating it from underlying connective tissue. The basal lamina attaches to a reticular lamina, which is secreted by the underlying connective tissue, forming a basement membrane that helps hold it all together.

Epithelial tissues are nearly completely avascular. For instance, no blood vessels cross the basement membrane to enter the tissue, and nutrients must come by diffusion or absorption from underlying tissues or the surface. Many epithelial tissues are capable of rapidly replacing damaged and dead cells. Sloughing off of damaged or dead cells is a characteristic of surface epithelium and allows our airways and digestive tracts to rapidly replace damaged cells with new cells.

Generalized Functions of Epithelial Tissue

Epithelial tissues provide the body’s first line of protection from physical, chemical, and biological wear and tear. The cells of an epithelium act as gatekeepers of the body controlling permeability and allowing selective transfer of materials across a physical barrier. All substances that enter the body must cross an epithelium. Some epithelia often include structural features that allow the selective transport of molecules and ions across their cell membranes.

Many epithelial cells are capable of secretion and release mucous and specific chemical compounds onto their apical surfaces. The epithelium of the small intestine releases digestive enzymes, for example. Cells lining the respiratory tract secrete mucous that traps incoming microorganisms and particles. A glandular epithelium contains many secretory cells.

The Epithelial Cell

Epithelial cells are typically characterized by the polarized distribution of organelles and membrane-bound proteins between their basal and apical surfaces. Particular structures found in some epithelial cells are an adaptation to specific functions. Certain organelles are segregated to the basal sides, whereas other organelles and extensions, such as cilia, when present, are on the apical surface.

Cilia are microscopic extensions of the apical cell membrane that are supported by microtubules. They beat in unison and move fluids as well as trapped particles. Ciliated epithelium lines the ventricles of the brain where it helps circulate the cerebrospinal fluid. The ciliated epithelium of your airway forms a mucociliary escalator that sweeps particles of dust and pathogens trapped in the secreted mucous toward the throat. It is called an escalator because it continuously pushes mucous with trapped particles upward. In contrast, nasal cilia sweep the mucous blanket down towards your throat. In both cases, the transported materials are usually swallowed, and end up in the acidic environment of your stomach.

Cell to Cell Junctions

Cells of epithelia are closely connected and are not separated by intracellular material. Three basic types of connections allow varying degrees of interaction between the cells: tight junctions, anchoring junctions, and gap junctions (Figure 1).

[caption id="" align="aligncenter" width="500"]These three illustrations each show the edges of two vertical cell membranes. The cell membranes are viewed partially from the side so that the inside edge of the right cell membrane is visible. The upper left image shows a tight junction. The two cell membranes are bound by transmembrane protein strands. The proteins travel the inside edge of the right cell membrane and cross over to the left cell membrane, cinching the two membranes together. The cell membranes are still somewhat separated in between neighboring strands, creating intercellular spaces. The upper right diagram shows a gap junction. The gap junctions are composed of two interlocking connexins, which are round, hollow tubes that extend through the cell membranes. Two connexins, one from the left cell membrane and the other from the right cell membrane, meet between the two cells, forming a connexon. Even at the site of the connexon, there is a small gap between the cell membranes. On the inside edge of the right cell membrane, the gap junction appears as a depression. Three connexins are embedded into the membranes like buttons on a shirt. The bottom images show the three types of anchoring junctions. The left image shows a desmosome. Here, the inside edge of both the right and left cell membranes have brown, round plaques. Each plaque has tentacle-like intermediate filaments (keratin) that extend into each cell’s cytoplasm. The two plaques are connected across the intercellular space by several interlocking transmembrane glycoproteins (cadherin). The connected glycoproteins look similar to a zipped-up zipper between the right and left cell membranes. The right image shows an adheren. These are similar to desmosomes, with two plaques on the inside edge of each cell membrane connected across the intercellular space by glycoproteins. However, the plaques do not contain the tentacle-like intermediate filaments branching into the cytoplasm. Instead, the plaques are ribbed with green actin filaments. The filaments are neatly arranged in parallel, horizontal strands on the surface of the plaque facing the cytoplasm. The bottom image shows a hemidesmosome. Rather than located between two neighboring cells, the hemidesmosome is located between the bottom of a cell and the basement membrane. A hemidesmosome contains a single plaque on the inside edge of the cell membrane. Like the desmosome, intermediate filaments project from the plaque into the cytoplasm. The opposite side of the plaque has purple, knob-shaped integrins extending out to the basal lamina of the basement membrane. Figure 1. Types of Cell Junctions. The three basic types of cell-to-cell junctions are tight junctions, gap junctions, and anchoring junctions.[/caption]

At one end of the spectrum is the tight junction, which separates the cells into apical and basal compartments. An anchoring junction includes several types of cell junctions that help stabilize epithelial tissues. Anchoring junctions are common on the lateral and basal surfaces of cells where they provide strong and flexible connections. There are three types of anchoring junctions: desmosomes, hemidesmosomes, and adherens. Desmosomes occur in patches on the membranes of cells. The patches are structural proteins on the inner surface of the cell’s membrane. The adhesion molecule, cadherin, is embedded in these patches and projects through the cell membrane to link with the cadherin molecules of adjacent cells. These connections are especially important in holding cells together. Hemidesmosomes, which look like half a desmosome, link cells to the extracellular matrix, for example, the basal lamina. While similar in appearance to desmosomes, they include the adhesion proteins called integrins rather than cadherins. Adherens junctions use either cadherins or integrins depending on whether they are linking to other cells or matrix. The junctions are characterized by the presence of the contractile protein actin located on the cytoplasmic surface of the cell membrane. The actin can connect isolated patches or form a belt-like structure inside the cell. These junctions influence the shape and folding of the epithelial tissue.

In contrast with the tight and anchoring junctions, a gap junction forms an intercellular passageway between the membranes of adjacent cells to facilitate the movement of small molecules and ions between the cytoplasm of adjacent cells. These junctions allow electrical and metabolic coupling of adjacent cells, which coordinates function in large groups of cells.

Classification of Epithelial Tissues

Epithelial tissues are classified according to the shape of the cells and number of the cell layers formed (Figure 2). Cell shapes can be squamous (flattened and thin), cuboidal (boxy, as wide as it is tall), or columnar (rectangular, taller than it is wide). Similarly, the number of cell layers in the tissue can be one—where every cell rests on the basal lamina—which is a simple epithelium, or more than one, which is a stratified epithelium and only the basal layer of cells rests on the basal lamina. Pseudostratified (pseudo- = “false”) describes tissue with a single layer of irregularly shaped cells that give the appearance of more than one layer. Transitional describes a form of specialized stratified epithelium in which the shape of the cells can vary.

[caption id="" align="aligncenter" width="520"]This figure is a table showing the appearance of squamous, cuboidal and columnar epithelial tissues. Simple and compound forms are shown for each tissue type. In a simple squamous epithelium, the cells are flattened and single layered. In a simple cuboidal epithelium, the cells are cube shaped and single layered. In a simple columnar epithelium, the cells are rectangular and are attached to the basement membrane on one of their narrow sides, so that each cell is standing up like a column. There is only one layer of cells. In a pseudostratified columnar epithelium, the cells are column-like in appearance, but they vary in height. The taller cells bend over the tops of the shorter cells so that the top of the epithelial tissue is continuous. There is only one layer of cells. A stratified squamous epithelium contains many layers of flattened cells. Stratified cuboidal epithelium contains many layers of cube-shaped cells. Stratified columnar epithelium contains many layers of rectangular, column-shaped cells. Figure 2. Cells of Epithelial Tissue. Simple epithelial tissue is organized as a single layer of cells and stratified epithelial tissue is formed by several layers of cells.[/caption]

Simple Epithelium

The shape of the cells in the single cell layer of simple epithelium reflects the functioning of those cells. The cells in simple squamous epithelium have the appearance of thin scales. Squamous cell nuclei tend to be flat, horizontal, and elliptical, mirroring the form of the cell. The endothelium is the epithelial tissue that lines vessels of the lymphatic and cardiovascular system, and it is made up of a single layer of squamous cells. Simple squamous epithelium, because of the thinness of the cell, is present where rapid passage of chemical compounds is observed. The alveoli of lungs where gases diffuse, segments of kidney tubules, and the lining of capillaries are also made of simple squamous epithelial tissue. The mesothelium is a simple squamous epithelium that forms the surface layer of the serous membrane that lines body cavities and internal organs. Its primary function is to provide a smooth and protective surface. Mesothelial cells are squamous epithelial cells that secrete a fluid that lubricates the mesothelium.

In simple cuboidal epithelium, the nucleus of the box-like cells appears round and is generally located near the center of the cell. These epithelia are active in the secretion and absorptions of molecules. Simple cuboidal epithelia are observed in the lining of the kidney tubules and in the ducts of glands.

In simple columnar epithelium, the nucleus of the tall column-like cells tends to be elongated and located in the basal end of the cells. Like the cuboidal epithelia, this epithelium is active in the absorption and secretion of molecules. Simple columnar epithelium forms the lining of some sections of the digestive system and parts of the female reproductive tract. Ciliated columnar epithelium is composed of simple columnar epithelial cells with cilia on their apical surfaces. These epithelial cells are found in the lining of the fallopian tubes and parts of the respiratory system, where the beating of the cilia helps remove particulate matter.

Pseudostratified columnar epithelium is a type of epithelium that appears to be stratified but instead consists of a single layer of irregularly shaped and differently sized columnar cells. In pseudostratified epithelium, nuclei of neighboring cells appear at different levels rather than clustered in the basal end. The arrangement gives the appearance of stratification; but in fact all the cells are in contact with the basal lamina, although some do not reach the apical surface. Pseudostratified columnar epithelium is found in the respiratory tract, where some of these cells have cilia.

Both simple and pseudostratified columnar epithelia are heterogeneous epithelia because they include additional types of cells interspersed among the epithelial cells. For example, a goblet cell is a mucous-secreting unicellular “gland” interspersed between the columnar epithelial cells of mucous membranes (Figure 3).

[caption id="" align="aligncenter" width="250"]This illustration shows a diagram of a goblet cell. The goblet cell is shaped roughly like an upside down vase. The enlarged end at the top contains six finger like projections labeled microvilli. Between the microvilli, secretary vesicles containing mucin are moving from the upper half of the cell toward the microvilli. Below the secretory vesicles are several rough endoplasmic reticula and an irregularly shaped Golgi apparatus with secretory vesicles budding off of it. The narrow, lower half of the cell contains the oval-shaped nucleus as well as a few mitochondria and segments of the endoplasmic reticulum. Figure 3. Goblet Cell. (a) In the lining of the small intestine, columnar epithelium cells are interspersed with goblet cells. (b) The arrows in this micrograph point to the mucous-secreting goblet cells. LM × 1600. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]
The second image is a micrograph of the innermost lining of the small intestine. This innermost lining is a simple columnar epithelium, with a single layer of rectangular cells oriented in a line. Occasionally, the line of epithelial cells is interrupted by a goblet cell. Goblet cells are thinner than the epithelial cells and appear roughly pill shaped. In this micrograph, the cells did not stain as darkly as the epithelial cells.

Stratified Epithelium

A stratified epithelium consists of several stacked layers of cells. This epithelium protects against physical and chemical wear and tear. The stratified epithelium is named by the shape of the most apical layer of cells, closest to the free space. Stratified squamous epithelium is the most common type of stratified epithelium in the human body. The apical cells are squamous, whereas the basal layer contains either columnar or cuboidal cells. The top layer may be covered with dead cells filled with keratin. Mammalian skin is an example of this dry, keratinized, stratified squamous epithelium. The lining of the mouth cavity is an example of an unkeratinized, stratified squamous epithelium. Stratified cuboidal epithelium and stratified columnar epithelium can also be found in certain glands and ducts, but are uncommon in the human body.

Another kind of stratified epithelium is transitional epithelium, so-called because of the gradual changes in the shapes of the apical cells as the bladder fills with urine. It is found only in the urinary system, specifically the ureters and urinary bladder. When the bladder is empty, this epithelium is convoluted and has cuboidal apical cells with convex, umbrella shaped, apical surfaces. As the bladder fills with urine, this epithelium loses its convolutions and the apical cells transition from cuboidal to squamous. It appears thicker and more multi-layered when the bladder is empty, and more stretched out and less stratified when the bladder is full and distended. Figure 4 summarizes the different categories of epithelial cell tissue cells.
[caption id="" align="aligncenter" width="500"]This figure is a table with three columns and eight rows. The leftmost column is titled cells, and contains a drawing in each row showing how epithelial cells are arranged above a basement membrane. The middle column is titled location, while the rightmost column is titled function. In a simple squamous epithelium, the cells are flattened and single-layered. Simple squamous cells are found in the air sacs of the lungs, in the lining of the heart, blood vessels and lymphatic vessels. Their function is to allow materials to pass through by diffusion and filtration, as well as to secrete lubricating substances. In a simple cuboidal epithelium, the cells are cube shaped and single layered and located in ducts and secretory portions of small glands as well as in the kidney tubules. The function of simple cuboidal epithelium is to secrete and absorb. In a simple columnar epithelium, the cells are rectangular and are attached to the basement membrane on one of their narrow sides, so that each cell is standing up like a column. There is only one layer of cells. Simple columnar epithelium is found in ciliated tissues including the bronchi, uterine tubes, and uterus, as well as in smooth, nonciliated tissues such as the digestive tract bladder. The function of simple columnar epithelium is to absorb substances but also to secrete mucous and enzymes. In a pseudostratified columnar epithelium, the cells are column-like in appearance, but they vary in height. The taller cells bend over the tops of the shorter cells so that the top of the epithelial tissue is continuous. There is only one layer of cells. Pseudostratified columnar epithelium lines the trachea and much of the upper respiratory tract. The function of pseudostratified columnar epithelium is to secrete mucous and also move that mucus using the hair like cilia projecting from the top of each cell. A stratified squamous epithelium contains many layers of flattened cells. Stratified squamous epithelium lines the esophagus, mouth, and vagina. The function of stratified squamous epithelium is to protect against abrasion. Stratified cuboidal epithelium contains many layers of cube-shaped cells. Stratified cuboidal epithelium is found in the sweat glands, salivary glands, and mammary glands. The function of stratified cuboidal epithelium is to protect other tissues of the body. Stratified columnar epithelium contains many layers of rectangular, column-shaped cells. Stratified columnar epithelium is located in the male urethra and the ducts of some glands. The function of stratified columnar epithelium is to secrete and protect. Transitional epithelium consists of many layers of irregularly shaped cells with diverse sizes. Transitional epithelium is found lining the bladder, urethra and ureters. The function of transitional epithelium is to allow the urinary organs to expand and stretch. Figure 4. Summary of Epithelial Tissue Cells.[/caption]
[caption id="attachment_2994" align="aligncenter" width="150"] Watch this CrashCourse video to get an overview of epithelial tissue.[/caption]

Glandular Epithelium

A gland is a structure made up of one or more cells modified to synthesize and secrete chemical substances. Most glands consist of groups of epithelial cells. A gland can be classified as an endocrine gland, a ductless gland that releases secretions directly into surrounding tissues and fluids (endo- = “inside”), or an exocrine gland whose secretions leave through a duct that opens directly, or indirectly, to the external environment (exo- = “outside”).

Endocrine Glands

The secretions of endocrine glands are called hormones. Hormones are released into the interstitial fluid, diffused into the bloodstream, and delivered to targets, in other words, cells that have receptors to bind the hormones. The endocrine system is part of a major regulatory system coordinating the regulation and integration of body responses. A few examples of endocrine glands include the anterior pituitary, thymus, adrenal cortex, and gonads.

Exocrine Glands

Exocrine glands release their contents through a duct that leads to the epithelial surface. Mucous, sweat, saliva, and breast milk are all examples of secretions from exocrine glands. They are all discharged through tubular ducts. Secretions into the lumen of the gastrointestinal tract, technically outside of the body, are of the exocrine category.

Glandular Structure

Exocrine glands are classified as either unicellular or multicellular. The unicellular glands are scattered single cells, such as goblet cells, found in the mucous membranes of the small and large intestine.

The multicellular exocrine glands known as serous glands develop from simple epithelium to form a secretory surface that secretes directly into an inner cavity. These glands line the internal cavities of the abdomen and chest and release their secretions directly into the cavities. Other multicellular exocrine glands release their contents through a tubular duct. The duct is single in a simple gland but in compound glands is divided into one or more branches (Figure 5). In tubular glands, the ducts can be straight or coiled, whereas tubes that form pockets are alveolar (acinar), such as the exocrine portion of the pancreas. Combinations of tubes and pockets are known as tubuloalveolar (tubuloacinar) compound glands. In a branched gland, a duct is connected to more than one secretory group of cells.

[caption id="" align="aligncenter" width="550"]This table shows the different types of exocrine glands: alveolar (acinar) versus tubular and those with simple ducts versus compound ducts. Each diagram shows a single layer of columnar epithelial cells with a line of cells travelling along the surface of a tissue (surface epithelium) and then dipping into a hole in the tissue. The cells travel down the right side of the hole until they reach the bottom, then curve around the bottom of the hole and then travel up the left side. Finally, the cells emerge back onto the surface of the tissue. The surface epithelial cells are those that are on the surface of the tissue; the duct cells are those that line both walls of the hole. The gland cells are those that line the bottom of the hole. The shape of the hole differs in each gland. In the simple alvelolar (acinar) gland, the duct and gland cells are bulb shaped with the gland cells being the larger end of the bulb. Simple alveolar glands are not found in adults, as these represent an early developmental stage of simple, branched glands. In simple tubular glands, the duct and gland cells are U shaped. Simple tubular glands are found in the intestinal glands. In simple branched alveolar glands, the gland cells form three bulbs at the end of the duct, similar in appearance to a clover leaf. The sebaceous (oil) glands are examples of simple branched alveolar glands. In simple coiled tubular glands, the duct and gland cells form a U, however, the bottom of the U, which is all gland cells, is curved up to the right. Merocrine sweat glands are examples of simple coiled tubular glands. In simple branched tubular glands, the duct is very short and the gland cells divide into three lobes, similar in appearance to a bird’s foot. The gastric glands of the stomach and mucous glands of the esophagus, tongue and duodenum are examples of simple branched tubular glands. Among the glands with compound ducts, compound alveolar (acinar) glands have three sets of clover leaf bulbs, for a total of six bulbs. Two of the clover leaf shaped structures extend parallel to the surface epithelium in opposite directions to each other. The third clover leaf extends down into the tissue, perpendicular to the surface. The duct is cross-shaped. The mammary glands are an example of compound alveolar glands. Compound tubular glands have a similar structure to compound alveolar glands. However, instead of three cloverleaf shaped bulbs, the compound tubular gland has three bird’s foot shaped bulbs. The duct is also cross-shaped in the compound tubular gland. The mucous glands of the mouth and the bulbourethral glands of the male reproductive system are examples of compound tubular glands, which are also found in the seminiferous tubules of the testis. Compound tubuloalveolar glands are a hybrid between the compound alveolar gland and the compound tubular gland. The two sets of bulbs that run parallel to the surface are bird-foot shaped; however, the set of bulbs that runs perpendicularly below the surface is cloverleaf shaped. The salivary glands, glands of the respiratory passages and glands of the pancreas are all compound tubuloalveolar glands. Figure 5. Types of Exocrine Glands. Exocrine glands are classified by their structure.[/caption]

Methods and Types of Secretion

Exocrine glands can be classified by their mode of secretion and the nature of the substances released, as well as by the structure of the glands and shape of ducts (Figure 6). Merocrine secretion is the most common type of exocrine secretion. The secretions are enclosed in vesicles that move to the apical surface of the cell where the contents are released by exocytosis. For example, watery mucous containing the glycoprotein mucin, a lubricant that offers some pathogen protection is a merocrine secretion. The eccrine glands that produce and secrete sweat are another example.

[caption id="" align="aligncenter" width="400"]These three diagrams show the three modes of secretion. All three diagrams show three orange cells in a line with attached to a basement membrane. Each cell has a large nucleus in its lower half. The upper half of each cell contains a Golgi apparatus, which appears like an upside down jellyfish. Yellow secretory vesicles are budding from the top end of the Golgi apparatus. Each vesicle contains several orange circles, which are the secreted substance. In merocrine secretion, the secretory vesicles travel to the top edge of the cells and release the secretion from the cell by melding with the cell membrane. In apocrine secretion, the top third of the cell, which contains the secretory vesicles, pinches in at the sides and then completely disconnects above the Golgi complex. The pinched off portion of the cell is the secretion, as it contains the majority of the secretory vesicles. In holocrine secretion, the upper third of the cell, just above the Golgi complex, forms many finger like projections. Each projection contains several vesicles. The tips of the projections that contain secretory vesicles bud off from the cell. In this method of secretion, the mature cell eventually dies and becomes the secretory product. Figure 6. Modes of Glandular Secretion. (a) In merocrine secretion, the cell remains intact. (b) In apocrine secretion, the apical portion of the cell is released, as well. (c) In holocrine secretion, the cell is destroyed as it releases its product and the cell itself becomes part of the secretion.[/caption]

Apocrine secretion accumulates near the apical portion of the cell. That portion of the cell and its secretory contents pinch off from the cell and are released. The sweat glands of the armpit are classified as apocrine glands. Both merocrine and apocrine glands continue to produce and secrete their contents with little damage caused to the cell because the nucleus and golgi regions remain intact after secretion.

In contrast, the process of holocrine secretion involves the rupture and destruction of the entire gland cell. The cell accumulates its secretory products and releases them only when it bursts. New gland cells differentiate from cells in the surrounding tissue to replace those lost by secretion. The sebaceous glands that produce the oils on the skin and hair are holocrine glands/cells (Figure 7).

[caption id="" align="aligncenter" width="520"]Image A depicts a cross section of the skin layers. The surface of the skin is at the top of the diagram, with the outer layer occupying about one fifth of the cross section. The outer layer has an irregular border with the inner skin layer, which occupies the remainder of the cross section. A hair follicle is embedded within the inner layer. However, the outer layer actually invaginates into the inner layer around the outside of the follicle, completely sheathing the follicle. The follicle has a bulb at its bottom that is connected to blood vessels. The hair projects from the bulb and travels through the sheath to erupt from the skin surface. The sebaceous gland is an irregular, yellow structure attached at the midpoint of the hair shaft near the border between the inner and outer layers of skin. Its duct actually connects into the side of the hair follicle. Image B shows a micrograph of a sebaceous gland connected to a hair follicle. The bulb of the hair follicle is evident in the micrograph as a bundle of cell surrounding the growing hair at its center. The sebaceous gland is connected to the right of the follicle bulb. The gland appears as an oval shaped mass of pink staining, cube shaped cells with purple nuclei. Figure 7. Sebaceous Glands. These glands secrete oils that lubricate and protect the skin. They are holocrine glands and they are destroyed after releasing their contents. New glandular cells form to replace the cells that are lost. LM × 400. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Glands are also named after the products they produce. The serous gland produces watery, blood-plasma-like secretions rich in enzymes such as alpha amylase, whereas the mucous gland releases watery to viscous products rich in the glycoprotein mucin. Both serous and mucous glands are common in the salivary glands of the mouth. Mixed exocrine glands contain both serous and mucous glands and release both types of secretions.

]]> 2476 2467 1 0 12.1 Basic Structure and Function of the Nervous System https://pressbooks.bccampus.ca/apdouglas2017/chapter/12-1-basic-structure-and-function-of-the-nervous-system/ Mon, 17 Jul 2017 19:00:09 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2484

Learning Objectives

By the end of this section, you will be able to:
  • Identify the anatomical and functional divisions of the nervous system
  • Relate the functional and structural differences between gray matter and white matter structures of the nervous system to the structure of neurons
  • List the basic functions of the nervous system

The picture you have in your mind of the nervous system probably includes the brain, the nervous tissue contained within the cranium, and the spinal cord, the extension of nervous tissue within the vertebral column. That suggests it is made of two organs—and you may not even think of the spinal cord as an organ—but the nervous system is a very complex structure. Within the brain, many different and separate regions are responsible for many different and separate functions. It is as if the nervous system is composed of many organs that all look similar and can only be differentiated using tools such as the microscope or electrophysiology. In comparison, it is easy to see that the stomach is different than the esophagus or the liver, so you can imagine the digestive system as a collection of specific organs.

The Central and Peripheral Nervous Systems

The nervous system can be divided into two major regions: the central and peripheral nervous systems. The central nervous system (CNS) is the brain and spinal cord, and the peripheral nervous system (PNS) is everything else (Figure 1). The brain is contained within the cranial cavity of the skull, and the spinal cord is contained within the vertebral cavity of the vertebral column. It is a bit of an oversimplification to say that the CNS is what is inside these two cavities and the peripheral nervous system is outside of them, but that is one way to start to think about it. In actuality, there are some elements of the peripheral nervous system that are within the cranial or vertebral cavities. The peripheral nervous system is so named because it is on the periphery—meaning beyond the brain and spinal cord. Depending on different aspects of the nervous system, the dividing line between central and peripheral is not necessarily universal.
[caption id="" align="aligncenter" width="380"]This diagram shows a silhouette of a human highlighting the nervous system. The central nervous system is composed of the brain and spinal cord. The brain is a large mass of ridged and striated tissue within the head. The spinal cord extends down from the brain and travels through the torso, ending in the pelvis. Pairs of enlarged nervous tissue, labeled ganglia, flank the spinal cord as it travels through the rib area. The ganglia are part of the peripheral nervous system, along with the many thread-like nerves that radiate from the spinal cord and ganglia through the arms, abdomen and legs. Figure 1. Central and Peripheral Nervous System. The structures of the PNS are referred to as ganglia and nerves, which can be seen as distinct structures. The equivalent structures in the CNS are not obvious from this overall perspective and are best examined in prepared tissue under the microscope.[/caption]

Nervous tissue, present in both the CNS and PNS, contains two basic types of cells: neurons and glial cells. A glial cell is one of a variety of cells that provide a framework of tissue that supports the neurons and their activities. The neuron is the more functionally important of the two, in terms of the communicative function of the nervous system. To describe the functional divisions of the nervous system, it is important to understand the structure of a neuron. Neurons are cells and therefore have a soma, or cell body, but they also have extensions of the cell; each extension is generally referred to as a process. There is one important process that every neuron has called an axon, which is the fiber that connects a neuron with its target. Another type of process that branches off from the soma is the dendrite. Dendrites are responsible for receiving most of the input from other neurons. Looking at nervous tissue, there are regions that predominantly contain cell bodies and regions that are largely composed of just axons. These two regions within nervous system structures are often referred to as gray matter (the regions with many cell bodies and dendrites) or white matter (the regions with many axons). Figure 2 demonstrates the appearance of these regions in the brain and spinal cord. The colors ascribed to these regions are what would be seen in “fresh,” or unstained, nervous tissue. Gray matter is not necessarily gray. It can be pinkish because of blood content, or even slightly tan, depending on how long the tissue has been preserved. But white matter is white because axons are insulated by a lipid-rich substance called myelin. Lipids can appear as white (“fatty”) material, much like the fat on a raw piece of chicken or beef. Actually, gray matter may have that color ascribed to it because next to the white matter, it is just darker—hence, gray.

The distinction between gray matter and white matter is most often applied to central nervous tissue, which has large regions that can be seen with the unaided eye. When looking at peripheral structures, often a microscope is used and the tissue is stained with artificial colors. That is not to say that central nervous tissue cannot be stained and viewed under a microscope, but unstained tissue is most likely from the CNS—for example, a frontal section of the brain or cross section of the spinal cord.

[caption id="" align="aligncenter" width="380"]This photo shows an enlarged view of the dorsal side of a human brain. The right side of the occipital lobe has been shaved to reveal the white and gray matter beneath the surface blood vessels. The white matter branches though the shaved section like the limbs of a tree. The gray matter branches and curves on outside of the white matter, creating a buffer between the outer edges of the occipital lobe and the internal white matter. Figure 2. Gray Matter and White Matter. A brain removed during an autopsy, with a partial section removed, shows white matter surrounded by gray matter. Gray matter makes up the outer cortex of the brain. (credit: modification of work by “Suseno”/Wikimedia Commons)[/caption]

Regardless of the appearance of stained or unstained tissue, the cell bodies of neurons or axons can be located in discrete anatomical structures that need to be named. Those names are specific to whether the structure is central or peripheral. A localized collection of neuron cell bodies in the CNS is referred to as a nucleus. In the PNS, a cluster of neuron cell bodies is referred to as a ganglion. Figure 3 indicates how the term nucleus has a few different meanings within anatomy and physiology. It is the center of an atom, where protons and neutrons are found; it is the center of a cell, where the DNA is found; and it is a center of some function in the CNS. There is also a potentially confusing use of the word ganglion (plural = ganglia) that has a historical explanation. In the central nervous system, there is a group of nuclei that are connected together and were once called the basal ganglia before “ganglion” became accepted as a description for a peripheral structure. Some sources refer to this group of nuclei as the “basal nuclei” to avoid confusion.

[caption id="" align="aligncenter" width="395"]This figure shows two diagrams and a photo, labeled A, B, and C. Image A shows an atom composed of two neutrons and two protons surrounded by a hazy electron cloud. The nucleus of the atom is where the protons and neutrons are located. Image B shows a trumpet shaped cell with a large, oval nucleus near its narrow end. This is the nucleus of a cell. Image C shows an MRI capture of the brain. Two red areas near the center of the brain are highlighted in red. These are the nuclei within the brain. Figure 3. What Is a Nucleus? (a) The nucleus of an atom contains its protons and neutrons. (b) The nucleus of a cell is the organelle that contains DNA. (c) A nucleus in the CNS is a localized center of function with the cell bodies of several neurons, shown here circled in red. (credit c: “Was a bee”/Wikimedia Commons)[/caption]

Terminology applied to bundles of axons also differs depending on location. A bundle of axons, or fibers, found in the CNS is called a tract whereas the same thing in the PNS would be called a nerve. There is an important point to make about these terms, which is that they can both be used to refer to the same bundle of axons. When those axons are in the PNS, the term is nerve, but if they are CNS, the term is tract. The most obvious example of this is the axons that project from the retina into the brain. Those axons are called the optic nerve as they leave the eye, but when they are inside the cranium, they are referred to as the optic tract. There is a specific place where the name changes, which is the optic chiasm, but they are still the same axons (Figure 4). A similar situation outside of science can be described for some roads. Imagine a road called “Broad Street” in a town called “Anyville.” The road leaves Anyville and goes to the next town over, called “Hometown.” When the road crosses the line between the two towns and is in Hometown, its name changes to “Main Street.” That is the idea behind the naming of the retinal axons. In the PNS, they are called the optic nerve, and in the CNS, they are the optic tract. Table 1 helps to clarify which of these terms apply to the central or peripheral nervous systems.

[caption id="" align="aligncenter" width="380"]This illustration shows a superior view of a cross section of the brain. The anterior side of the brain is at the top of the diagram with the two eyes clearly visible. Each eye contains a left nerve tract and a right nerve tract. In the left eye, the left nerve tract travels straight back to the right side of the thalamus. It then enters the left occipital lobe. Conversely, the right nerve tract crosses to the right side of the brain through the optic chiasma. It travels through the right side of the thalamus and enters the right occipital lobe. In the right eye, the opposite is true. The left nerve tract crosses over to the left side of the brain at the optic chiasma, traveling into the left side of the thalamus and the left side of the occipital lobe. However, the right nerve tract leads straight back to the right side of the thalamus and the right occipital lobe. Therefore, the optic chiasma is where the right nerve tract from the right eye crosses over the left nerve tract from the left eye. Figure 4. Optic Nerve Versus Optic Tract. This drawing of the connections of the eye to the brain shows the optic nerve extending from the eye to the chiasm, where the structure continues as the optic tract. The same axons extend from the eye to the brain through these two bundles of fibers, but the chiasm represents the border between peripheral and central.[/caption]
Structures of the CNS and PNS (Table 1)
CNS PNS
Group of Neuron Cell Bodies (i.e., gray matter) Nucleus Ganglion
Bundle of Axons (i.e., white matter) Tract Nerve

Functional Divisions of the Nervous System

The nervous system can also be divided on the basis of its functions, but anatomical divisions and functional divisions are different. The CNS and the PNS both contribute to the same functions, but those functions can be attributed to different regions of the brain (such as the cerebral cortex or the hypothalamus) or to different ganglia in the periphery. The problem with trying to fit functional differences into anatomical divisions is that sometimes the same structure can be part of several functions. For example, the optic nerve carries signals from the retina that are either used for the conscious perception of visual stimuli, which takes place in the cerebral cortex, or for the reflexive responses of smooth muscle tissue that are processed through the hypothalamus.

There are two ways to consider how the nervous system is divided functionally. First, the basic functions of the nervous system are sensation, integration, and response. Secondly, control of the body can be somatic or autonomic—divisions that are largely defined by the structures that are involved in the response. There is also a region of the peripheral nervous system that is called the enteric nervous system that is responsible for a specific set of the functions within the realm of autonomic control related to gastrointestinal functions.

Basic Functions

The nervous system is involved in receiving information about the environment around us (sensation) and generating responses to that information (motor responses). The nervous system can be divided into regions that are responsible for sensation (sensory functions) and for the response (motor functions). But there is a third function that needs to be included. Sensory input needs to be integrated with other sensations, as well as with memories, emotional state, or learning (cognition). Some regions of the nervous system are termed integration or association areas. The process of integration combines sensory perceptions and higher cognitive functions such as memories, learning, and emotion to produce a response.

Sensation. The first major function of the nervous system is sensation—receiving information about the environment to gain input about what is happening outside the body (or, sometimes, within the body). The sensory functions of the nervous system register the presence of a change from homeostasis or a particular event in the environment, known as a stimulus. The senses we think of most are the “big five”: taste, smell, touch, sight, and hearing. The stimuli for taste and smell are both chemical substances (molecules, compounds, ions, etc.), touch is physical or mechanical stimuli that interact with the skin, sight is light stimuli, and hearing is the perception of sound, which is a physical stimulus similar to some aspects of touch. There are actually more senses than just those, but that list represents the major senses. Those five are all senses that receive stimuli from the outside world, and of which there is conscious perception. Additional sensory stimuli might be from the internal environment (inside the body), such as the stretch of an organ wall or the concentration of certain ions in the blood.

Response. The nervous system produces a response on the basis of the stimuli perceived by sensory structures. An obvious response would be the movement of muscles, such as withdrawing a hand from a hot stove, but there are broader uses of the term. The nervous system can cause the contraction of all three types of muscle tissue. For example, skeletal muscle contracts to move the skeleton, cardiac muscle is influenced as heart rate increases during exercise, and smooth muscle contracts as the digestive system moves food along the digestive tract. Responses also include the neural control of glands in the body as well, such as the production and secretion of sweat by the eccrine and merocrine sweat glands found in the skin to lower body temperature.

Responses can be divided into those that are voluntary or conscious (contraction of skeletal muscle) and those that are involuntary (contraction of smooth muscles, regulation of cardiac muscle, activation of glands). Voluntary responses are governed by the somatic nervous system and involuntary responses are governed by the autonomic nervous system, which are discussed in the next section.

Integration. Stimuli that are received by sensory structures are communicated to the nervous system where that information is processed. This is called integration. Stimuli are compared with, or integrated with, other stimuli, memories of previous stimuli, or the state of a person at a particular time. This leads to the specific response that will be generated. Seeing a baseball pitched to a batter will not automatically cause the batter to swing. The trajectory of the ball and its speed will need to be considered. Maybe the count is three balls and one strike, and the batter wants to let this pitch go by in the hope of getting a walk to first base. Or maybe the batter’s team is so far ahead, it would be fun to just swing away.

Controlling the Body

The nervous system can be divided into two parts mostly on the basis of a functional difference in responses. The somatic nervous system (SNS) is responsible for conscious perception and voluntary motor responses. Voluntary motor response means the contraction of skeletal muscle, but those contractions are not always voluntary in the sense that you have to want to perform them. Some somatic motor responses are reflexes, and often happen without a conscious decision to perform them. If your friend jumps out from behind a corner and yells “Boo!” you will be startled and you might scream or leap back. You didn’t decide to do that, and you may not have wanted to give your friend a reason to laugh at your expense, but it is a reflex involving skeletal muscle contractions. Other motor responses become automatic (in other words, unconscious) as a person learns motor skills (referred to as “habit learning” or “procedural memory”).

The autonomic nervous system (ANS) is responsible for involuntary control of the body, usually for the sake of homeostasis (regulation of the internal environment). Sensory input for autonomic functions can be from sensory structures tuned to external or internal environmental stimuli. The motor output extends to smooth and cardiac muscle as well as glandular tissue. The role of the autonomic system is to regulate the organ systems of the body, which usually means to control homeostasis. Sweat glands, for example, are controlled by the autonomic system. When you are hot, sweating helps cool your body down. That is a homeostatic mechanism. But when you are nervous, you might start sweating also. That is not homeostatic, it is the physiological response to an emotional state.

There is another division of the nervous system that describes functional responses. The enteric nervous system (ENS) is responsible for controlling the smooth muscle and glandular tissue in your digestive system. It is a large part of the PNS, and is not dependent on the CNS. It is sometimes valid, however, to consider the enteric system to be a part of the autonomic system because the neural structures that make up the enteric system are a component of the autonomic output that regulates digestion. There are some differences between the two, but for our purposes here there will be a good bit of overlap. See Figure 5 for examples of where these divisions of the nervous system can be found.

[caption id="" align="aligncenter" width="580"]This illustration shows a silhouette of a human with only the brain, spinal cord, PNS ganglia, nerves and a section of the digestive tract visible. The brain, which is part of the CNS, is the area of perception and processing of sensory stimuli (somatic/autonomic), the execution of voluntary motor responses (somatic), and the regulation of homeostatic mechanisms (autonomic). The spinal cord, which is part of the CNS, is the area where reflexes are initiated. The gray matter of the ventral horn initiates somatic reflexes while the gray matter of the lateral horn initiates autonomic reflexes. The spinal cord is also the somatic and autonomic pathway for sensory and motor functions between the PNS and the brain. The nerves, which are part of the PNS, are the fibers of sensory and motor neurons, which can be either somatic or autonomic. The ganglia, which are part of the PNS, are the areas for the reception of somatic and autonomic sensory stimuli. These are received by the dorsal root ganglia and cranial ganglia. The autonomic ganglia are also the relay for visceral motor responses. The digestive tract is part of the enteric nervous system, the ENS, which is located in the digestive tract and is responsible for autonomous function. The ENS can operate independent of the brain and spinal cord. Figure 5. Somatic, Autonomic, and Enteric Structures of the Nervous System. Somatic structures include the spinal nerves, both motor and sensory fibers, as well as the sensory ganglia (posterior root ganglia and cranial nerve ganglia). Autonomic structures are found in the nerves also, but include the sympathetic and parasympathetic ganglia. The enteric nervous system includes the nervous tissue within the organs of the digestive tract.[/caption]
[caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit this site to read about a woman that notices that her daughter is having trouble walking up the stairs.[/caption] [caption id="attachment_2996" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of the nervous system![/caption]
]]> 2484 2477 2 0 12.4 The Action Potential https://pressbooks.bccampus.ca/apdouglas2017/chapter/12-4-the-action-potential/ Mon, 17 Jul 2017 19:01:43 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2500

Learning Objectives

By the end of this section, you will be able to:
  • Describe the components of the membrane that establish the resting membrane potential
  • Describe the changes that occur to the membrane that result in the action potential

The functions of the nervous system—sensation, integration, and response—depend on the functions of the neurons underlying these pathways. To understand how neurons are able to communicate, it is necessary to describe the role of an excitable membrane in generating these signals. The basis of this communication is the action potential, which demonstrates how changes in the membrane can constitute a signal. Looking at the way these signals work in more variable circumstances involves a look at graded potentials, which will be covered in the next section.

Electrically Active Cell Membranes

Most cells in the body make use of charged particles, ions, to build up a charge across the cell membrane. Previously, this was shown to be a part of how muscle cells work. For skeletal muscles to contract, based on excitation–contraction coupling, requires input from a neuron. Both of the cells make use of the cell membrane to regulate ion movement between the extracellular fluid and cytosol.

As you learned in the chapter on cells, the cell membrane is primarily responsible for regulating what can cross the membrane and what stays on only one side. The cell membrane is a phospholipid bilayer, so only substances that can pass directly through the hydrophobic core can diffuse through unaided. Charged particles, which are hydrophilic by definition, cannot pass through the cell membrane without assistance (Figure 1). Transmembrane proteins, specifically channel proteins, make this possible. Several passive transport channels, as well as active transport pumps, are necessary to generate a transmembrane potential and an action potential. Of special interest is the carrier protein referred to as the sodium/potassium pump that moves sodium ions (Na+) out of a cell and potassium ions (K+) into a cell, thus regulating ion concentration on both sides of the cell membrane.

[caption id="" align="aligncenter" width="550"]This diagram shows a cross section of a cell membrane. The cell membrane proteins are large, blocky, objects. Peripheral proteins are not embedded in the phospholipid bilayer. The peripheral protein shown here is attached to the outside surface of another protein on the extracellular fluid side. Integral proteins are embedded between the phospholipids of the cell membrane. The transmembrane integral protein extends through both phospholipids layers. The opposite ends of this protein project into the cytosol and the extracellular fluid. A second, smaller integral protein only extends into the inner phospholipid layer. Its opposite end projects into the cytosol. This second protein is, therefore, not a transmembrane protein. The channel protein is cylinder shaped with a hollow internal tube labeled the pore. The sides of the channel protein can bulge inward to close the pore. Figure 1. Cell Membrane and Transmembrane Proteins. The cell membrane is composed of a phospholipid bilayer and has many transmembrane proteins, including different types of channel proteins that serve as ion channels.[/caption]

The sodium/potassium pump requires energy in the form of adenosine triphosphate (ATP), so it is also referred to as an ATPase. As was explained in the cell chapter, the concentration of Na+ is higher outside the cell than inside, and the concentration of K+ is higher inside the cell than outside. That means that this pump is moving the ions against the concentration gradients for sodium and potassium, which is why it requires energy. In fact, the pump basically maintains those concentration gradients.

Ion channels are pores that allow specific charged particles to cross the membrane in response to an existing concentration gradient. Proteins are capable of spanning the cell membrane, including its hydrophobic core, and can interact with the charge of ions because of the varied properties of amino acids found within specific domains or regions of the protein channel. Hydrophobic amino acids are found in the domains that are apposed to the hydrocarbon tails of the phospholipids. Hydrophilic amino acids are exposed to the fluid environments of the extracellular fluid and cytosol. Additionally, the ions will interact with the hydrophilic amino acids, which will be selective for the charge of the ion. Channels for cations (positive ions) will have negatively charged side chains in the pore. Channels for anions (negative ions) will have positively charged side chains in the pore. This is called electrochemical exclusion, meaning that the channel pore is charge-specific.

Ion channels can also be specified by the diameter of the pore. The distance between the amino acids will be specific for the diameter of the ion when it dissociates from the water molecules surrounding it. Because of the surrounding water molecules, larger pores are not ideal for smaller ions because the water molecules will interact, by hydrogen bonds, more readily than the amino acid side chains. This is called size exclusion. Some ion channels are selective for charge but not necessarily for size, and thus are called a nonspecific channel. These nonspecific channels allow cations—particularly Na+, K+, and Ca2+—to cross the membrane, but exclude anions.

Ion channels do not always freely allow ions to diffuse across the membrane. Some are opened by certain events, meaning the channels are gated. So another way that channels can be categorized is on the basis of how they are gated. Although these classes of ion channels are found primarily in the cells of nervous or muscular tissue, they also can be found in the cells of epithelial and connective tissues.

A ligand-gated channel opens because a signaling molecule, a ligand, binds to the extracellular region of the channel. This type of channel is also known as an ionotropic receptor because when the ligand, known as a neurotransmitter in the nervous system, binds to the protein, ions cross the membrane changing its charge (Figure 2).

[caption id="" align="aligncenter" width="580"]These two diagrams each show a channel protein embedded in the cell membrane. In the left diagram, there is a large number of sodium ions (NA plus) and calcium ions (CA two plus) in the extracellular fluid. Within the cytosol, there is a large number of potassium ions (K plus) but only a few sodium ions. In this diagram, the channel is closed. Two ACH molecules are floating in the extracellular fluid. Their label indicates that a neurotransmitter, a ligand, is required to open the ion channel. The neurotransmitter receptor site on the extracellular fluid side of the channel protein matches the shape of the ACH molecules. In the right diagram, the two ACH molecules attach to the neurotransmitter receptor sites on the channel protein. This opens the channel and the sodium and calcium ions diffuse through the channel and into the cytosol, down their concentration gradient. The potassium ions also diffuse through the channel in the opposite direction down their concentration gradient (out of the cell and into the extracellular fluid). Figure 2. Ligand-Gated Channels. When the ligand, in this case the neurotransmitter acetylcholine, binds to a specific location on the extracellular surface of the channel protein, the pore opens to allow select ions through. The ions, in this case, are cations of sodium, calcium, and potassium.[/caption]

A mechanically gated channel opens because of a physical distortion of the cell membrane. Many channels associated with the sense of touch (somatosensation) are mechanically gated. For example, as pressure is applied to the skin, these channels open and allow ions to enter the cell. Similar to this type of channel would be the channel that opens on the basis of temperature changes, as in testing the water in the shower (Figure 3).

[caption id="" align="aligncenter" width="580"]These two diagrams each show a channel protein embedded in the cell membrane. In the left diagram, there are a large number of sodium ions in the extracellular fluid, but only a few sodium ions in the cytosol. There is a large number of calcium ions in the cytosol but only a few calcium ions in the extracellular fluid. In this diagram, the channel is closed, as the extracellular side has a lid, somewhat resembling that on a trash can, that is closed over the channel opening. In the right diagram, the mechanically gated channel is open. This allows the sodium ions to flow from the extracellular fluid into the cell, down their concentration gradient. At the same time, the calcium ions are moving from the cytosol into the extracellular fluid, down their concentration gradient. Figure 3. Mechanically Gated Channels. When a mechanical change occurs in the surrounding tissue, such as pressure or touch, the channel is physically opened. Thermoreceptors work on a similar principle. When the local tissue temperature changes, the protein reacts by physically opening the channel.[/caption]

A voltage-gated channel is a channel that responds to changes in the electrical properties of the membrane in which it is embedded. Normally, the inner portion of the membrane is at a negative voltage. When that voltage becomes less negative, the channel begins to allow ions to cross the membrane (Figure 4).

[caption id="" align="aligncenter" width="500"]This is a two part diagram. Both diagrams show a voltage gated channel embedded in the lipid membrane bilayer. The channel contains a sphere shaped gate that is attached to a filament. In the first diagram there are several ions in the cytosol but only one ion in the extracellular fluid. The voltage across the membrane is currently minus seventy millivolts and the voltage gated channel is closed. In the second diagram, the voltage in the cytosol is minus fifty millivolts. This voltage change has caused the voltage gated channel to open, as the small sphere is no longer occluding the channel. One of the ions is moving through the channel, down its concentration gradient, and out into the extracellular fluid. Figure 4. Voltage-Gated Channels. Voltage-gated channels open when the transmembrane voltage changes around them. Amino acids in the structure of the protein are sensitive to charge and cause the pore to open to the selected ion.[/caption]

A leakage channel is randomly gated, meaning that it opens and closes at random, hence the reference to leaking. There is no actual event that opens the channel; instead, it has an intrinsic rate of switching between the open and closed states. Leakage channels contribute to the resting transmembrane voltage of the excitable membrane (Figure 5).

[caption id="" align="aligncenter" width="550"]This is a two part diagram. Both diagrams show a leakage channel embedded in the lipid membrane bilayer. The leakage channel is cylindrical with a large, central opening. In the first diagram there are several ions in the cytosol but only one ion in the extracellular fluid. No ions are moving through the leakage channel because the channel is closed. In the second diagram, the leakage channel randomly opens, allowing two ions to travel through the channel, down their concentration gradient, and out into the extracellular fluid. Figure 5. Leakage Channels. In certain situations, ions need to move across the membrane randomly. The particular electrical properties of certain cells are modified by the presence of this type of channel.[/caption]

The Membrane Potential

The electrical state of the cell membrane can have several variations. These are all variations in the membrane potential. A potential is a distribution of charge across the cell membrane, measured in millivolts (mV). The standard is to compare the inside of the cell relative to the outside, so the membrane potential is a value representing the charge on the intracellular side of the membrane based on the outside being zero, relatively speaking (Figure 6).

[caption id="" align="aligncenter" width="550"]This diagram shows a cross section of a cell membrane. The extracellular fluid side of the cell membrane is positively charged while the cytosol side of the membrane is negatively charged. There is a microelectrode embedded in the cell membrane. The microelectrode is attached to a voltmeter, which also has a reference electrode on the extracellular fluid side. The readout of the voltmeter is negative 70 millivolts. Figure 6. Measuring Charge across a Membrane with a Voltmeter. A recording electrode is inserted into the cell and a reference electrode is outside the cell. By comparing the charge measured by these two electrodes, the transmembrane voltage is determined. It is conventional to express that value for the cytosol relative to the outside.[/caption]

The concentration of ions in extracellular and intracellular fluids is largely balanced, with a net neutral charge. However, a slight difference in charge occurs right at the membrane surface, both internally and externally. It is the difference in this very limited region that has all the power in neurons (and muscle cells) to generate electrical signals, including action potentials.

Before these electrical signals can be described, the resting state of the membrane must be explained. When the cell is at rest, and the ion channels are closed (except for leakage channels which randomly open), ions are distributed across the membrane in a very predictable way. The concentration of Na+ outside the cell is 10 times greater than the concentration inside. Also, the concentration of K+ inside the cell is greater than outside. The cytosol contains a high concentration of anions, in the form of phosphate ions and negatively charged proteins. Large anions are a component of the inner cell membrane, including specialized phospholipids and proteins associated with the inner leaflet of the membrane (leaflet is a term used for one side of the lipid bilayer membrane). The negative charge is localized in the large anions.

With the ions distributed across the membrane at these concentrations, the difference in charge is measured at -70 mV, the value described as the resting membrane potential. The exact value measured for the resting membrane potential varies between cells, but -70 mV is most commonly used as this value. This voltage would actually be much lower except for the contributions of some important proteins in the membrane. Leakage channels allow Na+ to slowly move into the cell or K+ to slowly move out, and the Na+/K+ pump restores them. This may appear to be a waste of energy, but each has a role in maintaining the membrane potential.

The Action Potential

Resting membrane potential describes the steady state of the cell, which is a dynamic process that is balanced by ion leakage and ion pumping. Without any outside influence, it will not change. To get an electrical signal started, the membrane potential has to change.

This starts with a channel opening for Na+ in the membrane. Because the concentration of Na+ is higher outside the cell than inside the cell by a factor of 10, ions will rush into the cell that are driven largely by the concentration gradient. Because sodium is a positively charged ion, it will change the relative voltage immediately inside the cell relative to immediately outside. The resting potential is the state of the membrane at a voltage of -70 mV, so the sodium cation entering the cell will cause it to become less negative. This is known as depolarization, meaning the membrane potential moves toward zero.

The concentration gradient for Na+ is so strong that it will continue to enter the cell even after the membrane potential has become zero, so that the voltage immediately around the pore begins to become positive. The electrical gradient also plays a role, as negative proteins below the membrane attract the sodium ion. The membrane potential will reach +30 mV by the time sodium has entered the cell.

As the membrane potential reaches +30 mV, other voltage-gated channels are opening in the membrane. These channels are specific for the potassium ion. A concentration gradient acts on K+, as well. As K+ starts to leave the cell, taking a positive charge with it, the membrane potential begins to move back toward its resting voltage. This is called repolarization, meaning that the membrane voltage moves back toward the -70 mV value of the resting membrane potential.

Repolarization returns the membrane potential to the -70 mV value that indicates the resting potential, but it actually overshoots that value. Potassium ions reach equilibrium when the membrane voltage is below -70 mV, so a period of hyperpolarization occurs while the K+ channels are open. Those K+ channels are slightly delayed in closing, accounting for this short overshoot.

What has been described here is the action potential, which is presented as a graph of voltage over time in Figure 7. It is the electrical signal that nervous tissue generates for communication. The change in the membrane voltage from -70 mV at rest to +30 mV at the end of depolarization is a 100-mV change. That can also be written as a 0.1-V change. To put that value in perspective, think about a battery. An AA battery that you might find in a television remote has a voltage of 1.5 V, or a 9-V battery (the rectangular battery with two posts on one end) is, obviously, 9 V. The change seen in the action potential is one or two orders of magnitude less than the charge in these batteries. In fact, the membrane potential can be described as a battery. A charge is stored across the membrane that can be released under the correct conditions. A battery in your remote has stored a charge that is “released” when you push a button.

[caption id="" align="aligncenter" width="380"]This graph has membrane potential, in millivolts, on the X axis, ranging from negative 70 to positive thirty. Time is on the X axis. The plot line starts steadily at negative seventy and then increases to negative 55 millivolts. The plot line then increases quickly, peaking at positive thirty. This is the depolarization phase. The plot line then quickly drops back to negative seventy millivolts. This is the repolarization phase. The plot line continues to drop but then gradually increases back to negative seventy millivolts. The area where the plot line is below negative seventy millivolts is the hyperpolarization phase. Figure 7. Graph of Action Potential. Plotting voltage measured across the cell membrane against time, the action potential begins with depolarization, followed by repolarization, which goes past the resting potential into hyperpolarization, and finally the membrane returns to rest.[/caption]
[caption id="attachment_2998" align="aligncenter" width="150"] Watch this Crashcourse video to learn more about the action potential![/caption]

The question is, now, what initiates the action potential? The description above conveniently glosses over that point. But it is vital to understanding what is happening. The membrane potential will stay at the resting voltage until something changes. The description above just says that a Na+ channel opens. Now, to say “a channel opens” does not mean that one individual transmembrane protein changes. Instead, it means that one kind of channel opens. There are a few different types of channels that allow Na+ to cross the membrane. A ligand-gated Na+ channel will open when a neurotransmitter binds to it and a mechanically gated Na+ channel will open when a physical stimulus affects a sensory receptor (like pressure applied to the skin compresses a touch receptor). Whether it is a neurotransmitter binding to its receptor protein or a sensory stimulus activating a sensory receptor cell, some stimulus gets the process started. Sodium starts to enter the cell and the membrane becomes less negative.

A third type of channel that is an important part of depolarization in the action potential is the voltage-gated Na+ channel. The channels that start depolarizing the membrane because of a stimulus help the cell to depolarize from -70 mV to -55 mV. Once the membrane reaches that voltage, the voltage-gated Na+ channels open. This is what is known as the threshold. Any depolarization that does not change the membrane potential to -55 mV or higher will not reach threshold and thus will not result in an action potential. Also, any stimulus that depolarizes the membrane to -55 mV or beyond will cause a large number of channels to open and an action potential will be initiated.

Because of the threshold, the action potential can be likened to a digital event—it either happens or it does not. If the threshold is not reached, then no action potential occurs. If depolarization reaches -55 mV, then the action potential continues and runs all the way to +30 mV, at which K+ causes repolarization, including the hyperpolarizing overshoot. Also, those changes are the same for every action potential, which means that once the threshold is reached, the exact same thing happens. A stronger stimulus, which might depolarize the membrane well past threshold, will not make a “bigger” action potential. Action potentials are “all or none.” Either the membrane reaches the threshold and everything occurs as described above, or the membrane does not reach the threshold and nothing else happens. All action potentials peak at the same voltage (+30 mV), so one action potential is not bigger than another. Stronger stimuli will initiate multiple action potentials more quickly, but the individual signals are not bigger. Thus, for example, you will not feel a greater sensation of pain, or have a stronger muscle contraction, because of the size of the action potential because they are not different sizes.

As we have seen, the depolarization and repolarization of an action potential are dependent on two types of channels (the voltage-gated Na+ channel and the voltage-gated K+ channel). The voltage-gated Na+ channel actually has two gates. One is the activation gate, which opens when the membrane potential crosses -55 mV. The other gate is the inactivation gate, which closes after a specific period of time—on the order of a fraction of a millisecond. When a cell is at rest, the activation gate is closed and the inactivation gate is open. However, when the threshold is reached, the activation gate opens, allowing Na+ to rush into the cell. Timed with the peak of depolarization, the inactivation gate closes. During repolarization, no more sodium can enter the cell. When the membrane potential passes -55 mV again, the activation gate closes. After that, the inactivation gate re-opens, making the channel ready to start the whole process over again.

The voltage-gated K+ channel has only one gate, which is sensitive to a membrane voltage of -50 mV. However, it does not open as quickly as the voltage-gated Na+ channel does. It might take a fraction of a millisecond for the channel to open once that voltage has been reached. The timing of this coincides exactly with when the Na+ flow peaks, so voltage-gated K+ channels open just as the voltage-gated Na+ channels are being inactivated. As the membrane potential repolarizes and the voltage passes -50 mV again, the channel closes—again, with a little delay. Potassium continues to leave the cell for a short while and the membrane potential becomes more negative, resulting in the hyperpolarizing overshoot. Then the channel closes again and the membrane can return to the resting potential because of the ongoing activity of the non-gated channels and the Na+/K+ pump.

All of this takes place within approximately 2 milliseconds (Figure 8). While an action potential is in progress, another one cannot be initiated. That effect is referred to as the refractory period. There are two phases of the refractory period: the absolute refractory period and the relative refractory period. During the absolute phase, another action potential will not start. This is because of the inactivation gate of the voltage-gated Na+ channel. Once that channel is back to its resting conformation (less than -55 mV), a new action potential could be started, but only by a stronger stimulus than the one that initiated the current action potential. This is because of the flow of K+ out of the cell. Because that ion is rushing out, any Na+ that tries to enter will not depolarize the cell, but will only keep the cell from hyperpolarizing.

[caption id="" align="aligncenter" width="380"]This graph has membrane potential, in millivolts, on the X axis, ranging from negative 70 to positive thirty. Time is on the X axis. In step one, which is labeled at rest, the plot line is steady at negative seventy millivolts. In step 2, a stimulus is applied, causing the plot line to increase to positive 30 millivolts. The curve sharply increases at step three, labeled voltage rises. After peaking at positive thirty, the plot line then quickly drops back to negative 70. This is the fourth step, labeled voltage falls. The plot line continues to drop below negative 70 and this is step 5, labeled end of action potential. Finally, the plot line gradually increases back to negative seventy millivolts, which is step 6, labeled return to rest. Figure 8. Stages of an Action Potential. Plotting voltage measured across the cell membrane against time, the events of the action potential can be related to specific changes in the membrane voltage. (1) At rest, the membrane voltage is -70 mV. (2) The membrane begins to depolarize when an external stimulus is applied. (3) The membrane voltage begins a rapid rise toward +30 mV. (4) The membrane voltage starts to return to a negative value. (5) Repolarization continues past the resting membrane voltage, resulting in hyperpolarization. (6) The membrane voltage returns to the resting value shortly after hyperpolarization.[/caption]

Propagation of the Action Potential

The action potential is initiated at the beginning of the axon, at what is called the initial segment. There is a high density of voltage-gated Na+ channels so that rapid depolarization can take place here. Going down the length of the axon, the action potential is propagated because more voltage-gated Na+ channels are opened as the depolarization spreads. This spreading occurs because Na+ enters through the channel and moves along the inside of the cell membrane. As the Na+ moves, or flows, a short distance along the cell membrane, its positive charge depolarizes a little more of the cell membrane. As that depolarization spreads, new voltage-gated Na+ channels open and more ions rush into the cell, spreading the depolarization a little farther.

Because voltage-gated Na+ channels are inactivated at the peak of the depolarization, they cannot be opened again for a brief time—the absolute refractory period. Because of this, depolarization spreading back toward previously opened channels has no effect. The action potential must propagate toward the axon terminals; as a result, the polarity of the neuron is maintained, as mentioned above.

Propagation, as described above, applies to unmyelinated axons. When myelination is present, the action potential propagates differently. Sodium ions that enter the cell at the initial segment start to spread along the length of the axon segment, but there are no voltage-gated Na+ channels until the first node of Ranvier. Because there is not constant opening of these channels along the axon segment, the depolarization spreads at an optimal speed. The distance between nodes is the optimal distance to keep the membrane still depolarized above threshold at the next node. As Na+ spreads along the inside of the membrane of the axon segment, the charge starts to dissipate. If the node were any farther down the axon, that depolarization would have fallen off too much for voltage-gated Na+ channels to be activated at the next node of Ranvier. If the nodes were any closer together, the speed of propagation would be slower.

Propagation along an unmyelinated axon is referred to as continuous conduction; along the length of a myelinated axon, it is saltatory conduction. Continuous conduction is slow because there are always voltage-gated Na+ channels opening, and more and more Na+ is rushing into the cell. Saltatory conduction is faster because the action potential basically jumps from one node to the next (saltare = “to leap”), and the new influx of Na+ renews the depolarized membrane. Along with the myelination of the axon, the diameter of the axon can influence the speed of conduction. Much as water runs faster in a wide river than in a narrow creek, Na+-based depolarization spreads faster down a wide axon than down a narrow one. This concept is known as resistance and is generally true for electrical wires or plumbing, just as it is true for axons, although the specific conditions are different at the scales of electrons or ions versus water in a river.

Homeostatic Imbalances

Potassium Concentration Glial cells, especially astrocytes, are responsible for maintaining the chemical environment of the CNS tissue. The concentrations of ions in the extracellular fluid are the basis for how the membrane potential is established and changes in electrochemical signaling. If the balance of ions is upset, drastic outcomes are possible.

Normally the concentration of K+ is higher inside the neuron than outside. After the repolarizing phase of the action potential, K+ leakage channels and the Na+/K+ pump ensure that the ions return to their original locations. Following a stroke or other ischemic event, extracellular K+ levels are elevated. The astrocytes in the area are equipped to clear excess K+ to aid the pump. But when the level is far out of balance, the effects can be irreversible.

Astrocytes can become reactive in cases such as these, which impairs their ability to maintain the local chemical environment. The glial cells enlarge and their processes swell. They lose their K+ buffering ability and the function of the pump is affected, or even reversed. One of the early signs of cell disease is this "leaking" of sodium ions into the body cells. This sodium/potassium imbalance negatively affects the internal chemistry of cells, preventing them from functioning normally.

]]> 2500 2477 5 0 13.2 The Central Nervous System https://pressbooks.bccampus.ca/apdouglas2017/chapter/13-2-the-central-nervous-system/ Mon, 17 Jul 2017 19:04:45 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2522

Learning Objectives

By the end of this section, you will be able to:
  • Name the major regions of the adult brain
  • Describe the connections between the cerebrum and brain stem through the diencephalon, and from those regions into the spinal cord
  • Recognize the complex connections within the subcortical structures of the basal nuclei
  • Explain the arrangement of gray and white matter in the spinal cord

The brain and the spinal cord are the central nervous system, and they represent the main organs of the nervous system. The spinal cord is a single structure, whereas the adult brain is described in terms of four major regions: the cerebrum, the diencephalon, the brain stem, and the cerebellum. A person’s conscious experiences are based on neural activity in the brain. The regulation of homeostasis is governed by a specialized region in the brain. The coordination of reflexes depends on the integration of sensory and motor pathways in the spinal cord.

The Cerebrum

The iconic grey mantle of the human brain, which appears to make up most of the mass of the brain, is the cerebrum (Figure 1). The wrinkled portion is the cerebral cortex, and the rest of the structure is beneath that outer covering. There is a large separation between the two sides of the cerebrum called the longitudinal fissure. It separates the cerebrum into two distinct halves, a right and left cerebral hemisphere. Deep within the cerebrum, the white matter of the corpus callosum provides the major pathway for communication between the two hemispheres of the cerebral cortex.

[caption id="" align="aligncenter" width="600"]This figure shows the lateral view on the left panel and anterior view on the right panel of the brain. The major parts including the cerebrum are labeled. Figure 1. The Cerebrum. The cerebrum is a large component of the CNS in humans, and the most obvious aspect of it is the folded surface called the cerebral cortex.[/caption]

Many of the higher neurological functions, such as memory, emotion, and consciousness, are the result of cerebral function. The complexity of the cerebrum is different across vertebrate species. The cerebrum of the most primitive vertebrates is not much more than the connection for the sense of smell. In mammals, the cerebrum comprises the outer grey matter that is the cortex (from the Latin word meaning “bark of a tree”) and several deep nuclei that belong to three important functional groups. The basal nuclei are responsible for cognitive processing, the most important function being that associated with planning movements. The basal forebrain contains nuclei that are important in learning and memory. The limbic cortex is the region of the cerebral cortex that is part of the limbic system, a collection of structures involved in emotion, memory, and behavior.

Cerebral Cortex

The cerebrum is covered by a continuous layer of grey matter that wraps around either side of the forebrain—the cerebral cortex. This thin, extensive region of wrinkled gray matter is responsible for the higher functions of the nervous system. A gyrus (plural = gyri) is the ridge of one of those wrinkles, and a sulcus (plural = sulci) is the groove between two gyri. The pattern of these folds of tissue indicates specific regions of the cerebral cortex.

The head is limited by the size of the birth canal, and the brain must fit inside the cranial cavity of the skull. Extensive folding in the cerebral cortex enables more grey matter to fit into this limited space. If the grey matter of the cortex were peeled off of the cerebrum and laid out flat, its surface area would be roughly equal to one square meter.

The folding of the cortex maximizes the amount of grey matter in the cranial cavity. During embryonic development, as the telencephalon expands within the skull, the brain goes through a regular course of growth that results in everyone’s brain having a similar pattern of folds. The surface of the brain can be mapped on the basis of the locations of large gyri and sulci. Using these landmarks, the cortex can be separated into four major regions, or lobes (Figure 2). The lateral sulcus that separates the temporal lobe from the other regions is one such landmark. Superior to the lateral sulcus are the parietal lobe and frontal lobe, which are separated from each other by the central sulcus. The posterior region of the cortex is the occipital lobe, which has no obvious anatomical border between it and the parietal or temporal lobes on the lateral surface of the brain. From the medial surface, an obvious landmark separating the parietal and occipital lobes is called the parieto-occipital sulcus. The fact that there is no obvious anatomical border between these lobes is consistent with the functions of these regions being interrelated.

[caption id="" align="aligncenter" width="380"]This figure shows the lateral view of the brain and the major lobes are labeled. Figure 2. Lobes of the Cerebral Cortex. The cerebral cortex is divided into four lobes. Extensive folding increases the surface area available for cerebral functions.[/caption]

Different regions of the cerebral cortex can be associated with particular functions, a concept known as localization of function. In the early 1900s, a German neuroscientist named Korbinian Brodmann performed an extensive study of the microscopic anatomy—the cytoarchitecture—of the cerebral cortex and divided the cortex into 52 separate regions on the basis of the histology of the cortex. His work resulted in a system of classification known as Brodmann’s areas, which is still used today to describe the anatomical distinctions within the cortex (Figure 3). The results from Brodmann’s work on the anatomy align very well with the functional differences within the cortex. Areas 17 and 18 in the occipital lobe are responsible for primary visual perception. That visual information is complex, so it is processed in the temporal and parietal lobes as well.

The temporal lobe is associated with primary auditory sensation, known as Brodmann’s areas 41 and 42 in the superior temporal lobe. Because regions of the temporal lobe are part of the limbic system, memory is an important function associated with that lobe. Memory is essentially a sensory function; memories are recalled sensations such as the smell of Mom’s baking or the sound of a barking dog. Even memories of movement are really the memory of sensory feedback from those movements, such as stretching muscles or the movement of the skin around a joint. Structures in the temporal lobe are responsible for establishing long-term memory, but the ultimate location of those memories is usually in the region in which the sensory perception was processed.

The main sensation associated with the parietal lobe is somatosensation, meaning the general sensations associated with the body. Posterior to the central sulcus is the postcentral gyrus, the primary somatosensory cortex, which is identified as Brodmann’s areas 1, 2, and 3. All of the tactile senses are processed in this area, including touch, pressure, tickle, pain, itch, and vibration, as well as more general senses of the body such as proprioception and kinesthesia, which are the senses of body position and movement, respectively.

Anterior to the central sulcus is the frontal lobe, which is primarily associated with motor functions. The precentral gyrus is the primary motor cortex. Cells from this region of the cerebral cortex are the upper motor neurons that instruct cells in the spinal cord to move skeletal muscles. Anterior to this region are a few areas that are associated with planned movements. The premotor area is responsible for thinking of a movement to be made. The frontal eye fields are important in eliciting eye movements and in attending to visual stimuli. Broca’s area is responsible for the production of language, or controlling movements responsible for speech; in the vast majority of people, it is located only on the left side. Anterior to these regions is the prefrontal lobe, which serves cognitive functions that can be the basis of personality, short-term memory, and consciousness. The prefrontal lobotomy is an outdated mode of treatment for personality disorders (psychiatric conditions) that profoundly affected the personality of the patient.

[caption id="" align="aligncenter" width="550"]In this figure, the Brodmann areas, identifying the functional regions of the brain, are mapped. The left panel shows the lateral surface of the brain and the right panel shows the medial surface. Figure 3. Brodmann's Areas of the Cerebral Cortex. Brodmann mapping of functionally distinct regions of the cortex was based on its cytoarchitecture at a microscopic level.[/caption]

Subcortical structures

Beneath the cerebral cortex are sets of nuclei known as subcortical nuclei that augment cortical processes. The nuclei of the basal forebrain serve as the primary location for acetylcholine production, which modulates the overall activity of the cortex, possibly leading to greater attention to sensory stimuli. Alzheimer’s disease is associated with a loss of neurons in the basal forebrain. The hippocampus and amygdala are medial-lobe structures that, along with the adjacent cortex, are involved in long-term memory formation and emotional responses. The basal nuclei are a set of nuclei in the cerebrum responsible for comparing cortical processing with the general state of activity in the nervous system to influence the likelihood of movement taking place. For example, while a student is sitting in a classroom listening to a lecture, the basal nuclei will keep the urge to jump up and scream from actually happening. (The basal nuclei are also referred to as the basal ganglia, although that is potentially confusing because the term ganglia is typically used for peripheral structures.)

The major structures of the basal nuclei that control movement are the caudate, putamen, and globus pallidus, which are located deep in the cerebrum. The caudate is a long nucleus that follows the basic C-shape of the cerebrum from the frontal lobe, through the parietal and occipital lobes, into the temporal lobe. The putamen is mostly deep in the anterior regions of the frontal and parietal lobes. Together, the caudate and putamen are called the striatum. The globus pallidus is a layered nucleus that lies just medial to the putamen; they are called the lenticular nuclei because they look like curved pieces fitting together like lenses. The globus pallidus has two subdivisions, the external and internal segments, which are lateral and medial, respectively. These nuclei are depicted in a frontal section of the brain in Figure 4.

[caption id="" align="aligncenter" width="380"]This diagram shows the frontal section of the brain and identifies the major components of the basal nuclei. Figure 4. Frontal Section of Cerebral Cortex and Basal Nuclei. The major components of the basal nuclei, shown in a frontal section of the brain, are the caudate (just lateral to the lateral ventricle), the putamen (inferior to the caudate and separated by the large white-matter structure called the internal capsule), and the globus pallidus (medial to the putamen).[/caption]

The basal nuclei in the cerebrum are connected with a few more nuclei in the brain stem that together act as a functional group that forms a motor pathway. Two streams of information processing take place in the basal nuclei. All input to the basal nuclei is from the cortex into the striatum (Figure 5). The direct pathway is the projection of axons from the striatum to the globus pallidus internal segment (GPi) and the substantia nigra pars reticulata (SNr). The GPi/SNr then projects to the thalamus, which projects back to the cortex. The indirect pathway is the projection of axons from the striatum to the globus pallidus external segment (GPe), then to the subthalamic nucleus (STN), and finally to GPi/SNr. The two streams both target the GPi/SNr, but one has a direct projection and the other goes through a few intervening nuclei. The direct pathway causes the disinhibition of the thalamus (inhibition of one cell on a target cell that then inhibits the first cell), whereas the indirect pathway causes, or reinforces, the normal inhibition of the thalamus. The thalamus then can either excite the cortex (as a result of the direct pathway) or fail to excite the cortex (as a result of the indirect pathway).

[caption id="" align="aligncenter" width="295"]This flowchart shows the connection between the different regions of the brain such as the cortex, striatum and the thalamus. Figure 5. Connections of Basal Nuclei. Input to the basal nuclei is from the cerebral cortex, which is an excitatory connection releasing glutamate as a neurotransmitter. This input is to the striatum, or the caudate and putamen. In the direct pathway, the striatum projects to the internal segment of the globus pallidus and the substantia nigra pars reticulata (GPi/SNr). This is an inhibitory pathway, in which GABA is released at the synapse, and the target cells are hyperpolarized and less likely to fire. The output from the basal nuclei is to the thalamus, which is an inhibitory projection using GABA.[/caption]

The switch between the two pathways is the substantia nigra pars compacta, which projects to the striatum and releases the neurotransmitter dopamine. Dopamine receptors are either excitatory (D1-type receptors) or inhibitory (D2-type receptors). The direct pathway is activated by dopamine, and the indirect pathway is inhibited by dopamine. When the substantia nigra pars compacta is firing, it signals to the basal nuclei that the body is in an active state, and movement will be more likely. When the substantia nigra pars compacta is silent, the body is in a passive state, and movement is inhibited. To illustrate this situation, while a student is sitting listening to a lecture, the substantia nigra pars compacta would be silent and the student less likely to get up and walk around. Likewise, while the professor is lecturing, and walking around at the front of the classroom, the professor’s substantia nigra pars compacta would be active, in keeping with his or her activity level.

The Diencephalon

The diencephalon is the one region of the adult brain that retains its name from embryologic development. The etymology of the word diencephalon translates to “through brain.” It is the connection between the cerebrum and the rest of the nervous system, with one exception. The rest of the brain, the spinal cord, and the PNS all send information to the cerebrum through the diencephalon. Output from the cerebrum passes through the diencephalon. The single exception is the system associated with olfaction, or the sense of smell, which connects directly with the cerebrum. In the earliest vertebrate species, the cerebrum was not much more than olfactory bulbs that received peripheral information about the chemical environment (to call it smell in these organisms is imprecise because they lived in the ocean). The diencephalon is deep beneath the cerebrum and constitutes the walls of the third ventricle. The diencephalon can be described as any region of the brain with “thalamus” in its name. The two major regions of the diencephalon are the thalamus itself and the hypothalamus (Figure 6). There are other structures, such as the epithalamus, which contains the pineal gland, or the subthalamus, which includes the subthalamic nucleus that is part of the basal nuclei.

Thalamus

The thalamus is a collection of nuclei that relay information between the cerebral cortex and the periphery, spinal cord, or brain stem. All sensory information, except for the sense of smell, passes through the thalamus before processing by the cortex. Axons from the peripheral sensory organs, or intermediate nuclei, synapse in the thalamus, and thalamic neurons project directly to the cerebrum. It is a requisite synapse in any sensory pathway, except for olfaction. The thalamus does not just pass the information on, it also processes that information. For example, the portion of the thalamus that receives visual information will influence what visual stimuli are important, or what receives attention.

The cerebrum also sends information down to the thalamus, which usually communicates motor commands. This involves interactions with the cerebellum and other nuclei in the brain stem. The cerebrum interacts with the basal nuclei, which involves connections with the thalamus. The primary output of the basal nuclei is to the thalamus, which relays that output to the cerebral cortex. The cortex also sends information to the thalamus that will then influence the effects of the basal nuclei.

Hypothalamus

Inferior and slightly anterior to the thalamus is the hypothalamus, the other major region of the diencephalon. The hypothalamus is a collection of nuclei that are largely involved in regulating homeostasis. The hypothalamus is the executive region in charge of the autonomic nervous system and the endocrine system through its regulation of the anterior pituitary gland. Other parts of the hypothalamus are involved in memory and emotion as part of the limbic system.

[caption id="" align="aligncenter" width="450"]This figure shows the location of the thalamus, hypothalamus and pituitary gland in the brain. Figure 6. The Diencephalon. The diencephalon is composed primarily of the thalamus and hypothalamus, which together define the walls of the third ventricle. The thalami are two elongated, ovoid structures on either side of the midline that make contact in the middle. The hypothalamus is inferior and anterior to the thalamus, culminating in a sharp angle to which the pituitary gland is attached.[/caption]

Brain Stem

The midbrain and hindbrain (composed of the pons and the medulla) are collectively referred to as the brain stem (Figure 7). The structure emerges from the ventral surface of the forebrain as a tapering cone that connects the brain to the spinal cord. Attached to the brain stem, but considered a separate region of the adult brain, is the cerebellum. The midbrain coordinates sensory representations of the visual, auditory, and somatosensory perceptual spaces. The pons is the main connection with the cerebellum. The pons and the medulla regulate several crucial functions, including the cardiovascular and respiratory systems and rates.

The cranial nerves connect through the brain stem and provide the brain with the sensory input and motor output associated with the head and neck, including most of the special senses. The major ascending and descending pathways between the spinal cord and brain, specifically the cerebrum, pass through the brain stem.
[caption id="" align="aligncenter" width="450"]This figure shows the location of the midbrain, pons and the medulla in the brain. Figure 7. The Brain Stem. The brain stem comprises three regions: the midbrain, the pons, and the medulla.[/caption]

Midbrain

One of the original regions of the embryonic brain, the midbrain is a small region between the thalamus and pons. It is separated into the tectum and tegmentum, from the Latin words for roof and floor, respectively. The cerebral aqueduct passes through the center of the midbrain, such that these regions are the roof and floor of that canal.

The tectum is composed of four bumps known as the colliculi (singular = colliculus), which means “little hill” in Latin. The inferior colliculus is the inferior pair of these enlargements and is part of the auditory brain stem pathway. Neurons of the inferior colliculus project to the thalamus, which then sends auditory information to the cerebrum for the conscious perception of sound. The superior colliculus is the superior pair and combines sensory information about visual space, auditory space, and somatosensory space. Activity in the superior colliculus is related to orienting the eyes to a sound or touch stimulus. If you are walking along the sidewalk on campus and you hear chirping, the superior colliculus coordinates that information with your awareness of the visual location of the tree right above you. That is the correlation of auditory and visual maps. If you suddenly feel something wet fall on your head, your superior colliculus integrates that with the auditory and visual maps and you know that the chirping bird just relieved itself on you. You want to look up to see the culprit, but do not.

The tegmentum is continuous with the gray matter of the rest of the brain stem. Throughout the midbrain, pons, and medulla, the tegmentum contains the nuclei that receive and send information through the cranial nerves, as well as regions that regulate important functions such as those of the cardiovascular and respiratory systems.

Pons

The word pons comes from the Latin word for bridge. It is visible on the anterior surface of the brain stem as the thick bundle of white matter attached to the cerebellum. The pons is the main connection between the cerebellum and the brain stem. The bridge-like white matter is only the anterior surface of the pons; the gray matter beneath that is a continuation of the tegmentum from the midbrain. Gray matter in the tegmentum region of the pons contains neurons receiving descending input from the forebrain that is sent to the cerebellum.

Medulla

The medulla is the region known as the myelencephalon in the embryonic brain. The initial portion of the name, “myel,” refers to the significant white matter found in this region—especially on its exterior, which is continuous with the white matter of the spinal cord. The tegmentum of the midbrain and pons continues into the medulla because this gray matter is responsible for processing cranial nerve information. A diffuse region of gray matter throughout the brain stem, known as the reticular formation, is related to sleep and wakefulness, such as general brain activity and attention.

The Cerebellum

The cerebellum, as the name suggests, is the “little brain.” It is covered in gyri and sulci like the cerebrum, and looks like a miniature version of that part of the brain (Figure 8). The cerebellum is largely responsible for comparing information from the cerebrum with sensory feedback from the periphery through the spinal cord. It accounts for approximately 10 percent of the mass of the brain.

[caption id="" align="aligncenter" width="420"]This figure shows the location of the cerebellum in the brain. In the top panel, a lateral view labels the location of the cerebellum and the deep cerebellar white matter. In the bottom panel, a photograph of a brain, with the cerebellum in pink is shown. Figure 8. The Cerebellum. The cerebellum is situated on the posterior surface of the brain stem. Descending input from the cerebellum enters through the large white matter structure of the pons. Ascending input from the periphery and spinal cord enters through the fibers of the inferior olive. Output goes to the midbrain, which sends a descending signal to the spinal cord.[/caption]

Descending fibers from the cerebrum have branches that connect to neurons in the pons. Those neurons project into the cerebellum, providing a copy of motor commands sent to the spinal cord. Sensory information from the periphery, which enters through spinal or cranial nerves, is copied to a nucleus in the medulla known as the inferior olive. Fibers from this nucleus enter the cerebellum and are compared with the descending commands from the cerebrum. If the primary motor cortex of the frontal lobe sends a command down to the spinal cord to initiate walking, a copy of that instruction is sent to the cerebellum. Sensory feedback from the muscles and joints, proprioceptive information about the movements of walking, and sensations of balance are sent to the cerebellum through the inferior olive and the cerebellum compares them. If walking is not coordinated, perhaps because the ground is uneven or a strong wind is blowing, then the cerebellum sends out a corrective command to compensate for the difference between the original cortical command and the sensory feedback. The output of the cerebellum is into the midbrain, which then sends a descending input to the spinal cord to correct the messages going to skeletal muscles.

The Spinal Cord

The description of the CNS is concentrated on the structures of the brain, but the spinal cord is another major organ of the system. Whereas the brain develops out of expansions of the neural tube into primary and then secondary vesicles, the spinal cord maintains the tube structure and is only specialized into certain regions. As the spinal cord continues to develop in the newborn, anatomical features mark its surface. The anterior midline is marked by the anterior median fissure, and the posterior midline is marked by the posterior median sulcus. Axons enter the posterior side through the dorsal (posterior) nerve root, which marks the posterolateral sulcus on either side. The axons emerging from the anterior side do so through the ventral (anterior) nerve root. Note that it is common to see the terms dorsal (dorsal = “back”) and ventral (ventral = “belly”) used interchangeably with posterior and anterior, particularly in reference to nerves and the structures of the spinal cord. You should learn to be comfortable with both.

On the whole, the posterior regions are responsible for sensory functions and the anterior regions are associated with motor functions. This comes from the initial development of the spinal cord, which is divided into the basal plate and the alar plate. The basal plate is closest to the ventral midline of the neural tube, which will become the anterior face of the spinal cord and gives rise to motor neurons. The alar plate is on the dorsal side of the neural tube and gives rise to neurons that will receive sensory input from the periphery.

The length of the spinal cord is divided into regions that correspond to the regions of the vertebral column. The name of a spinal cord region corresponds to the level at which spinal nerves pass through the intervertebral foramina. Immediately adjacent to the brain stem is the cervical region, followed by the thoracic, then the lumbar, and finally the sacral region. The spinal cord is not the full length of the vertebral column because the spinal cord does not grow significantly longer after the first or second year, but the skeleton continues to grow. The nerves that emerge from the spinal cord pass through the intervertebral formina at the respective levels. As the vertebral column grows, these nerves grow with it and result in a long bundle of nerves that resembles a horse’s tail and is named the cauda equina. The sacral spinal cord is at the level of the upper lumbar vertebral bones. The spinal nerves extend from their various levels to the proper level of the vertebral column.

Gray Horns

In cross-section, the gray matter of the spinal cord has the appearance of an ink-blot test, with the spread of the gray matter on one side replicated on the other—a shape reminiscent of a bulbous capital “H.” As shown in Figure 9, the gray matter is subdivided into regions that are referred to as horns. The posterior horn is responsible for sensory processing. The anterior horn sends out motor signals to the skeletal muscles. The lateral horn, which is only found in the thoracic, upper lumbar, and sacral regions, is the central component of the sympathetic division of the autonomic nervous system.

Some of the largest neurons of the spinal cord are the multipolar motor neurons in the anterior horn. The fibers that cause contraction of skeletal muscles are the axons of these neurons. The motor neuron that causes contraction of the big toe, for example, is located in the sacral spinal cord. The axon that has to reach all the way to the belly of that muscle may be a meter in length. The neuronal cell body that maintains that long fiber must be quite large, possibly several hundred micrometers in diameter, making it one of the largest cells in the body.
[caption id="" align="aligncenter" width="380"]This figure shows the cross section of the spinal cord. The top panel shows a diagram of the cross section and the major parts are labeled. The bottom panel shows an ultrasound image of the spinal cord cross section. Figure 9. Cross-section of Spinal Cord. The cross-section of a thoracic spinal cord segment shows the posterior, anterior, and lateral horns of gray matter, as well as the posterior, anterior, and lateral columns of white matter. LM × 40. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

White Columns

Just as the gray matter is separated into horns, the white matter of the spinal cord is separated into columns. Ascending tracts of nervous system fibers in these columns carry sensory information up to the brain, whereas descending tracts carry motor commands from the brain. Looking at the spinal cord longitudinally, the columns extend along its length as continuous bands of white matter. Between the two posterior horns of gray matter are the posterior columns. Between the two anterior horns, and bounded by the axons of motor neurons emerging from that gray matter area, are the anterior columns. The white matter on either side of the spinal cord, between the posterior horn and the axons of the anterior horn neurons, are the lateral columns. The posterior columns are composed of axons of ascending tracts. The anterior and lateral columns are composed of many different groups of axons of both ascending and descending tracts—the latter carrying motor commands down from the brain to the spinal cord to control output to the periphery.

[caption id="attachment_3000" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of the central nervous system![/caption]
]]> 2522 2512 1 0 14.1 Sensory Perception https://pressbooks.bccampus.ca/apdouglas2017/chapter/14-1-sensory-perception/ Mon, 17 Jul 2017 19:09:01 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2553

Learning Objectives

By the end of this section, you will be able to:
  • Describe different types of sensory receptors
  • Describe the structures responsible for the special senses of taste, smell, hearing, balance, and vision
  • Distinguish how different tastes are transduced
  • Describe the means of mechanoreception for hearing and balance
  • List the supporting structures around the eye and describe the structure of the eyeball
  • Describe the processes of phototransduction

A major role of sensory receptors is to help us learn about the environment around us, or about the state of our internal environment. Stimuli from varying sources, and of different types, are received and changed into the electrochemical signals of the nervous system. This occurs when a stimulus changes the cell membrane potential of a sensory neuron. The stimulus causes the sensory cell to produce an action potential that is relayed into the central nervous system (CNS), where it is integrated with other sensory information—or sometimes higher cognitive functions—to become a conscious perception of that stimulus. The central integration may then lead to a motor response.

Describing sensory function with the term sensation or perception is a deliberate distinction. Sensation is the activation of sensory receptor cells at the level of the stimulus. Perception is the central processing of sensory stimuli into a meaningful pattern. Perception is dependent on sensation, but not all sensations are perceived. Receptors are the cells or structures that detect sensations. A receptor cell is changed directly by a stimulus. A transmembrane protein receptor is a protein in the cell membrane that mediates a physiological change in a neuron, most often through the opening of ion channels or changes in the cell signaling processes. Transmembrane receptors are activated by chemicals called ligands. For example, a molecule in food can serve as a ligand for taste receptors. Other transmembrane proteins, which are not accurately called receptors, are sensitive to mechanical or thermal changes. Physical changes in these proteins increase ion flow across the membrane, and can generate an action potential or a graded potential in the sensory neurons.

Sensory Receptors

Stimuli in the environment activate specialized receptor cells in the peripheral nervous system. Different types of stimuli are sensed by different types of receptor cells. Receptor cells can be classified into types on the basis of three different criteria: cell type, position, and function. Receptors can be classified structurally on the basis of cell type and their position in relation to stimuli they sense. They can also be classified functionally on the basis of the transduction of stimuli, or how the mechanical stimulus, light, or chemical changed the cell membrane potential.

Structural Receptor Types

The cells that interpret information about the environment can be either (1) a neuron that has a free nerve ending, with dendrites embedded in tissue that would receive a sensation; (2) a neuron that has an encapsulated ending in which the sensory nerve endings are encapsulated in connective tissue that enhances their sensitivity; or (3) a specialized receptor cell, which has distinct structural components that interpret a specific type of stimulus (Figure 1). The pain and temperature receptors in the dermis of the skin are examples of neurons that have free nerve endings. Also located in the dermis of the skin are lamellated corpuscles, neurons with encapsulated nerve endings that respond to pressure and touch. The cells in the retina that respond to light stimuli are an example of a specialized receptor, a photoreceptor.

[caption id="" align="aligncenter" width="490"]This figure shows the different types of receptors. The top panel shows a neuron receptor with free receptor endings, the middle panel shows a neuron receptor with encapsulated nerve endings, and the bottom panel shows a specialized receptor cell. Figure 1. Receptor Classification by Cell Type. Receptor cell types can be classified on the basis of their structure. Sensory neurons can have either (a) free nerve endings or (b) encapsulated endings. Photoreceptors in the eyes, such as rod cells, are examples of (c) specialized receptor cells. These cells release neurotransmitters onto a bipolar cell, which then synapses with the optic nerve neurons.[/caption]

Another way that receptors can be classified is based on their location relative to the stimuli. An exteroceptor is a receptor that is located near a stimulus in the external environment, such as the somatosensory receptors that are located in the skin. An interoceptor is one that interprets stimuli from internal organs and tissues, such as the receptors that sense the increase in blood pressure in the aorta or carotid sinus. Finally, a proprioceptor is a receptor located near a moving part of the body, such as a muscle, that interprets the positions of the tissues as they move.

Functional Receptor Types

A third classification of receptors is by how the receptor transduces stimuli into membrane potential changes. Stimuli are of three general types. Some stimuli are ions and macromolecules that affect transmembrane receptor proteins when these chemicals diffuse across the cell membrane. Some stimuli are physical variations in the environment that affect receptor cell membrane potentials. Other stimuli include the electromagnetic radiation from visible light. For humans, the only electromagnetic energy that is perceived by our eyes is visible light. Some other organisms have receptors that humans lack, such as the heat sensors of snakes, the ultraviolet light sensors of bees, or magnetic receptors in migratory birds.

Receptor cells can be further categorized on the basis of the type of stimuli they transduce. Chemical stimuli can be interpreted by a chemoreceptor that interprets chemical stimuli, such as an object’s taste or smell. Osmoreceptors respond to solute concentrations of body fluids. Additionally, pain is primarily a chemical sense that interprets the presence of chemicals from tissue damage, or similar intense stimuli, through a nociceptor. Physical stimuli, such as pressure and vibration, as well as the sensation of sound and body position (balance), are interpreted through a mechanoreceptor. Another physical stimulus that has its own type of receptor is temperature, which is sensed through a thermoreceptor that is either sensitive to temperatures above (heat) or below (cold) normal body temperature.

Sensory Modalities

Ask anyone what the senses are, and they are likely to list the five major senses—taste, smell, touch, hearing, and sight. However, these are not all of the senses. The most obvious omission from this list is balance. Also, what is referred to simply as touch can be further subdivided into pressure, vibration, stretch, and hair-follicle position, on the basis of the type of mechanoreceptors that perceive these touch sensations. Other overlooked senses include temperature perception by thermoreceptors and pain perception by nociceptors.

Within the realm of physiology, senses can be classified as either general or specific. A general sense is one that is distributed throughout the body and has receptor cells within the structures of other organs. Mechanoreceptors in the skin, muscles, or the walls of blood vessels are examples of this type. General senses often contribute to the sense of touch, as described above, or to proprioception (body movement) and kinesthesia (body movement), or to a visceral sense, which is most important to autonomic functions. A special sense is one that has a specific organ devoted to it, namely the eye, inner ear, tongue, or nose.

Each of the senses is referred to as a sensory modality. Modality refers to the way that information is encoded, which is similar to the idea of transduction. The main sensory modalities can be described on the basis of how each is transduced. The chemical senses are taste and smell. The general sense that is usually referred to as touch includes chemical sensation in the form of nociception, or pain. Pressure, vibration, muscle stretch, and the movement of hair by an external stimulus, are all sensed by mechanoreceptors. Hearing and balance are also sensed by mechanoreceptors. Finally, vision involves the activation of photoreceptors.

Listing all the different sensory modalities, which can number as many as 17, involves separating the five major senses into more specific categories, or submodalities, of the larger sense. An individual sensory modality represents the sensation of a specific type of stimulus. For example, the general sense of touch, which is known as somatosensation, can be separated into light pressure, deep pressure, vibration, itch, pain, temperature, or hair movement.

Gustation (Taste)

Only a few recognized submodalities exist within the sense of taste, or gustation. Until recently, only four tastes were recognized: sweet, salty, sour, and bitter. Research at the turn of the 20th century led to recognition of the fifth taste, umami, during the mid-1980s. Umami is a Japanese word that means “delicious taste,” and is often translated to mean savory. Very recent research has suggested that there may also be a sixth taste for fats, or lipids.

Gustation is the special sense associated with the tongue. The surface of the tongue, along with the rest of the oral cavity, is lined by a stratified squamous epithelium. Raised bumps called papillae (singular = papilla) contain the structures for gustatory transduction. There are four types of papillae, based on their appearance (Figure 2): circumvallate, foliate, filiform, and fungiform. Within the structure of the papillae are taste buds that contain specialized gustatory receptor cells for the transduction of taste stimuli. These receptor cells are sensitive to the chemicals contained within foods that are ingested, and they release neurotransmitters based on the amount of the chemical in the food. Neurotransmitters from the gustatory cells can activate sensory neurons in the facial, glossopharyngeal, and vagus cranial nerves.

[caption id="" align="aligncenter" width="525"]The left panel shows the image of a tongue with callouts that show magnified views of different parts of the tongue. The top right panel shows a micrograph of the circumvallate papilla, and the bottom right panel shows the structure of a taste bud. Figure 2. The Tongue. The tongue is covered with small bumps, called papillae, which contain taste buds that are sensitive to chemicals in ingested food or drink. Different types of papillae are found in different regions of the tongue. The taste buds contain specialized gustatory receptor cells that respond to chemical stimuli dissolved in the saliva. These receptor cells activate sensory neurons that are part of the facial and glossopharyngeal nerves. LM × 1600. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Salty taste is simply the perception of sodium ions (Na+) in the saliva. When you eat something salty, the salt crystals dissociate into the component ions Na+ and Cl–, which dissolve into the saliva in your mouth. The Na+ concentration becomes high outside the gustatory cells, creating a strong concentration gradient that drives the diffusion of the ion into the cells. The entry of Na+ into these cells results in the depolarization of the cell membrane and the generation of a receptor potential.

Sour taste is the perception of H+ concentration. Just as with sodium ions in salty flavors, these hydrogen ions enter the cell and trigger depolarization. Sour flavors are, essentially, the perception of acids in our food. Increasing hydrogen ion concentrations in the saliva (lowering saliva pH) triggers progressively stronger graded potentials in the gustatory cells. For example, orange juice—which contains citric acid—will taste sour because it has a pH value of approximately 3. Of course, it is often sweetened so that the sour taste is masked.

The first two tastes (salty and sour) are triggered by the cations Na+ and H+. The other tastes result from food molecules binding to a G protein–coupled receptor. A G protein signal transduction system ultimately leads to depolarization of the gustatory cell. The sweet taste is the sensitivity of gustatory cells to the presence of glucose dissolved in the saliva. Other monosaccharides such as fructose, or artificial sweeteners such as aspartame (NutraSweet™), saccharine, or sucralose (Splenda™) also activate the sweet receptors. The affinity for each of these molecules varies, and some will taste sweeter than glucose because they bind to the G protein–coupled receptor differently.

Bitter taste is similar to sweet in that food molecules bind to G protein–coupled receptors. However, there are a number of different ways in which this can happen because there are a large diversity of bitter-tasting molecules. Some bitter molecules depolarize gustatory cells, whereas others hyperpolarize gustatory cells. Likewise, some bitter molecules increase G protein activation within the gustatory cells, whereas other bitter molecules decrease G protein activation. The specific response depends on which molecule is binding to the receptor.

One major group of bitter-tasting molecules are alkaloids. Alkaloids are nitrogen containing molecules that are commonly found in bitter-tasting plant products, such as coffee, hops (in beer), tannins (in wine), tea, and aspirin. By containing toxic alkaloids, the plant is less susceptible to microbe infection and less attractive to herbivores.

Therefore, the function of bitter taste may primarily be related to stimulating the gag reflex to avoid ingesting poisons. Because of this, many bitter foods that are normally ingested are often combined with a sweet component to make them more palatable (cream and sugar in coffee, for example). The highest concentration of bitter receptors appear to be in the posterior tongue, where a gag reflex could still spit out poisonous food.

The taste known as umami is often referred to as the savory taste. Like sweet and bitter, it is based on the activation of G protein–coupled receptors by a specific molecule. The molecule that activates this receptor is the amino acid L-glutamate. Therefore, the umami flavor is often perceived while eating protein-rich foods. Not surprisingly, dishes that contain meat are often described as savory.

Once the gustatory cells are activated by the taste molecules, they release neurotransmitters onto the dendrites of sensory neurons. These neurons are part of the facial and glossopharyngeal cranial nerves, as well as a component within the vagus nerve dedicated to the gag reflex. The facial nerve connects to taste buds in the anterior third of the tongue. The glossopharyngeal nerve connects to taste buds in the posterior two thirds of the tongue. The vagus nerve connects to taste buds in the extreme posterior of the tongue, verging on the pharynx, which are more sensitive to noxious stimuli such as bitterness.

Olfaction (Smell)

Like taste, the sense of smell, or olfaction, is also responsive to chemical stimuli. The olfactory receptor neurons are located in a small region within the superior nasal cavity (Figure 3). This region is referred to as the olfactory epithelium and contains bipolar sensory neurons. Each olfactory sensory neuron has dendrites that extend from the apical surface of the epithelium into the mucus lining the cavity. As airborne molecules are inhaled through the nose, they pass over the olfactory epithelial region and dissolve into the mucus. These odorant molecules bind to proteins that keep them dissolved in the mucus and help transport them to the olfactory dendrites. The odorant–protein complex binds to a receptor protein within the cell membrane of an olfactory dendrite. These receptors are G protein–coupled, and will produce a graded membrane potential in the olfactory neurons.

The axon of an olfactory neuron extends from the basal surface of the epithelium, through an olfactory foramen in the cribriform plate of the ethmoid bone, and into the brain. The group of axons called the olfactory tract connect to the olfactory bulb on the ventral surface of the frontal lobe. From there, the axons split to travel to several brain regions. Some travel to the cerebrum, specifically to the primary olfactory cortex that is located in the inferior and medial areas of the temporal lobe. Others project to structures within the limbic system and hypothalamus, where smells become associated with long-term memory and emotional responses. This is how certain smells trigger emotional memories, such as the smell of food associated with one’s birthplace. Smell is the one sensory modality that does not synapse in the thalamus before connecting to the cerebral cortex. This intimate connection between the olfactory system and the cerebral cortex is one reason why smell can be a potent trigger of memories and emotion.

The nasal epithelium, including the olfactory cells, can be harmed by airborne toxic chemicals. Therefore, the olfactory neurons are regularly replaced within the nasal epithelium, after which the axons of the new neurons must find their appropriate connections in the olfactory bulb. These new axons grow along the axons that are already in place in the cranial nerve.
[caption id="" align="aligncenter" width="500"]The top left panel of this image shows the side view of a person’s face with a cup containing a beverage underneath the nose. The image shows how the aroma of the beverage passes through the nasal cavity. The top right panel shows a detailed ultrastructure of the olfactory bulb. The bottom panel shows a micrograph of the nasal cavity. Figure 3. The Olfactory System. (a) The olfactory system begins in the peripheral structures of the nasal cavity. (b) The olfactory receptor neurons are within the olfactory epithelium. (c) Axons of the olfactory receptor neurons project through the cribriform plate of the ethmoid bone and synapse with the neurons of the olfactory bulb (tissue source: simian). LM × 812. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]
[caption id="attachment_3002" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of taste and smell![/caption]

Audition (Hearing)

Hearing, or audition, is the transduction of sound waves into a neural signal that is made possible by the structures of the ear (Figure 4). The large, fleshy structure on the lateral aspect of the head is known as the auricle. Some sources will also refer to this structure as the pinna, though that term is more appropriate for a structure that can be moved, such as the external ear of a cat. The C-shaped curves of the auricle direct sound waves toward the auditory canal. The canal enters the skull through the external auditory meatus of the temporal bone. At the end of the auditory canal is the tympanic membrane, or ear drum, which vibrates after it is struck by sound waves. The auricle, ear canal, and tympanic membrane are often referred to as the external ear. The middle ear consists of a space spanned by three small bones called the ossicles. The three ossicles are the malleus, incus, and stapes, which are Latin names that roughly translate to hammer, anvil, and stirrup. The malleus is attached to the tympanic membrane and articulates with the incus. The incus, in turn, articulates with the stapes. The stapes is then attached to the inner ear, where the sound waves will be transduced into a neural signal. The middle ear is connected to the pharynx through the Eustachian tube, which helps equilibrate air pressure across the tympanic membrane. The tube is normally closed but will pop open when the muscles of the pharynx contract during swallowing or yawning.

[caption id="" align="aligncenter" width="425"]This image shows the structure of the ear with the major parts labeled. Figure 4. Structures of the Ear. The external ear contains the auricle, ear canal, and tympanic membrane. The middle ear contains the ossicles and is connected to the pharynx by the Eustachian tube. The inner ear contains the cochlea and vestibule, which are responsible for audition and equilibrium, respectively.[/caption]

The inner ear is often described as a bony labyrinth, as it is composed of a series of canals embedded within the temporal bone. It has two separate regions, the cochlea and the vestibule, which are responsible for hearing and balance, respectively. The neural signals from these two regions are relayed to the brain stem through separate fiber bundles. However, these two distinct bundles travel together from the inner ear to the brain stem as the vestibulocochlear nerve. Sound is transduced into neural signals within the cochlear region of the inner ear, which contains the sensory neurons of the spiral ganglia. These ganglia are located within the spiral-shaped cochlea of the inner ear. The cochlea is attached to the stapes through the oval window.

The oval window is located at the beginning of a fluid-filled tube within the cochlea called the scala vestibuli. The scala vestibuli extends from the oval window, travelling above the cochlear duct, which is the central cavity of the cochlea that contains the sound-transducing neurons. At the uppermost tip of the cochlea, the scala vestibuli curves over the top of the cochlear duct. The fluid-filled tube, now called the scala tympani, returns to the base of the cochlea, this time travelling under the cochlear duct. The scala tympani ends at the round window, which is covered by a membrane that contains the fluid within the scala. As vibrations of the ossicles travel through the oval window, the fluid of the scala vestibuli and scala tympani moves in a wave-like motion. The frequency of the fluid waves match the frequencies of the sound waves (Figure 5). The membrane covering the round window will bulge out or pucker in with the movement of the fluid within the scala tympani.

[caption id="" align="aligncenter" width="520"]This diagram shows how sound waves travel through the ear, and each step details the process. Figure 5. Transmission of Sound Waves to Cochlea. A sound wave causes the tympanic membrane to vibrate. This vibration is amplified as it moves across the malleus, incus, and stapes. The amplified vibration is picked up by the oval window causing pressure waves in the fluid of the scala vestibuli and scala tympani. The complexity of the pressure waves is determined by the changes in amplitude and frequency of the sound waves entering the ear.[/caption]

A cross-sectional view of the cochlea shows that the scala vestibuli and scala tympani run along both sides of the cochlear duct (Figure 6). The cochlear duct contains several organs of Corti, which tranduce the wave motion of the two scala into neural signals. The organs of Corti lie on top of the basilar membrane, which is the side of the cochlear duct located between the organs of Corti and the scala tympani. As the fluid waves move through the scala vestibuli and scala tympani, the basilar membrane moves at a specific spot, depending on the frequency of the waves. Higher frequency waves move the region of the basilar membrane that is close to the base of the cochlea. Lower frequency waves move the region of the basilar membrane that is near the tip of the cochlea.

[caption id="" align="aligncenter" width="480"]This diagram shows the structure of the cochlea in the inner ear. Figure 6. Cross Section of the Cochlea. The three major spaces within the cochlea are highlighted. The scala tympani and scala vestibuli lie on either side of the cochlear duct. The organ of Corti, containing the mechanoreceptor hair cells, is adjacent to the scala tympani, where it sits atop the basilar membrane.[/caption]

The organs of Corti contain hair cells, which are named for the hair-like stereocilia extending from the cell’s apical surfaces (Figure 7). The stereocilia are an array of microvilli-like structures arranged from tallest to shortest. Protein fibers tether adjacent hairs together within each array, such that the array will bend in response to movements of the basilar membrane. The stereocilia extend up from the hair cells to the overlying tectorial membrane, which is attached medially to the organ of Corti. When the pressure waves from the scala move the basilar membrane, the tectorial membrane slides across the stereocilia. This bends the stereocilia either toward or away from the tallest member of each array. When the stereocilia bend toward the tallest member of their array, tension in the protein tethers opens ion channels in the hair cell membrane. This will depolarize the hair cell membrane, triggering nerve impulses that travel down the afferent nerve fibers attached to the hair cells. When the stereocilia bend toward the shortest member of their array, the tension on the tethers slackens and the ion channels close. When no sound is present, and the stereocilia are standing straight, a small amount of tension still exists on the tethers, keeping the membrane potential of the hair cell slightly depolarized.

[caption id="" align="aligncenter" width="500"]This diagram shows the structure of the hair cell. The right panel shows a magnified view of the hair cell. Figure 7. Hair Cell. The hair cell is a mechanoreceptor with an array of stereocilia emerging from its apical surface. The stereocilia are tethered together by proteins that open ion channels when the array is bent toward the tallest member of their array, and closed when the array is bent toward the shortest member of their array.[/caption]
[caption id="" align="aligncenter" width="450"]This micrograph shows the ultrastructure of the cochlea. Figure 8. Cochlea and Organ of Corti. LM × 412. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

As stated above, a given region of the basilar membrane will only move if the incoming sound is at a specific frequency. Because the tectorial membrane only moves where the basilar membrane moves, the hair cells in this region will also only respond to sounds of this specific frequency. Therefore, as the frequency of a sound changes, different hair cells are activated all along the basilar membrane. The cochlea encodes auditory stimuli for frequencies between 20 and 20,000 Hz, which is the range of sound that human ears can detect. The unit of Hertz measures the frequency of sound waves in terms of cycles produced per second. Frequencies as low as 20 Hz are detected by hair cells at the apex, or tip, of the cochlea. Frequencies in the higher ranges of 20 KHz are encoded by hair cells at the base of the cochlea, close to the round and oval windows ([link]). Most auditory stimuli contain a mixture of sounds at a variety of frequencies and intensities (represented by the amplitude of the sound wave). The hair cells along the length of the cochlear duct, which are each sensitive to a particular frequency, allow the cochlea to separate auditory stimuli by frequency, just as a prism separates visible light into its component colors.

[caption id="" align="aligncenter" width="500"]This diagram shows how different sound frequencies are coded in the cochlea. Figure 9. Frequency Coding in the Cochlea. The standing sound wave generated in the cochlea by the movement of the oval window deflects the basilar membrane on the basis of the frequency of sound. Therefore, hair cells at the base of the cochlea are activated only by high frequencies, whereas those at the apex of the cochlea are activated only by low frequencies.[/caption]

Equilibrium (Balance)

Along with audition, the inner ear is responsible for encoding information about equilibrium, the sense of balance. A similar mechanoreceptor—a hair cell with stereocilia—senses head position, head movement, and whether our bodies are in motion. These cells are located within the vestibule of the inner ear. Head position is sensed by the utricle and saccule, whereas head movement is sensed by the semicircular canals. The neural signals generated in the vestibular ganglion are transmitted through the vestibulocochlear nerve to the brain stem and cerebellum.

The utricle and saccule are both largely composed of macula tissue (plural = maculae). The macula is composed of hair cells surrounded by support cells. The stereocilia of the hair cells extend into a viscous gel called the otolithic membrane (Figure 10). On top of the otolithic membrane is a layer of calcium carbonate crystals, called otoliths. The otoliths essentially make the otolithic membrane top-heavy. The otolithic membrane moves separately from the macula in response to head movements. Tilting the head causes the otolithic membrane to slide over the macula in the direction of gravity. The moving otolithic membrane, in turn, bends the sterocilia, causing some hair cells to depolarize as others hyperpolarize. The exact position of the head is interpreted by the brain based on the pattern of hair-cell depolarization.

[caption id="" align="aligncenter" width="520"]This diagram shows how the macula orients itself to allow for equilibrium. The top left panel shows the inner ear. The bottom left panel shows the cellular structure of the macula. In the top right panel, a person’s head is shown in the side view along with the orientation of the macula. In the bottom right panel, a person’s head is shown with the head tilted forward and depicts the orientation of the macula to account for the tilt. Figure 10. Linear Acceleration Coding by Maculae. The maculae are specialized for sensing linear acceleration, such as when gravity acts on the tilting head, or if the head starts moving in a straight line. The difference in inertia between the hair cell stereocilia and the otolithic membrane in which they are embedded leads to a shearing force that causes the stereocilia to bend in the direction of that linear acceleration.[/caption]

The semicircular canals are three ring-like extensions of the vestibule. One is oriented in the horizontal plane, whereas the other two are oriented in the vertical plane. The anterior and posterior vertical canals are oriented at approximately 45 degrees relative to the sagittal plane (Figure 11). The base of each semicircular canal, where it meets with the vestibule, connects to an enlarged region known as the ampulla. The ampulla contains the hair cells that respond to rotational movement, such as turning the head while saying “no.” The stereocilia of these hair cells extend into the cupula, a membrane that attaches to the top of the ampulla. As the head rotates in a plane parallel to the semicircular canal, the fluid lags, deflecting the cupula in the direction opposite to the head movement. The semicircular canals contain several ampullae, with some oriented horizontally and others oriented vertically. By comparing the relative movements of both the horizontal and vertical ampullae, the vestibular system can detect the direction of most head movements within three-dimensional (3-D) space.

[caption id="" align="aligncenter" width="520"]The left panel of this image shows a person’s head in a still position. Underneath this, the ampullary nerve is shown. The right panel shows a person rotating his head, and the below that, the direction of movement of the cupula is shown. Figure 11. Rotational Coding by Semicircular Canals. Rotational movement of the head is encoded by the hair cells in the base of the semicircular canals. As one of the canals moves in an arc with the head, the internal fluid moves in the opposite direction, causing the cupula and stereocilia to bend. The movement of two canals within a plane results in information about the direction in which the head is moving, and activation of all six canals can give a very precise indication of head movement in three dimensions.[/caption]
[caption id="attachment_3002" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of hearing and balance![/caption]

Somatosensation (Touch)

Somatosensation is considered a general sense, as opposed to the special senses discussed in this section. Somatosensation is the group of sensory modalities that are associated with touch, proprioception, and interoception. These modalities include pressure, vibration, light touch, tickle, itch, temperature, pain, proprioception, and kinesthesia. This means that its receptors are not associated with a specialized organ, but are instead spread throughout the body in a variety of organs. Many of the somatosensory receptors are located in the skin, but receptors are also found in muscles, tendons, joint capsules, ligaments, and in the walls of visceral organs.

Two types of somatosensory signals that are transduced by free nerve endings are pain and temperature. These two modalities use thermoreceptors and nociceptors to transduce temperature and pain stimuli, respectively. Temperature receptors are stimulated when local temperatures differ from body temperature. Some thermoreceptors are sensitive to just cold and others to just heat. Nociception is the sensation of potentially damaging stimuli. Mechanical, chemical, or thermal stimuli beyond a set threshold will elicit painful sensations. Stressed or damaged tissues release chemicals that activate receptor proteins in the nociceptors. For example, the sensation of heat associated with spicy foods involves capsaicin, the active molecule in hot peppers. Capsaicin molecules bind to a transmembrane ion channel in nociceptors that is sensitive to temperatures above 37°C. The dynamics of capsaicin binding with this transmembrane ion channel is unusual in that the molecule remains bound for a long time. Because of this, it will decrease the ability of other stimuli to elicit pain sensations through the activated nociceptor. For this reason, capsaicin can be used as a topical analgesic, such as in products such as Icy Hot™.

If you drag your finger across a textured surface, the skin of your finger will vibrate. Such low frequency vibrations are sensed by mechanoreceptors called Merkel cells, also known as type I cutaneous mechanoreceptors. Merkel cells are located in the stratum basale of the epidermis. Deep pressure and vibration is transduced by lamellated (Pacinian) corpuscles, which are receptors with encapsulated endings found deep in the dermis, or subcutaneous tissue. Light touch is transduced by the encapsulated endings known as tactile (Meissner) corpuscles. Follicles are also wrapped in a plexus of nerve endings known as the hair follicle plexus. These nerve endings detect the movement of hair at the surface of the skin, such as when an insect may be walking along the skin. Stretching of the skin is transduced by stretch receptors known as bulbous corpuscles. Bulbous corpuscles are also known as Ruffini corpuscles, or type II cutaneous mechanoreceptors.

Other somatosensory receptors are found in the joints and muscles. Stretch receptors monitor the stretching of tendons, muscles, and the components of joints. For example, have you ever stretched your muscles before or after exercise and noticed that you can only stretch so far before your muscles spasm back to a less stretched state? This spasm is a reflex that is initiated by stretch receptors to avoid muscle tearing. Such stretch receptors can also prevent over-contraction of a muscle. In skeletal muscle tissue, these stretch receptors are called muscle spindles. Golgi tendon organs similarly transduce the stretch levels of tendons. Bulbous corpuscles are also present in joint capsules, where they measure stretch in the components of the skeletal system within the joint. The types of nerve endings, their locations, and the stimuli they transduce are presented in Table 1.

*No corresponding eponymous name.
Mechanoreceptors of Somatosensation (Table 1)
Name Historical (eponymous) name Location(s) Stimuli
Free nerve endings * Dermis, cornea, tongue, joint capsules, visceral organs Pain, temperature, mechanical deformation
Mechanoreceptors Merkel’s discs Epidermal–dermal junction, mucosal membranes Low frequency vibration (5–15 Hz)
Bulbous corpuscle Ruffini’s corpuscle Dermis, joint capsules Stretch
Tactile corpuscle Meissner’s corpuscle Papillary dermis, especially in the fingertips and lips Light touch, vibrations below 50 Hz
Lamellated corpuscle Pacinian corpuscle Deep dermis, subcutaneous tissue Deep pressure, high-frequency vibration (around 250 Hz)
Hair follicle plexus * Wrapped around hair follicles in the dermis Movement of hair
Muscle spindle * In line with skeletal muscle fibers Muscle contraction and stretch
Tendon stretch organ Golgi tendon organ In line with tendons Stretch of tendons

Vision

Vision is the special sense of sight that is based on the transduction of light stimuli received through the eyes. The eyes are located within either orbit in the skull. The bony orbits surround the eyeballs, protecting them and anchoring the soft tissues of the eye (Figure 12). The eyelids, with lashes at their leading edges, help to protect the eye from abrasions by blocking particles that may land on the surface of the eye. The inner surface of each lid is a thin membrane known as the palpebral conjunctiva. The conjunctiva extends over the white areas of the eye (the sclera), connecting the eyelids to the eyeball. Tears are produced by the lacrimal gland, located beneath the lateral edges of the nose. Tears produced by this gland flow through the lacrimal duct to the medial corner of the eye, where the tears flow over the conjunctiva, washing away foreign particles.

[caption id="" align="aligncenter" width="430"]This diagram shows the lateral view of the eye. The major parts are labeled. Figure 12. The Eye in the Orbit. The eye is located within the orbit and surrounded by soft tissues that protect and support its function. The orbit is surrounded by cranial bones of the skull.[/caption]

Movement of the eye within the orbit is accomplished by the contraction of six extraocular muscles that originate from the bones of the orbit and insert into the surface of the eyeball (Figure 13). Four of the muscles are arranged at the cardinal points around the eye and are named for those locations. They are the superior rectus, medial rectus, inferior rectus, and lateral rectus. When each of these muscles contract, the eye to moves toward the contracting muscle. For example, when the superior rectus contracts, the eye rotates to look up. The superior oblique originates at the posterior orbit, near the origin of the four rectus muscles. However, the tendon of the oblique muscles threads through a pulley-like piece of cartilage known as the trochlea. The tendon inserts obliquely into the superior surface of the eye. The angle of the tendon through the trochlea means that contraction of the superior oblique rotates the eye medially. The inferior oblique muscle originates from the floor of the orbit and inserts into the inferolateral surface of the eye. When it contracts, it laterally rotates the eye, in opposition to the superior oblique. Rotation of the eye by the two oblique muscles is necessary because the eye is not perfectly aligned on the sagittal plane. When the eye looks up or down, the eye must also rotate slightly to compensate for the superior rectus pulling at approximately a 20-degree angle, rather than straight up. The same is true for the inferior rectus, which is compensated by contraction of the inferior oblique. A seventh muscle in the orbit is the levator palpebrae superioris, which is responsible for elevating and retracting the upper eyelid, a movement that usually occurs in concert with elevation of the eye by the superior rectus (see Figure 12).

The extraocular muscles are innervated by three cranial nerves. The lateral rectus, which causes abduction of the eye, is innervated by the abducens nerve. The superior oblique is innervated by the trochlear nerve. All of the other muscles are innervated by the oculomotor nerve, as is the levator palpebrae superioris. The motor nuclei of these cranial nerves connect to the brain stem, which coordinates eye movements.

[caption id="" align="aligncenter" width="520"]This image shows the muscles surrounding the eye. The left panel shows the lateral view, and the right panel shows the anterior view of the right eye. Figure 13. Extraocular Muscles. The extraocular muscles move the eye within the orbit.[/caption]

The eye itself is a hollow sphere composed of three layers of tissue. The outermost layer is the fibrous tunic, which includes the white sclera and clear cornea. The sclera accounts for five sixths of the surface of the eye, most of which is not visible, though humans are unique compared with many other species in having so much of the “white of the eye” visible (Figure 14). The transparent cornea covers the anterior tip of the eye and allows light to enter the eye. The middle layer of the eye is the vascular tunic, which is mostly composed of the choroid, ciliary body, and iris. The choroid is a layer of highly vascularized connective tissue that provides a blood supply to the eyeball. The choroid is posterior to the ciliary body, a muscular structure that is attached to the lens by zonule fibers. These two structures bend the lens, allowing it to focus light on the back of the eye. Overlaying the ciliary body, and visible in the anterior eye, is the iris—the colored part of the eye. The iris is a smooth muscle that opens or closes the pupil, which is the hole at the center of the eye that allows light to enter. The iris constricts the pupil in response to bright light and dilates the pupil in response to dim light. The innermost layer of the eye is the neural tunic, or retina, which contains the nervous tissue responsible for photoreception.

The eye is also divided into two cavities: the anterior cavity and the posterior cavity. The anterior cavity is the space between the cornea and lens, including the iris and ciliary body. It is filled with a watery fluid called the aqueous humor. The posterior cavity is the space behind the lens that extends to the posterior side of the interior eyeball, where the retina is located. The posterior cavity is filled with a more viscous fluid called the vitreous humor.

The retina is composed of several layers and contains specialized cells for the initial processing of visual stimuli. The photoreceptors (rods and cones) change their membrane potential when stimulated by light energy. The change in membrane potential alters the amount of neurotransmitter that the photoreceptor cells release onto bipolar cells in the outer synaptic layer. It is the bipolar cell in the retina that connects a photoreceptor to a retinal ganglion cell (RGC) in the inner synaptic layer. There, amacrine cells additionally contribute to retinal processing before an action potential is produced by the RGC. The axons of RGCs, which lie at the innermost layer of the retina, collect at the optic disc and leave the eye as the optic nerve (see Figure 14). Because these axons pass through the retina, there are no photoreceptors at the very back of the eye, where the optic nerve begins. This creates a “blind spot” in the retina, and a corresponding blind spot in our visual field.

[caption id="" align="aligncenter" width="520"]This diagram shows the structure of the eye with the major parts labeled. Figure 14. Structure of the Eye. The sphere of the eye can be divided into anterior and posterior chambers. The wall of the eye is composed of three layers: the fibrous tunic, vascular tunic, and neural tunic. Within the neural tunic is the retina, with three layers of cells and two synaptic layers in between. The center of the retina has a small indentation known as the fovea.[/caption]

Note that the photoreceptors in the retina (rods and cones) are located behind the axons, RGCs, bipolar cells, and retinal blood vessels. A significant amount of light is absorbed by these structures before the light reaches the photoreceptor cells. However, at the exact center of the retina is a small area known as the fovea. At the fovea, the retina lacks the supporting cells and blood vessels, and only contains photoreceptors. Therefore, visual acuity, or the sharpness of vision, is greatest at the fovea. This is because the fovea is where the least amount of incoming light is absorbed by other retinal structures (see Figure 14). As one moves in either direction from this central point of the retina, visual acuity drops significantly. In addition, each photoreceptor cell of the fovea is connected to a single RGC. Therefore, this RGC does not have to integrate inputs from multiple photoreceptors, which reduces the accuracy of visual transduction. Toward the edges of the retina, several photoreceptors converge on RGCs (through the bipolar cells) up to a ratio of 50 to 1. The difference in visual acuity between the fovea and peripheral retina is easily evidenced by looking directly at a word in the middle of this paragraph. The visual stimulus in the middle of the field of view falls on the fovea and is in the sharpest focus. Without moving your eyes off that word, notice that words at the beginning or end of the paragraph are not in focus. The images in your peripheral vision are focused by the peripheral retina, and have vague, blurry edges and words that are not as clearly identified. As a result, a large part of the neural function of the eyes is concerned with moving the eyes and head so that important visual stimuli are centered on the fovea.

Light falling on the retina causes chemical changes to pigment molecules in the photoreceptors, ultimately leading to a change in the activity of the RGCs. Photoreceptor cells have two parts, the inner segment and the outer segment (Figure 15). The inner segment contains the nucleus and other common organelles of a cell, whereas the outer segment is a specialized region in which photoreception takes place. There are two types of photoreceptors—rods and cones—which differ in the shape of their outer segment. The rod-shaped outer segments of the rod photoreceptor contain a stack of membrane-bound discs that contain the photosensitive pigment rhodopsin. The cone-shaped outer segments of the cone photoreceptor contain their photosensitive pigments in infoldings of the cell membrane. There are three cone photopigments, called opsins, which are each sensitive to a particular wavelength of light. The wavelength of visible light determines its color. The pigments in human eyes are specialized in perceiving three different primary colors: red, green, and blue.

[caption id="" align="aligncenter" width="450"]The top panel shows the cellular structure of the different cells in the eye. The bottom panel shows a micrograph of the cellular structure. Figure 15. Photoreceptor. (a) All photoreceptors have inner segments containing the nucleus and other important organelles and outer segments with membrane arrays containing the photosensitive opsin molecules. Rod outer segments are long columnar shapes with stacks of membrane-bound discs that contain the rhodopsin pigment. Cone outer segments are short, tapered shapes with folds of membrane in place of the discs in the rods. (b) Tissue of the retina shows a dense layer of nuclei of the rods and cones. LM × 800. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

At the molecular level, visual stimuli cause changes in the photopigment molecule that lead to changes in membrane potential of the photoreceptor cell. A single unit of light is called a photon, which is described in physics as a packet of energy with properties of both a particle and a wave. The energy of a photon is represented by its wavelength, with each wavelength of visible light corresponding to a particular color. Visible light is electromagnetic radiation with a wavelength between 380 and 720 nm. Wavelengths of electromagnetic radiation longer than 720 nm fall into the infrared range, whereas wavelengths shorter than 380 nm fall into the ultraviolet range. Light with a wavelength of 380 nm is blue whereas light with a wavelength of 720 nm is dark red. All other colors fall between red and blue at various points along the wavelength scale.

Opsin pigments are actually transmembrane proteins that contain a cofactor known as retinal. Retinal is a hydrocarbon molecule related to vitamin A. When a photon hits retinal, the long hydrocarbon chain of the molecule is biochemically altered. Specifically, photons cause some of the double-bonded carbons within the chain to switch from a cis to a trans conformation. This process is called photoisomerization. Before interacting with a photon, retinal’s flexible double-bonded carbons are in the cis conformation. This molecule is referred to as 11-cis-retinal. A photon interacting with the molecule causes the flexible double-bonded carbons to change to the trans- conformation, forming all-trans-retinal, which has a straight hydrocarbon chain (Figure 16).

The shape change of retinal in the photoreceptors initiates visual transduction in the retina. Activation of retinal and the opsin proteins result in activation of a G protein. The G protein changes the membrane potential of the photoreceptor cell, which then releases less neurotransmitter into the outer synaptic layer of the retina. Until the retinal molecule is changed back to the 11-cis-retinal shape, the opsin cannot respond to light energy, which is called bleaching. When a large group of photopigments is bleached, the retina will send information as if opposing visual information is being perceived. After a bright flash of light, afterimages are usually seen in negative. The photoisomerization is reversed by a series of enzymatic changes so that the retinal responds to more light energy.

[caption id="" align="aligncenter" width="520"]This figure shows a rod cell on the left and then shows a magnified view of the discs in the rod cells. Further magnified images show the reaction cycle required to convert cis-retinal to trans-retinal. Chemical structures of both these molecules are shown. Figure 16. Retinal Isomers. The retinal molecule has two isomers, (a) one before a photon interacts with it and (b) one that is altered through photoisomerization.[/caption]

The opsins are sensitive to limited wavelengths of light. Rhodopsin, the photopigment in rods, is most sensitive to light at a wavelength of 498 nm. The three color opsins have peak sensitivities of 564 nm, 534 nm, and 420 nm corresponding roughly to the primary colors of red, green, and blue (Figure 17). The absorbance of rhodopsin in the rods is much more sensitive than in the cone opsins; specifically, rods are sensitive to vision in low light conditions, and cones are sensitive to brighter conditions. In normal sunlight, rhodopsin will be constantly bleached while the cones are active. In a darkened room, there is not enough light to activate cone opsins, and vision is entirely dependent on rods. Rods are so sensitive to light that a single photon can result in an action potential from a rod’s corresponding RGC.

The three types of cone opsins, being sensitive to different wavelengths of light, provide us with color vision. By comparing the activity of the three different cones, the brain can extract color information from visual stimuli. For example, a bright blue light that has a wavelength of approximately 450 nm would activate the “red” cones minimally, the “green” cones marginally, and the “blue” cones predominantly. The relative activation of the three different cones is calculated by the brain, which perceives the color as blue. However, cones cannot react to low-intensity light, and rods do not sense the color of light. Therefore, our low-light vision is—in essence—in grayscale. In other words, in a dark room, everything appears as a shade of gray. If you think that you can see colors in the dark, it is most likely because your brain knows what color something is and is relying on that memory.

[caption id="" align="aligncenter" width="420"]This graph shows the normalized absorbance versus wavelength for different cell types in the eye. Figure 17. Comparison of Color Sensitivity of Photopigments. Comparing the peak sensitivity and absorbance spectra of the four photopigments suggests that they are most sensitive to particular wavelengths.[/caption]
[caption id="attachment_3004" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of vision![/caption]

Sensory Nerves

Once any sensory cell transduces a stimulus into a nerve impulse, that impulse has to travel along axons to reach the CNS. In many of the special senses, the axons leaving the sensory receptors have a topographical arrangement, meaning that the location of the sensory receptor relates to the location of the axon in the nerve. For example, in the retina, axons from RGCs in the fovea are located at the center of the optic nerve, where they are surrounded by axons from the more peripheral RGCs.

Spinal Nerves

Generally, spinal nerves contain afferent axons from sensory receptors in the periphery, such as from the skin, mixed with efferent axons travelling to the muscles or other effector organs. As the spinal nerve nears the spinal cord, it splits into dorsal and ventral roots. The dorsal root contains only the axons of sensory neurons, whereas the ventral roots contain only the axons of the motor neurons. Some of the branches will synapse with local neurons in the dorsal root ganglion, posterior (dorsal) horn, or even the anterior (ventral) horn, at the level of the spinal cord where they enter. Other branches will travel a short distance up or down the spine to interact with neurons at other levels of the spinal cord. A branch may also turn into the posterior (dorsal) column of the white matter to connect with the brain. For the sake of convenience, we will use the terms ventral and dorsal in reference to structures within the spinal cord that are part of these pathways. This will help to underscore the relationships between the different components. Typically, spinal nerve systems that connect to the brain are contralateral, in that the right side of the body is connected to the left side of the brain and the left side of the body to the right side of the brain.

Cranial Nerves

Cranial nerves convey specific sensory information from the head and neck directly to the brain. For sensations below the neck, the right side of the body is connected to the left side of the brain and the left side of the body to the right side of the brain. Whereas spinal information is contralateral, cranial nerve systems are mostly ipsilateral, meaning that a cranial nerve on the right side of the head is connected to the right side of the brain. Some cranial nerves contain only sensory axons, such as the olfactory, optic, and vestibulocochlear nerves. Other cranial nerves contain both sensory and motor axons, including the trigeminal, facial, glossopharyngeal, and vagus nerves (however, the vagus nerve is not associated with the somatic nervous system). The general senses of somatosensation for the face travel through the trigeminal system.

]]> 2553 2535 1 0 14.3 Motor Responses https://pressbooks.bccampus.ca/apdouglas2017/chapter/14-3-motor-responses/ Mon, 17 Jul 2017 19:10:35 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2568

Learning Objectives

By the end of this section, you will be able to:
  • List the components of the basic processing stream for the motor system
  • Describe the pathway of descending motor commands from the cortex to the skeletal muscles
  • Compare different descending pathways, both by structure and function
  • Explain the initiation of movement from the neurological connections
  • Describe several reflex arcs and their functional roles

The defining characteristic of the somatic nervous system is that it controls skeletal muscles. Somatic senses inform the nervous system about the external environment, but the response to that is through voluntary muscle movement. The term “voluntary” suggests that there is a conscious decision to make a movement. However, some aspects of the somatic system use voluntary muscles without conscious control. One example is the ability of our breathing to switch to unconscious control while we are focused on another task. However, the muscles that are responsible for the basic process of breathing are also utilized for speech, which is entirely voluntary.

Cortical Responses

Let’s start with sensory stimuli that have been registered through receptor cells and the information relayed to the CNS along ascending pathways. In the cerebral cortex, the initial processing of sensory perception progresses to associative processing and then integration in multimodal areas of cortex. These levels of processing can lead to the incorporation of sensory perceptions into memory, but more importantly, they lead to a response. The completion of cortical processing through the primary, associative, and integrative sensory areas initiates a similar progression of motor processing, usually in different cortical areas.

Whereas the sensory cortical areas are located in the occipital, temporal, and parietal lobes, motor functions are largely controlled by the frontal lobe. The most anterior regions of the frontal lobe—the prefrontal areas—are important for executive functions, which are those cognitive functions that lead to goal-directed behaviors. These higher cognitive processes include working memory, which has been called a “mental scratch pad,” that can help organize and represent information that is not in the immediate environment. The prefrontal lobe is responsible for aspects of attention, such as inhibiting distracting thoughts and actions so that a person can focus on a goal and direct behavior toward achieving that goal.

The functions of the prefrontal cortex are integral to the personality of an individual, because it is largely responsible for what a person intends to do and how they accomplish those plans. A famous case of damage to the prefrontal cortex is that of Phineas Gage, dating back to 1848. He was a railroad worker who had a metal spike impale his prefrontal cortex (Figure 1). He survived the accident, but according to second-hand accounts, his personality changed drastically. Friends described him as no longer acting like himself. Whereas he was a hardworking, amiable man before the accident, he turned into an irritable, temperamental, and lazy man after the accident. Many of the accounts of his change may have been inflated in the retelling, and some behavior was likely attributable to alcohol used as a pain medication. However, the accounts suggest that some aspects of his personality did change. Also, there is new evidence that though his life changed dramatically, he was able to become a functioning stagecoach driver, suggesting that the brain has the ability to recover even from major trauma such as this.

Phineas Gage
The victim of an accident while working on a railroad in 1848, Phineas Gage had a large iron rod impaled through the prefrontal cortex of his frontal lobe. After the accident, his personality appeared to change, but he eventually learned to cope with the trauma and lived as a coach driver even after such a traumatic event. (credit b: John M. Harlow, MD)
This photo shows Phineas Gage holding the metal spike that impaled his prefrontal cortex.
[caption id="" align="aligncenter" width="300"]The image on the right shows a drawing of the skull with the metal spike inserted like it would have been when he was injured. Figure 1. Phineas Gage. The victim of an accident while working on a railroad in 1848, Phineas Gage had a large iron rod impaled through the prefrontal cortex of his frontal lobe. After the accident, his personality appeared to change, but he eventually learned to cope with the trauma and lived as a coach driver even after such a traumatic event. (credit b: John M. Harlow, MD)[/caption]

Secondary Motor Cortices

In generating motor responses, the executive functions of the prefrontal cortex will need to initiate actual movements. One way to define the prefrontal area is any region of the frontal lobe that does not elicit movement when electrically stimulated. These are primarily in the anterior part of the frontal lobe. The regions of the frontal lobe that remain are the regions of the cortex that produce movement. The prefrontal areas project into the secondary motor cortices, which include the premotor cortex and the supplemental motor area.

Two important regions that assist in planning and coordinating movements are located adjacent to the primary motor cortex. The premotor cortex is more lateral, whereas the supplemental motor area is more medial and superior. The premotor area aids in controlling movements of the core muscles to maintain posture during movement, whereas the supplemental motor area is hypothesized to be responsible for planning and coordinating movement. The supplemental motor area also manages sequential movements that are based on prior experience (that is, learned movements). Neurons in these areas are most active leading up to the initiation of movement. For example, these areas might prepare the body for the movements necessary to drive a car in anticipation of a traffic light changing.

Adjacent to these two regions are two specialized motor planning centers. The frontal eye fields are responsible for moving the eyes in response to visual stimuli. There are direct connections between the frontal eye fields and the superior colliculus. Also, anterior to the premotor cortex and primary motor cortex is Broca’s area. This area is responsible for controlling movements of the structures of speech production. The area is named after a French surgeon and anatomist who studied patients who could not produce speech. They did not have impairments to understanding speech, only to producing speech sounds, suggesting a damaged or underdeveloped Broca’s area.

Primary Motor Cortex

The primary motor cortex is located in the precentral gyrus of the frontal lobe. A neurosurgeon, Walter Penfield, described much of the basic understanding of the primary motor cortex by electrically stimulating the surface of the cerebrum. Penfield would probe the surface of the cortex while the patient was only under local anesthesia so that he could observe responses to the stimulation. This led to the belief that the precentral gyrus directly stimulated muscle movement. We now know that the primary motor cortex receives input from several areas that aid in planning movement, and its principle output stimulates spinal cord neurons to stimulate skeletal muscle contraction.

The primary motor cortex is arranged in a similar fashion to the primary somatosensory cortex, in that it has a topographical map of the body, creating a motor homunculus (see Chapter 14.2 Figure 5). The neurons responsible for musculature in the feet and lower legs are in the medial wall of the precentral gyrus, with the thighs, trunk, and shoulder at the crest of the longitudinal fissure. The hand and face are in the lateral face of the gyrus. Also, the relative space allotted for the different regions is exaggerated in muscles that have greater enervation. The greatest amount of cortical space is given to muscles that perform fine, agile movements, such as the muscles of the fingers and the lower face. The “power muscles” that perform coarser movements, such as the buttock and back muscles, occupy much less space on the motor cortex.

Descending Pathways

The motor output from the cortex descends into the brain stem and to the spinal cord to control the musculature through motor neurons. Neurons located in the primary motor cortex, named Betz cells, are large cortical neurons that synapse with lower motor neurons in the spinal cord or the brain stem. The two descending pathways travelled by the axons of Betz cells are the corticospinal tract and the corticobulbar tract. Both tracts are named for their origin in the cortex and their targets—either the spinal cord or the brain stem (the term “bulbar” refers to the brain stem as the bulb, or enlargement, at the top of the spinal cord).

These two descending pathways are responsible for the conscious or voluntary movements of skeletal muscles. Any motor command from the primary motor cortex is sent down the axons of the Betz cells to activate upper motor neurons in either the cranial motor nuclei or in the ventral horn of the spinal cord. The axons of the corticobulbar tract are ipsilateral, meaning they project from the cortex to the motor nucleus on the same side of the nervous system. Conversely, the axons of the corticospinal tract are largely contralateral, meaning that they cross the midline of the brain stem or spinal cord and synapse on the opposite side of the body. Therefore, the right motor cortex of the cerebrum controls muscles on the left side of the body, and vice versa.

The corticospinal tract descends from the cortex through the deep white matter of the cerebrum. It then passes between the caudate nucleus and putamen of the basal nuclei as a bundle called the internal capsule. The tract then passes through the midbrain as the cerebral peduncles, after which it burrows through the pons. Upon entering the medulla, the tracts make up the large white matter tract referred to as the pyramids (Figure 2). The defining landmark of the medullary-spinal border is the pyramidal decussation, which is where most of the fibers in the corticospinal tract cross over to the opposite side of the brain. At this point, the tract separates into two parts, which have control over different domains of the musculature.

[caption id="" align="aligncenter" width="325"]This diagram shows how the motor neurons thread their way through the spinal cord and into the brain. It also shows the the different connections they make along the way. Figure 2. Corticospinal Tract. The major descending tract that controls skeletal muscle movements is the corticospinal tract. It is composed of two neurons, the upper motor neuron and the lower motor neuron. The upper motor neuron has its cell body in the primary motor cortex of the frontal lobe and synapses on the lower motor neuron, which is in the ventral horn of the spinal cord and projects to the skeletal muscle in the periphery.[/caption]

Appendicular Control

The lateral corticospinal tract is composed of the fibers that cross the midline at the pyramidal decussation (see Figure 2). The axons cross over from the anterior position of the pyramids in the medulla to the lateral column of the spinal cord. These axons are responsible for controlling appendicular muscles.

This influence over the appendicular muscles means that the lateral corticospinal tract is responsible for moving the muscles of the arms and legs. The ventral horn in both the lower cervical spinal cord and the lumbar spinal cord both have wider ventral horns, representing the greater number of muscles controlled by these motor neurons. The cervical enlargement is particularly large because there is greater control over the fine musculature of the upper limbs, particularly of the fingers. The lumbar enlargement is not as significant in appearance because there is less fine motor control of the lower limbs.

Axial Control

The anterior corticospinal tract is responsible for controlling the muscles of the body trunk (see Figure 2). These axons do not decussate in the medulla. Instead, they remain in an anterior position as they descend the brain stem and enter the spinal cord. These axons then travel to the spinal cord level at which they synapse with a lower motor neuron. Upon reaching the appropriate level, the axons decussate, entering the ventral horn on the opposite side of the spinal cord from which they entered. In the ventral horn, these axons synapse with their corresponding lower motor neurons. The lower motor neurons are located in the medial regions of the ventral horn, because they control the axial muscles of the trunk.

Because movements of the body trunk involve both sides of the body, the anterior corticospinal tract is not entirely contralateral. Some collateral branches of the tract will project into the ipsilateral ventral horn to control synergistic muscles on that side of the body, or to inhibit antagonistic muscles through interneurons within the ventral horn. Through the influence of both sides of the body, the anterior corticospinal tract can coordinate postural muscles in broad movements of the body. These coordinating axons in the anterior corticospinal tract are often considered bilateral, as they are both ipsilateral and contralateral.

Extrapyramidal Controls

Other descending connections between the brain and the spinal cord are called the extrapyramidal system. The name comes from the fact that this system is outside the corticospinal pathway, which includes the pyramids in the medulla. A few pathways originating from the brain stem contribute to this system.

The tectospinal tract projects from the midbrain to the spinal cord and is important for postural movements that are driven by the superior colliculus. The name of the tract comes from an alternate name for the superior colliculus, which is the tectum. The reticulospinal tract connects the reticular system, a diffuse region of gray matter in the brain stem, with the spinal cord. This tract influences trunk and proximal limb muscles related to posture and locomotion. The reticulospinal tract also contributes to muscle tone and influences autonomic functions. The vestibulospinal tract connects the brain stem nuclei of the vestibular system with the spinal cord. This allows posture, movement, and balance to be modulated on the basis of equilibrium information provided by the vestibular system.

The pathways of the extrapyramidal system are influenced by subcortical structures. For example, connections between the secondary motor cortices and the extrapyramidal system modulate spine and cranium movements. The basal nuclei, which are important for regulating movement initiated by the CNS, influence the extrapyramidal system as well as its thalamic feedback to the motor cortex.

The conscious movement of our muscles is more complicated than simply sending a single command from the precentral gyrus down to the proper motor neurons. During the movement of any body part, our muscles relay information back to the brain, and the brain is constantly sending “revised” instructions back to the muscles. The cerebellum is important in contributing to the motor system because it compares cerebral motor commands with proprioceptive feedback. The corticospinal fibers that project to the ventral horn of the spinal cord have branches that also synapse in the pons, which project to the cerebellum. Also, the proprioceptive sensations of the dorsal column system have a collateral projection to the medulla that projects to the cerebellum. These two streams of information are compared in the cerebellar cortex. Conflicts between the motor commands sent by the cerebrum and body position information provided by the proprioceptors cause the cerebellum to stimulate the red nucleus of the midbrain. The red nucleus then sends corrective commands to the spinal cord along the rubrospinal tract. The name of this tract comes from the word for red that is seen in the English word “ruby.”

A good example of how the cerebellum corrects cerebral motor commands can be illustrated by walking in water. An original motor command from the cerebrum to walk will result in a highly coordinated set of learned movements. However, in water, the body cannot actually perform a typical walking movement as instructed. The cerebellum can alter the motor command, stimulating the leg muscles to take larger steps to overcome the water resistance. The cerebellum can make the necessary changes through the rubrospinal tract. Modulating the basic command to walk also relies on spinal reflexes, but the cerebellum is responsible for calculating the appropriate response. When the cerebellum does not work properly, coordination and balance are severely affected. The most dramatic example of this is during the overconsumption of alcohol. Alcohol inhibits the ability of the cerebellum to interpret proprioceptive feedback, making it more difficult to coordinate body movements, such as walking a straight line, or guide the movement of the hand to touch the tip of the nose. [caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit this site to read about an elderly woman who starts to lose the ability to control fine movements, such as speech and the movement of limbs.[/caption]

Ventral Horn Output

The somatic nervous system provides output strictly to skeletal muscles. The lower motor neurons, which are responsible for the contraction of these muscles, are found in the ventral horn of the spinal cord. These large, multipolar neurons have a corona of dendrites surrounding the cell body and an axon that extends out of the ventral horn. This axon travels through the ventral nerve root to join the emerging spinal nerve. The axon is relatively long because it needs to reach muscles in the periphery of the body. The diameters of cell bodies may be on the order of hundreds of micrometers to support the long axon; some axons are a meter in length, such as the lumbar motor neurons that innervate muscles in the first digits of the feet.

The axons will also branch to innervate multiple muscle fibers. Together, the motor neuron and all the muscle fibers that it controls make up a motor unit. Motor units vary in size. Some may contain up to 1000 muscle fibers, such as in the quadriceps, or they may only have 10 fibers, such as in an extraocular muscle. The number of muscle fibers that are part of a motor unit corresponds to the precision of control of that muscle. Also, muscles that have finer motor control have more motor units connecting to them, and this requires a larger topographical field in the primary motor cortex.

Motor neuron axons connect to muscle fibers at a neuromuscular junction. This is a specialized synaptic structure at which multiple axon terminals synapse with the muscle fiber sarcolemma. The synaptic end bulbs of the motor neurons secrete acetylcholine, which binds to receptors on the sarcolemma. The binding of acetylcholine opens ligand-gated ion channels, increasing the movement of cations across the sarcolemma. This depolarizes the sarcolemma, initiating muscle contraction. Whereas other synapses result in graded potentials that must reach a threshold in the postsynaptic target, activity at the neuromuscular junction reliably leads to muscle fiber contraction with every nerve impulse received from a motor neuron. However, the strength of contraction and the number of fibers that contract can be affected by the frequency of the motor neuron impulses.

Reflexes

This chapter began by introducing reflexes as an example of the basic elements of the somatic nervous system. Simple somatic reflexes do not include the higher centers discussed for conscious or voluntary aspects of movement. Reflexes can be spinal or cranial, depending on the nerves and central components that are involved. The example described at the beginning of the chapter involved heat and pain sensations from a hot stove causing withdrawal of the arm through a connection in the spinal cord that leads to contraction of the biceps brachii. The description of this withdrawal reflex was simplified, for the sake of the introduction, to emphasize the parts of the somatic nervous system. But to consider reflexes fully, more attention needs to be given to this example.

As you withdraw your hand from the stove, you do not want to slow that reflex down. As the biceps brachii contracts, the antagonistic triceps brachii needs to relax. Because the neuromuscular junction is strictly excitatory, the biceps will contract when the motor nerve is active. Skeletal muscles do not actively relax. Instead the motor neuron needs to “quiet down,” or be inhibited. In the hot-stove withdrawal reflex, this occurs through an interneuron in the spinal cord. The interneuron’s cell body is located in the dorsal horn of the spinal cord. The interneuron receives a synapse from the axon of the sensory neuron that detects that the hand is being burned. In response to this stimulation from the sensory neuron, the interneuron then inhibits the motor neuron that controls the triceps brachii. This is done by releasing a neurotransmitter or other signal that hyperpolarizes the motor neuron connected to the triceps brachii, making it less likely to initiate an action potential. With this motor neuron being inhibited, the triceps brachii relaxes. Without the antagonistic contraction, withdrawal from the hot stove is faster and keeps further tissue damage from occurring.

Another example of a withdrawal reflex occurs when you step on a painful stimulus, like a tack or a sharp rock. The nociceptors that are activated by the painful stimulus activate the motor neurons responsible for contraction of the tibialis anterior muscle. This causes dorsiflexion of the foot. An inhibitory interneuron, activated by a collateral branch of the nociceptor fiber, will inhibit the motor neurons of the gastrocnemius and soleus muscles to cancel plantar flexion. An important difference in this reflex is that plantar flexion is most likely in progress as the foot is pressing down onto the tack. Contraction of the tibialis anterior is not the most important aspect of the reflex, as continuation of plantar flexion will result in further damage from stepping onto the tack.

Another type of reflex is a stretch reflex. In this reflex, when a skeletal muscle is stretched, a muscle spindle receptor is activated. The axon from this receptor structure will cause direct contraction of the muscle. A collateral of the muscle spindle fiber will also inhibit the motor neuron of the antagonist muscles. The reflex helps to maintain muscles at a constant length. A common example of this reflex is the knee jerk that is elicited by a rubber hammer struck against the patellar ligament in a physical exam.

A specialized reflex to protect the surface of the eye is the corneal reflex, or the eye blink reflex. When the cornea is stimulated by a tactile stimulus, or even by bright light in a related reflex, blinking is initiated. The sensory component travels through the trigeminal nerve, which carries somatosensory information from the face, or through the optic nerve, if the stimulus is bright light. The motor response travels through the facial nerve and innervates the orbicularis oculi on the same side. This reflex is commonly tested during a physical exam using an air puff or a gentle touch of a cotton-tipped applicator.

[caption id="attachment_3007" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about reflexes and the peripheral nervous system![/caption]
]]> 2568 2535 3 0 15.1 Divisions of the Autonomic Nervous System - 1203 https://pressbooks.bccampus.ca/apdouglas2017/chapter/15-1-divisions-of-the-autonomic-nervous-system-1203/ Mon, 17 Jul 2017 19:13:36 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2574

Learning Objectives

By the end of this section, you will be able to:
  • Name the components that generate the sympathetic and parasympathetic responses of the autonomic nervous system
  • Explain the differences in output connections within the two divisions of the autonomic nervous system
  • Describe the signaling molecules and receptor proteins involved in communication within the two divisions of the autonomic nervous system

The nervous system can be divided into two functional parts: the somatic nervous system and the autonomic nervous system. The major differences between the two systems are evident in the responses that each produces. The somatic nervous system causes contraction of skeletal muscles. The autonomic nervous system controls cardiac and smooth muscle, as well as glandular tissue. The somatic nervous system is associated with voluntary responses (though many can happen without conscious awareness, like breathing), and the autonomic nervous system is associated with involuntary responses, such as those related to homeostasis.

The autonomic nervous system regulates many of the internal organs through a balance of two aspects, or divisions. In addition to the endocrine system, the autonomic nervous system is instrumental in homeostatic mechanisms in the body. The two divisions of the autonomic nervous system are the sympathetic division and the parasympathetic division. The sympathetic system is associated with the fight-or-flight response, and parasympathetic activity is referred to by the epithet of rest and digest. Homeostasis is the balance between the two systems. At each target effector, dual innervation determines activity. For example, the heart receives connections from both the sympathetic and parasympathetic divisions. One causes heart rate to increase, whereas the other causes heart rate to decrease.

Sympathetic Division of the Autonomic Nervous System

To respond to a threat—to fight or to run away—the sympathetic system causes divergent effects as many different effector organs are activated together for a common purpose. More oxygen needs to be inhaled and delivered to skeletal muscle. The respiratory, cardiovascular, and musculoskeletal systems are all activated together. Additionally, sweating keeps the excess heat that comes from muscle contraction from causing the body to overheat. The digestive system shuts down so that blood is not absorbing nutrients when it should be delivering oxygen to skeletal muscles. To coordinate all these responses, the connections in the sympathetic system diverge from a limited region of the central nervous system (CNS) to a wide array of ganglia that project to the many effector organs simultaneously. The complex set of structures that compose the output of the sympathetic system make it possible for these disparate effectors to come together in a coordinated, systemic change.

The sympathetic division of the autonomic nervous system influences the various organ systems of the body through connections emerging from the thoracic and upper lumbar spinal cord. It is referred to as the thoracolumbar system to reflect this anatomical basis. A central neuron in the lateral horn of any of these spinal regions projects to ganglia adjacent to the vertebral column through the ventral spinal roots. The majority of ganglia of the sympathetic system belong to a network of sympathetic chain ganglia that runs alongside the vertebral column. The ganglia appear as a series of clusters of neurons linked by axonal bridges. There are typically 23 ganglia in the chain on either side of the spinal column. Three correspond to the cervical region, 12 are in the thoracic region, four are in the lumbar region, and four correspond to the sacral region. The cervical and sacral levels are not connected to the spinal cord directly through the spinal roots, but through ascending or descending connections through the bridges within the chain.

A diagram that shows the connections of the sympathetic system is somewhat like a circuit diagram that shows the electrical connections between different receptacles and devices. In Figure 1, the “circuits” of the sympathetic system are intentionally simplified.

[caption id="" align="aligncenter" width="520"]This diagram shows the spinal cord, and the connections from the spinal cord to the different target organs. The target organs are listed on the right. Figure 1. Connections of Sympathetic Division of the Autonomic Nervous System. Neurons from the lateral horn of the spinal cord (preganglionic nerve fibers - solid lines)) project to the chain ganglia on either side of the vertebral column or to collateral (prevertebral) ganglia that are anterior to the vertebral column in the abdominal cavity. Axons from these ganglionic neurons (postganglionic nerve fibers - dotted lines) then project to target effectors throughout the body.[/caption]

To continue with the analogy of the circuit diagram, there are three different types of “junctions” that operate within the sympathetic system (Figure 2). The first type is most direct: the sympathetic nerve projects to the chain ganglion at the same level as the target effector (the organ, tissue, or gland to be innervated). An example of this type is spinal nerve T1 that synapses with the T1 chain ganglion to innervate the trachea. The fibers of this branch are called white rami communicantes (singular = ramus communicans); they are myelinated and therefore referred to as white (see Figure 2a). The axon from the central neuron (the preganglionic fiber shown as a solid line) synapses with the ganglionic neuron (with the postganglionic fiber shown as a dashed line). This neuron then projects to a target effector—in this case, the trachea—via gray rami communicantes, which are unmyelinated axons.

In some cases, the target effectors are located superior or inferior to the spinal segment at which the preganglionic fiber emerges. With respect to the “wiring” involved, the synapse with the ganglionic neuron occurs at chain ganglia superior or inferior to the location of the central neuron. An example of this is spinal nerve T1 that innervates the eye. The spinal nerve tracks up through the chain until it reaches the superior cervical ganglion, where it synapses with the postganglionic neuron (see Figure 2b). The cervical ganglia are referred to as paravertebral ganglia, given their location adjacent to prevertebral ganglia in the sympathetic chain.

Not all axons from the central neurons terminate in the chain ganglia. Additional branches from the ventral nerve root continue through the chain and on to one of the collateral ganglia as the greater splanchnic nerve or lesser splanchnic nerve. For example, the greater splanchnic nerve at the level of T5 synapses with a collateral ganglion outside the chain before making the connection to the postganglionic nerves that innervate the stomach (see Figure 2c).

Collateral ganglia, also called prevertebral ganglia, are situated anterior to the vertebral column and receive inputs from splanchnic nerves as well as central sympathetic neurons. They are associated with controlling organs in the abdominal cavity, and are also considered part of the enteric nervous system. The three collateral ganglia are the celiac ganglion, the superior mesenteric ganglion, and the inferior mesenteric ganglion (see Figure 1). The word celiac is derived from the Latin word “coelom,” which refers to a body cavity (in this case, the abdominal cavity), and the word mesenteric refers to the digestive system.

[caption id="" align="aligncenter" width="520"]This table shows the connections between the spinal cord and the ganglia. The top panel shows the connection between a central neuron and a chain ganglion at the same lever. The center panel shows the connection between a central neuron and a synapse with a superior or inferior ganglion. The bottom panel shows the projection of a central neuron into the white ramus. Figure 2. Sympathetic Connections and Chain Ganglia. The axon from a central sympathetic neuron in the spinal cord can project to the periphery in a number of different ways. (a) The fiber can project out to the ganglion at the same level and synapse on a ganglionic neuron. (b) A branch can project to more superior or inferior ganglion in the chain. (c) A branch can project through the white ramus communicans, but not terminate on a ganglionic neuron in the chain. Instead, it projects through one of the splanchnic nerves to a collateral ganglion or the adrenal medulla (not pictured).[/caption]
An axon from the central neuron that projects to a sympathetic ganglion is referred to as a preganglionic fiber or neuron, and represents the output from the CNS to the ganglion. Because the sympathetic ganglia are adjacent to the vertebral column, preganglionic sympathetic fibers are relatively short, and they are myelinated. A postganglionic fiber—the axon from a ganglionic neuron that projects to the target effector—represents the output of a ganglion that directly influences the organ. Compared with the preganglionic fibers, postganglionic sympathetic fibers are long because of the relatively greater distance from the ganglion to the target effector. These fibers are unmyelinated. (Note that the term “postganglionic neuron” may be used to describe the projection from a ganglion to the target. The problem with that usage is that the cell body is in the ganglion, and only the fiber is postganglionic. Typically, the term neuron applies to the entire cell.)

One type of preganglionic sympathetic fiber does not terminate in a ganglion. These are the axons from central sympathetic neurons that project to the adrenal medulla, the interior portion of the adrenal gland. These axons are still referred to as preganglionic fibers, but the target is not a ganglion. The adrenal medulla releases signaling molecules into the bloodstream, rather than using axons to communicate with target structures. The cells in the adrenal medulla that are contacted by the preganglionic fibers are called chromaffin cells. These cells are neurosecretory cells that develop from the neural crest along with the sympathetic ganglia, reinforcing the idea that the gland is, functionally, a sympathetic ganglion.

The projections of the sympathetic division of the autonomic nervous system diverge widely, resulting in a broad influence of the system throughout the body. As a response to a threat, the sympathetic system would increase heart rate and breathing rate and cause blood flow to the skeletal muscle to increase and blood flow to the digestive system to decrease. Sweat gland secretion should also increase as part of an integrated response. All of those physiological changes are going to be required to occur together to run away from the hunting lioness, or the modern equivalent. This divergence is seen in the branching patterns of preganglionic sympathetic neurons—a single preganglionic sympathetic neuron may have 10–20 targets. An axon that leaves a central neuron of the lateral horn in the thoracolumbar spinal cord will pass through the white ramus communicans and enter the sympathetic chain, where it will branch toward a variety of targets. At the level of the spinal cord at which the preganglionic sympathetic fiber exits the spinal cord, a branch will synapse on a neuron in the adjacent chain ganglion. Some branches will extend up or down to a different level of the chain ganglia. Other branches will pass through the chain ganglia and project through one of the splanchnic nerves to a collateral ganglion. Finally, some branches may project through the splanchnic nerves to the adrenal medulla. All of these branches mean that one preganglionic neuron can influence different regions of the sympathetic system very broadly, by acting on widely distributed organs.

Parasympathetic Division of the Autonomic Nervous System

The parasympathetic division of the autonomic nervous system is named because its central neurons are located on either side of the thoracolumbar region of the spinal cord (para- = “beside” or “near”). The parasympathetic system can also be referred to as the craniosacral system (or outflow) because the preganglionic neurons are located in nuclei of the brain stem and the lateral horn of the sacral spinal cord.

The connections, or “circuits,” of the parasympathetic division are similar to the general layout of the sympathetic division with a few specific differences (Figure 3). The preganglionic fibers from the cranial region travel in cranial nerves, whereas preganglionic fibers from the sacral region travel in spinal nerves. The targets of these fibers are terminal ganglia, which are located near—or even within—the target effector. These ganglia are often referred to as intramural ganglia when they are found within the walls of the target organ. The postganglionic fiber projects from the terminal ganglia a short distance to the target effector, or to the specific target tissue within the organ. Comparing the relative lengths of axons in the parasympathetic system, the preganglionic fibers are long and the postganglionic fibers are short because the ganglia are close to—and sometimes within—the target effectors.

The cranial component of the parasympathetic system is based in particular nuclei of the brain stem. In the midbrain, the Edinger–Westphal nucleus is part of the oculomotor complex, and axons from those neurons travel with the fibers in the oculomotor nerve (cranial nerve III) that innervate the extraocular muscles. The preganglionic parasympathetic fibers within cranial nerve III terminate in the ciliary ganglion, which is located in the posterior orbit. The postganglionic parasympathetic fibers then project to the smooth muscle of the iris to control pupillary size. In the upper medulla, the salivatory nuclei contain neurons with axons that project through the facial and glossopharyngeal nerves to ganglia that control salivary glands. Tear production is influenced by parasympathetic fibers in the facial nerve, which activate a ganglion, and ultimately the lacrimal (tear) gland. Neurons in the dorsal nucleus of the vagus nerve and the nucleus ambiguus project through the vagus nerve (cranial nerve X) to the terminal ganglia of the thoracic and abdominal cavities. Parasympathetic preganglionic fibers primarily influence the heart, bronchi, and esophagus in the thoracic cavity and the stomach, liver, pancreas, gall bladder, and small intestine of the abdominal cavity. The postganglionic fibers from the ganglia activated by the vagus nerve are often incorporated into the structure of the organ, such as the mesenteric plexus of the digestive tract organs and the intramural ganglia.

[caption id="" align="aligncenter" width="520"]This diagram shows the spinal cord and has different central nerves emerging from it. The central nerves target different effector organs that are listed on the right. Figure 3. Connections of Parasympathetic Division of the Autonomic Nervous System. Neurons from brain-stem nuclei, or from the lateral horn of the sacral spinal cord, project to terminal ganglia near or within the various organs of the body. Axons from these ganglionic neurons then project the short distance to those target effectors.[/caption]

Chemical Signaling in the Autonomic Nervous System

Where an autonomic neuron connects with a target, there is a synapse. The electrical signal of the action potential causes the release of a signaling molecule, which will bind to receptor proteins on the target cell. Synapses of the autonomic system are classified as either cholinergic, meaning that acetylcholine (ACh) is released, or adrenergic, meaning that norepinephrine is released. The terms cholinergic and adrenergic refer not only to the signaling molecule that is released but also to the class of receptors that each binds.

The cholinergic system includes two classes of receptor: the nicotinic receptor and the muscarinic receptor. Both receptor types bind to ACh and cause changes in the target cell. The nicotinic receptor is a ligand-gated cation channel and the muscarinic receptor is a G protein–coupled receptor. The receptors are named for, and differentiated by, other molecules that bind to them. Whereas nicotine will bind to the nicotinic receptor, and muscarine will bind to the muscarinic receptor, there is no cross-reactivity between the receptors. The situation is similar to locks and keys. Imagine two locks—one for a classroom and the other for an office—that are opened by two separate keys. The classroom key will not open the office door and the office key will not open the classroom door. This is similar to the specificity of nicotine and muscarine for their receptors. However, a master key can open multiple locks, such as a master key for the Biology Department that opens both the classroom and the office doors. This is similar to ACh that binds to both types of receptors. The molecules that define these receptors are not crucial—they are simply tools for researchers to use in the laboratory. These molecules are exogenous, meaning that they are made outside of the human body, so a researcher can use them without any confounding endogenous results (results caused by the molecules produced in the body).

The adrenergic system also has two types of receptors, named the alpha (α)-adrenergic receptor and beta (β)-adrenergic receptor. Unlike cholinergic receptors, these receptor types are not classified by which drugs can bind to them. All of them are G protein–coupled receptors. There are three types of α-adrenergic receptors, termed α1, α2, and α3, and there are two types of β-adrenergic receptors, termed β1 and β2. An additional aspect of the adrenergic system is that there is a second signaling molecule called epinephrine. The chemical difference between norepinephrine and epinephrine is the addition of a methyl group (CH3) in epinephrine. The prefix “nor-” actually refers to this chemical difference, in which a methyl group is missing.

The term adrenergic should remind you of the word adrenaline, which is associated with the fight-or-flight response described at the beginning of the chapter. Adrenaline and epinephrine are two names for the same molecule. The adrenal gland (in Latin, ad- = “on top of”; renal = “kidney”) secretes adrenaline. The ending “-ine” refers to the chemical being derived, or extracted, from the adrenal gland. A similar construction from Greek instead of Latin results in the word epinephrine (epi- = “above”; nephr- = “kidney”). In scientific usage, epinephrine is preferred in the United States, whereas adrenaline is preferred in Great Britain, because “adrenalin” was once a registered, proprietary drug name in the United States. Though the drug is no longer sold, the convention of referring to this molecule by the two different names persists. Similarly, norepinephrine and noradrenaline are two names for the same molecule.

Having understood the cholinergic and adrenergic systems, their role in the autonomic system is relatively simple to understand. All preganglionic fibers, both sympathetic and parasympathetic, release ACh. All ganglionic neurons—the targets of these preganglionic fibers—have nicotinic receptors in their cell membranes. The nicotinic receptor is a ligand-gated cation channel that results in depolarization of the postsynaptic membrane. The postganglionic parasympathetic fibers also release ACh, but the receptors on their targets are muscarinic receptors, which are G protein–coupled receptors and do not exclusively cause depolarization of the postsynaptic membrane. Postganglionic sympathetic fibers release norepinephrine, except for fibers that project to sweat glands and to blood vessels associated with skeletal muscles, which release ACh (table 1).

Autonomic System Signaling Molecules (Table 1)
Sympathetic Parasympathetic
Preganglionic Acetylcholine → nicotinic receptor Acetylcholine → nicotinic receptor
Postganglionic Norepinephrine → α- or β-adrenergic receptors
Acetylcholine → muscarinic receptor (associated with sweat glands and the blood vessels associated with skeletal muscles only		 Acetylcholine → muscarinic receptor

Signaling molecules can belong to two broad groups. Neurotransmitters are released at synapses, whereas hormones are released into the bloodstream. These are simplistic definitions, but they can help to clarify this point. Acetylcholine can be considered a neurotransmitter because it is released by axons at synapses. The adrenergic system, however, presents a challenge. Postganglionic sympathetic fibers release norepinephrine, which can be considered a neurotransmitter. But the adrenal medulla releases epinephrine and norepinephrine into circulation, so they should be considered hormones.

What are referred to here as synapses may not fit the strictest definition of synapse. Some sources will refer to the connection between a postganglionic fiber and a target effector as neuroeffector junctions; neurotransmitters, as defined above, would be called neuromodulators. The structure of postganglionic connections are not the typical synaptic end bulb that is found at the neuromuscular junction, but rather are chains of swellings along the length of a postganglionic fiber called a varicosity (Figure 4).

[caption id="" align="aligncenter" width="520"]This figure shows the connection between autonomic fibers and the target effectors. The left image shows a slice of smooth muscle with the postganglionic varicosities and the postganglionic axons labeled. The right panel shows a magnified view of the synaptic vesicles, neurotransmitters, and the sarcolemma. Figure 4. Autonomic Varicosities. The connection between autonomic fibers and target effectors is not the same as the typical synapse, such as the neuromuscular junction. Instead of a synaptic end bulb, a neurotransmitter is released from swellings along the length of a fiber that makes an extended network of connections in the target effector.[/caption]
[caption id="attachment_3009" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of the autonomic nervous system![/caption]
]]> 2574 2569 1 0 17.1 An Overview of the Endocrine System https://pressbooks.bccampus.ca/apdouglas2017/chapter/17-1-an-overview-of-the-endocrine-system/ Mon, 17 Jul 2017 19:14:52 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2588

Learning Objectives

By the end of this section, you will be able to:
  • Distinguish the types of intercellular communication, their importance, mechanisms, and effects
  • Identify the major organs and tissues of the endocrine system and their location in the body

Communication is a process in which a sender transmits signals to one or more receivers to control and coordinate actions. In the human body, two major organ systems participate in relatively “long distance” communication: the nervous system and the endocrine system. Together, these two systems are primarily responsible for maintaining homeostasis in the body.

Neural and Endocrine Signaling

The nervous system uses two types of intercellular communication—electrical and chemical signaling—either by the direct action of an electrical potential, or in the latter case, through the action of chemical neurotransmitters such as serotonin or norepinephrine. Neurotransmitters act locally and rapidly. When an electrical signal in the form of an action potential arrives at the synaptic terminal, they diffuse across the synaptic cleft (the gap between a sending neuron and a receiving neuron or muscle cell). Once the neurotransmitters interact (bind) with receptors on the receiving (post-synaptic) cell, the receptor stimulation is transduced into a response such as continued electrical signaling or modification of cellular response. The target cell responds within milliseconds of receiving the chemical “message”; this response then ceases very quickly once the neural signaling ends. In this way, neural communication enables body functions that involve quick, brief actions, such as movement, sensation, and cognition.In contrast, the endocrine system uses just one method of communication: chemical signaling. These signals are sent by the endocrine organs, which secrete chemicals—the hormone—into the extracellular fluid. Hormones are transported primarily via the bloodstream throughout the body, where they bind to receptors on target cells, inducing a characteristic response. As a result, endocrine signaling requires more time than neural signaling to prompt a response in target cells, though the precise amount of time varies with different hormones. For example, the hormones released when you are confronted with a dangerous or frightening situation, called the fight-or-flight response, occur by the release of adrenal hormones—epinephrine and norepinephrine—within seconds. In contrast, it may take up to 48 hours for target cells to respond to certain reproductive hormones.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit this link to watch an animation of the events that occur when a hormone binds to a cell membrane receptor.[/caption]

In addition, endocrine signaling is typically less specific than neural signaling. The same hormone may play a role in a variety of different physiological processes depending on the target cells involved. For example, the hormone oxytocin promotes uterine contractions in women in labor. It is also important in breastfeeding, and may be involved in the sexual response and in feelings of emotional attachment in both males and females.

In general, the nervous system involves quick responses to rapid changes in the external environment, and the endocrine system is usually slower acting—taking care of the internal environment of the body, maintaining homeostasis, and controlling reproduction (Table 1). So how does the fight-or-flight response that was mentioned earlier happen so quickly if hormones are usually slower acting? It is because the two systems are connected. It is the fast action of the nervous system in response to the danger in the environment that stimulates the adrenal glands to secrete their hormones. As a result, the nervous system can cause rapid endocrine responses to keep up with sudden changes in both the external and internal environments when necessary.

Endocrine and Nervous Systems (Table 1)
Endocrine system Nervous system
Signaling mechanism(s) Chemical Chemical/electrical
Primary chemical signal Hormones Neurotransmitters
Distance traveled Long or short Always short
Response time Fast or slow Always fast
Environment targeted Internal Internal and external

Structures of the Endocrine System

The endocrine system consists of cells, tissues, and organs that secrete hormones as a primary or secondary function. The endocrine gland is the major player in this system. The primary function of these ductless glands is to secrete their hormones directly into the surrounding fluid. The interstitial fluid and the blood vessels then transport the hormones throughout the body. The endocrine system includes the pituitary, thyroid, parathyroid, adrenal, and pineal glands (Figure 1). Some of these glands have both endocrine and non-endocrine functions. For example, the pancreas contains cells that function in digestion as well as cells that secrete the hormones insulin and glucagon, which regulate blood glucose levels. The hypothalamus, thymus, heart, kidneys, stomach, small intestine, liver, skin, female ovaries, and male testes are other organs that contain cells with endocrine function. Moreover, adipose tissue has long been known to produce hormones, and recent research has revealed that even bone tissue has endocrine functions.

[caption id="" align="aligncenter" width="520"]This diagram shows the endocrine glands and cells that are located throughout the body. The endocrine system organs include the pineal gland and pituitary gland in the brain. The pituitary is located on the anterior side of the thalamus while the pineal gland is located on the posterior side of the thalamus. The thyroid gland is a butterfly-shaped gland that wraps around the trachea within the neck. Four small, disc-shaped parathyroid glands are embedded into the posterior side of the thyroid. The adrenal glands are located on top of the kidneys. The pancreas is located at the center of the abdomen. In females, the two ovaries are connected to the uterus by two long, curved, tubes in the pelvic region. In males, the two testes are located in the scrotum below the penis. Figure 1. Endocrine System. Endocrine glands and cells are located throughout the body and play an important role in homeostasis.[/caption]

The ductless endocrine glands are not to be confused with the body’s exocrine system, whose glands release their secretions through ducts. Examples of exocrine glands include the sebaceous and sweat glands of the skin. As just noted, the pancreas also has an exocrine function: most of its cells secrete pancreatic juice through the pancreatic and accessory ducts to the lumen of the small intestine.

Other Types of Chemical Signaling

In endocrine signaling, hormones secreted into the extracellular fluid diffuse into the blood or lymph, and can then travel great distances throughout the body. In contrast, autocrine signaling takes place within the same cell. An autocrine (auto- = “self”) is a chemical that elicits a response in the same cell that secreted it. Interleukin-1, or IL-1, is a signaling molecule that plays an important role in inflammatory response. The cells that secrete IL-1 have receptors on their cell surface that bind these molecules, resulting in autocrine signaling.

Local intercellular communication is the province of the paracrine, also called a paracrine factor, which is a chemical that induces a response in neighboring cells. Although paracrines may enter the bloodstream, their concentration is generally too low to elicit a response from distant tissues. A familiar example to those with asthma is histamine, a paracrine that is released by immune cells in the bronchial tree. Histamine causes the smooth muscle cells of the bronchi to constrict, narrowing the airways. Another example is the neurotransmitters of the nervous system, which act only locally within the synaptic cleft.

[caption id="attachment_3011" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of the endocrine system![/caption]
]]> 2588 2585 1 0 22.5 Transport of Gases https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-5-transport-of-gases-2/ Mon, 17 Jul 2017 19:16:40 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2630

Learning Objectives

By the end of this section, you will be able to:
  • Describe the principles of oxygen transport
  • Describe the structure of hemoglobin
  • Compare and contrast fetal and adult hemoglobin
  • Describe the principles of carbon dioxide transport
The other major activity in the lungs is the process of respiration, the process of gas exchange. The function of respiration is to provide oxygen for use by body cells during cellular respiration and to eliminate carbon dioxide, a waste product of cellular respiration, from the body. In order for the exchange of oxygen and carbon dioxide to occur, both gases must be transported between the external and internal respiration sites. Although carbon dioxide is more soluble than oxygen in blood, both gases require a specialized transport system for the majority of the gas molecules to be moved between the lungs and other tissues.

Oxygen Transport in the Blood

Even though oxygen is transported via the blood, you may recall that oxygen is not very soluble in liquids. A small amount of oxygen does dissolve in the blood and is transported in the bloodstream, but it is only about 1.5% of the total amount. The majority of oxygen molecules are carried from the lungs to the body’s tissues by a specialized transport system, which relies on the erythrocyte—the red blood cell. Erythrocytes contain a metalloprotein, hemoglobin, which serves to bind oxygen molecules to the erythrocyte (Figure 1). Heme is the portion of hemoglobin that contains iron, and it is heme that binds oxygen. One hemoglobin molecule contains iron-containing Heme molecules, and because of this, each hemoglobin molecule is capable of carrying up to four molecules of oxygen. As oxygen diffuses across the respiratory membrane from the alveolus to the capillary, it also diffuses into the red blood cell and is bound by hemoglobin. The following reversible chemical reaction describes the production of the final product, oxyhemoglobin (Hb–O2), which is formed when oxygen binds to hemoglobin. Oxyhemoglobin is a bright red-colored molecule that contributes to the bright red color of oxygenated blood.
Hb + O2 ↔ Hb − O2
In this formula, Hb represents reduced hemoglobin, that is, hemoglobin that does not have oxygen bound to it. There are multiple factors involved in how readily heme binds to and dissociates from oxygen, which will be discussed in the subsequent sections.
[caption id="" align="aligncenter" width="300"]This diagram shows a red blood cell and the structure of a hemoglobin molecule. Figure 1. Erythrocyte and Hemoglobin. Hemoglobin consists of four subunits, each of which contains one molecule of iron.[/caption]

Function of Hemoglobin

Hemoglobin is composed of subunits, a protein structure that is referred to as a quaternary structure. Each of the four subunits that make up hemoglobin is arranged in a ring-like fashion, with an iron atom covalently bound to the heme in the center of each subunit. Binding of the first oxygen molecule causes a conformational change in hemoglobin that allows the second molecule of oxygen to bind more readily. As each molecule of oxygen is bound, it further facilitates the binding of the next molecule, until all four heme sites are occupied by oxygen. The opposite occurs as well: After the first oxygen molecule dissociates and is “dropped off” at the tissues, the next oxygen molecule dissociates more readily. When all four heme sites are occupied, the hemoglobin is said to be saturated. When one to three heme sites are occupied, the hemoglobin is said to be partially saturated. Therefore, when considering the blood as a whole, the percent of the available heme units that are bound to oxygen at a given time is called hemoglobin saturation. Hemoglobin saturation of 100 percent means that every heme unit in all of the erythrocytes of the body is bound to oxygen. In a healthy individual with normal hemoglobin levels, hemoglobin saturation generally ranges from 95 percent to 99 percent.

Oxygen Dissociation from Hemoglobin

Partial pressure is an important aspect of the binding of oxygen to and disassociation from heme. An oxygen–hemoglobin dissociation curve is a graph that describes the relationship of partial pressure to the binding of oxygen to heme and its subsequent dissociation from heme (Figure 2). Remember that gases travel from an area of higher partial pressure to an area of lower partial pressure. In addition, the affinity of an oxygen molecule for heme increases as more oxygen molecules are bound. Therefore, in the oxygen–hemoglobin saturation curve, as the partial pressure of oxygen increases, a proportionately greater number of oxygen molecules are bound by heme. Not surprisingly, the oxygen–hemoglobin saturation/dissociation curve also shows that the lower the partial pressure of oxygen, the fewer oxygen molecules are bound to heme. As a result, the partial pressure of oxygen plays a major role in determining the degree of binding of oxygen to heme at the site of the respiratory membrane, as well as the degree of dissociation of oxygen from heme at the site of body tissues.
The top panel of this figure shows a graph with oxygen saturation of the y-axis and partial pressure of oxygen on the x-axis.The middle panel shows oxygen saturation versus partial pressure of oxygen as a function of pH. [caption id="" align="aligncenter" width="400"]The bottom panel shows the same relationship as a function of temperature. Figure 2. Oxygen-Hemoglobin Dissociation and Effects of pH and Temperature. These three graphs show (a) the relationship between the partial pressure of oxygen and hemoglobin saturation, (b) the effect of pH on the oxygen–hemoglobin dissociation curve, and (c) the effect of temperature on the oxygen–hemoglobin dissociation curve.[/caption]
The mechanisms behind the oxygen–hemoglobin saturation/dissociation curve also serve as automatic control mechanisms that regulate how much oxygen is delivered to different tissues throughout the body. This is important because some tissues have a higher metabolic rate than others. Highly active tissues, such as muscle, rapidly use oxygen to produce ATP, lowering the partial pressure of oxygen in the tissue to about 20 mm Hg. The partial pressure of oxygen inside capillaries is about 100 mm Hg, so the difference between the two becomes quite high, about 80 mm Hg. As a result, a greater number of oxygen molecules dissociate from hemoglobin and enter the tissues. The reverse is true of tissues, such as adipose (body fat), which have lower metabolic rates. Because less oxygen is used by these cells, the partial pressure of oxygen within such tissues remains relatively high, resulting in fewer oxygen molecules dissociating from hemoglobin and entering the tissue interstitial fluid. Although venous blood is said to be deoxygenated, some oxygen is still bound to hemoglobin in its red blood cells. This provides an oxygen reserve that can be used when tissues suddenly demand more oxygen. Factors other than partial pressure also affect the oxygen–hemoglobin saturation/dissociation curve. For example, a higher temperature promotes hemoglobin and oxygen to dissociate faster, whereas a lower temperature inhibits dissociation (see Figure 2, middle). However, the human body tightly regulates temperature, so this factor may not affect gas exchange throughout the body. The exception to this is in highly active tissues, which may release a larger amount of energy than is given off as heat. As a result, oxygen readily dissociates from hemoglobin, which is a mechanism that helps to provide active tissues with more oxygen. Certain hormones, such as androgens, epinephrine, thyroid hormones, and growth hormone, can affect the oxygen–hemoglobin saturation/disassociation curve by stimulating the production of a compound called 2,3-bisphosphoglycerate (BPG) by erythrocytes. BPG is a byproduct of glycolysis. Because erythrocytes do not contain mitochondria, glycolysis is the sole method by which these cells produce ATP. BPG promotes the disassociation of oxygen from hemoglobin. Therefore, the greater the concentration of BPG, the more readily oxygen dissociates from hemoglobin, despite its partial pressure. The pH of the blood is another factor that influences the oxygen–hemoglobin saturation/dissociation curve (see Figure 2). The Bohr effect is a phenomenon that arises from the relationship between pH and oxygen’s affinity for hemoglobin: A lower, more acidic pH promotes oxygen dissociation from hemoglobin. In contrast, a higher, or more basic, pH inhibits oxygen dissociation from hemoglobin. The greater the amount of carbon dioxide in the blood, the more molecules that must be converted, which in turn generates hydrogen ions and thus lowers blood pH. Furthermore, blood pH may become more acidic when certain byproducts of cell metabolism, such as lactic acid, carbonic acid, and carbon dioxide, are released into the bloodstream.

Hemoglobin of the Fetus

The fetus has its own circulation with its own erythrocytes; however, it is dependent on the mother for oxygen. Blood is supplied to the fetus by way of the umbilical cord, which is connected to the placenta and separated from maternal blood by the chorion. The mechanism of gas exchange at the chorion is similar to gas exchange at the respiratory membrane. However, the partial pressure of oxygen is lower in the maternal blood in the placenta, at about 35 to 50 mm Hg, than it is in maternal arterial blood. The difference in partial pressures between maternal and fetal blood is not large, as the partial pressure of oxygen in fetal blood at the placenta is about 20 mm Hg. Therefore, there is not as much diffusion of oxygen into the fetal blood supply. The fetus’ hemoglobin overcomes this problem by having a greater affinity for oxygen than maternal hemoglobin (Figure 3). Both fetal and adult hemoglobin have four subunits, but two of the subunits of fetal hemoglobin have a different structure that causes fetal hemoglobin to have a greater affinity for oxygen than does adult hemoglobin.
[caption id="" align="aligncenter" width="430"]This graph shows the oxygen saturation versus the partial pressure of oxygen in fetal hemoglobin and adult hemoglobin. Figure 3. Oxygen-Hemoglobin Dissociation Curves in Fetus and Adult. Fetal hemoglobin has a greater affinity for oxygen than does adult hemoglobin.[/caption]

Carbon Dioxide Transport in the Blood

Carbon dioxide is transported by three major mechanisms. The first mechanism of carbon dioxide transport is by blood plasma, as some carbon dioxide molecules dissolve in the blood. The second mechanism is transport in the form of bicarbonate (HCO3–), which also dissolves in plasma. The third mechanism of carbon dioxide transport is similar to the transport of oxygen by erythrocytes (Figure 4).
[caption id="" align="aligncenter" width="435"]This figure shows how carbon dioxide is transported from the tissue to the red blood cell. Figure 4: Carbon Dioxide Transport Carbon dioxide is transported by three different methods: (a) in erythrocytes; (b) after forming carbonic acid (H2CO3 ), which is dissolved in plasma; (c) and in plasma.[/caption]

Dissolved Carbon Dioxide

Although carbon dioxide is not considered to be highly soluble in blood, a small fraction—about 7 to 10 percent—of the carbon dioxide that diffuses into the blood from the tissues dissolves in plasma. The dissolved carbon dioxide then travels in the bloodstream and when the blood reaches the pulmonary capillaries, the dissolved carbon dioxide diffuses across the respiratory membrane into the alveoli, where it is then exhaled during pulmonary ventilation.

Bicarbonate Buffer

A large fraction—about 70 percent—of the carbon dioxide molecules that diffuse into the blood is transported to the lungs as bicarbonate. Most bicarbonate is produced in erythrocytes after carbon dioxide diffuses into the capillaries, and subsequently into red blood cells. Carbonic anhydrase (CA) causes carbon dioxide and water to form carbonic acid (H2CO3), which dissociates into two ions: bicarbonate (HCO3–) and hydrogen (H+). The following formula depicts this reaction:
CO2 + H2O CA ↔ H2CO3↔H+ + HCO3−
Bicarbonate tends to build up in the erythrocytes, so that there is a greater concentration of bicarbonate in the erythrocytes than in the surrounding blood plasma. As a result, some of the bicarbonate will leave the erythrocytes and move down its concentration gradient into the plasma in exchange for chloride (Cl–) ions. This phenomenon is referred to as the chloride shift and occurs because by exchanging one negative ion for another negative ion, neither the electrical charge of the erythrocytes nor that of the blood is altered. At the pulmonary capillaries, the chemical reaction that produced bicarbonate (shown above) is reversed, and carbon dioxide and water are the products. Much of the bicarbonate in the plasma re-enters the erythrocytes in exchange for chloride ions. Hydrogen ions and bicarbonate ions join to form carbonic acid, which is converted into carbon dioxide and water by carbonic anhydrase. Carbon dioxide diffuses out of the erythrocytes and into the plasma, where it can further diffuse across the respiratory membrane into the alveoli to be exhaled during pulmonary ventilation.

Carbaminohemoglobin

About 20 percent of carbon dioxide is bound by hemoglobin and is transported to the lungs. Carbon dioxide does not bind to iron as oxygen does; instead, carbon dioxide binds amino acid moieties on the globin portions of hemoglobin to form carbaminohemoglobin, which forms when hemoglobin and carbon dioxide bind. When hemoglobin is not transporting oxygen, it tends to have a bluish-purple tone to it, creating the darker maroon color typical of deoxygenated blood. The following formula depicts this reversible reaction:
CO2 + Hb ↔ HbCO2
Similar to the transport of oxygen by heme, the binding and dissociation of carbon dioxide to and from hemoglobin is dependent on the partial pressure of carbon dioxide. Because carbon dioxide is released from the lungs, blood that leaves the lungs and reaches body tissues has a lower partial pressure of carbon dioxide than is found in the tissues. As a result, carbon dioxide leaves the tissues because of its higher partial pressure, enters the blood, and then moves into red blood cells, binding to hemoglobin. In contrast, in the pulmonary capillaries, the partial pressure of carbon dioxide is high compared to within the alveoli. As a result, carbon dioxide dissociates readily from hemoglobin and diffuses across the respiratory membrane into the air. In addition to the partial pressure of carbon dioxide, the oxygen saturation of hemoglobin and the partial pressure of oxygen in the blood also influence the affinity of hemoglobin for carbon dioxide. The Haldane effect is a phenomenon that arises from the relationship between the partial pressure of oxygen and the affinity of hemoglobin for carbon dioxide. Hemoglobin that is saturated with oxygen does not readily bind carbon dioxide. However, when oxygen is not bound to heme and the partial pressure of oxygen is low, hemoglobin readily binds to carbon dioxide.
[caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this video to see the transport of oxygen from the lungs to the tissues.[/caption] [caption id="attachment_3013" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about the transport of oxygen in the body![/caption]
]]> 2630 2618 2 0 24.1 Overview of Metabolic Reactions https://pressbooks.bccampus.ca/apdouglas2017/chapter/24-1-overview-of-metabolic-reactions/ Mon, 17 Jul 2017 19:29:13 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2637

Learning Objectives

By the end of this section, you will be able to:
  • Describe the process by which polymers are broken down into monomers
  • Describe the process by which monomers are combined into polymers
  • Discuss the role of ATP in metabolism
  • Explain oxidation-reduction reactions
  • Describe the hormones that regulate anabolic and catabolic reactions

Metabolic processes are constantly taking place in the body. Metabolism is the sum of all of the chemical reactions that are involved in catabolism and anabolism. The reactions governing the breakdown of food to obtain energy are called catabolic reactions. Conversely, anabolic reactions use the energy produced by catabolic reactions to synthesize larger molecules from smaller ones, such as when the body forms proteins by stringing together amino acids. Both sets of reactions are critical to maintaining life.

Because catabolic reactions produce energy and anabolic reactions use energy, ideally, energy usage would balance the energy produced. If the net energy change is positive (catabolic reactions release more energy than the anabolic reactions use), then the body stores the excess energy by building fat molecules for long-term storage. On the other hand, if the net energy change is negative (catabolic reactions release less energy than anabolic reactions use), the body uses stored energy to compensate for the deficiency of energy released by catabolism.

Catabolic Reactions

Catabolic reactions break down large organic molecules into smaller molecules, releasing the energy contained in the chemical bonds. These energy releases (conversions) are not 100 percent efficient. The amount of energy released is less than the total amount contained in the molecule. Approximately 40 percent of energy yielded from catabolic reactions is directly transferred to the high-energy molecule adenosine triphosphate (ATP). ATP, the energy currency of cells, can be used immediately to power molecular machines that support cell, tissue, and organ function. This includes building new tissue and repairing damaged tissue. ATP can also be stored to fulfill future energy demands. The remaining 60 percent of the energy released from catabolic reactions is given off as heat, which tissues and body fluids absorb.

Structurally, ATP molecules consist of an adenine, a ribose, and three phosphate groups (Figure 1). The chemical bond between the second and third phosphate groups, termed a high-energy bond, represents the greatest source of energy in a cell. It is the first bond that catabolic enzymes break when cells require energy to do work. The products of this reaction are a molecule of adenosine diphosphate (ADP) and a lone phosphate group (Pi). ATP, ADP, and Pi are constantly being cycled through reactions that build ATP and store energy, and reactions that break down ATP and release energy.

[caption id="" align="aligncenter" width="550"]This diagram shows the chemical structure of adenosine triphosphate, and how different reactions add or remove phosphate groups. Figure 1. Structure of ATP Molecule. Adenosine triphosphate (ATP) is the energy molecule of the cell. During catabolic reactions, ATP is created and energy is stored until needed during anabolic reactions.[/caption]

The energy from ATP drives all bodily functions, such as contracting muscles, maintaining the electrical potential of nerve cells, and absorbing food in the gastrointestinal tract. The metabolic reactions that produce ATP come from various sources (Figure 2).

[caption id="" align="aligncenter" width="520"]This flowchart shows how food is modified into lipids, carbohydrates, and protein, and the various catabolic reactions which convert food into energy. Figure 2. Sources of ATP. During catabolic reactions, proteins are broken down into amino acids, lipids are broken down into fatty acids, and polysaccharides are broken down into monosaccharides. These building blocks are then used for the synthesis of molecules in anabolic reactions.[/caption]

Of the four major macromolecular groups (carbohydrates, lipids, proteins, and nucleic acids) that are processed by digestion, carbohydrates are considered the most common source of energy to fuel the body. They take the form of either complex carbohydrates, polysaccharides like starch and glycogen, or simple sugars (monosaccharides) like glucose and fructose. Sugar catabolism breaks polysaccharides down into their individual monosaccharides. Among the monosaccharides, glucose is the most common fuel for ATP production in cells, and as such, there are a number of endocrine control mechanisms to regulate glucose concentration in the bloodstream. Excess glucose is either stored as an energy reserve in the liver and skeletal muscles as the complex polymer glycogen, or it is converted into fat (triglyceride) in adipose cells (adipocytes).

Among the lipids (fats), triglycerides are most often used for energy via a metabolic process called β-oxidation. About one-half of excess fat is stored in adipocytes that accumulate in the subcutaneous tissue under the skin, whereas the rest is stored in adipocytes in other tissues and organs.

Proteins, which are polymers, can be broken down into their monomers, individual amino acids. Amino acids can be used as building blocks of new proteins or broken down further for the production of ATP. When one is chronically starving, this use of amino acids for energy production can lead to a wasting away of the body, as more and more proteins are broken down. Nucleic acids are present in most of the foods you eat. During digestion, nucleic acids including DNA and various RNAs are broken down into their constituent nucleotides. These nucleotides are readily absorbed and transported throughout the body to be used by individual cells during nucleic acid metabolism.

Anabolic Reactions

In contrast to catabolic reactions, anabolic reactions involve the joining of smaller molecules into larger ones. Anabolic reactions combine monosaccharides to form polysaccharides, fatty acids to form triglycerides, amino acids to form proteins, and nucleotides to form nucleic acids. These processes require energy in the form of ATP molecules generated by catabolic reactions. Anabolic reactions, also called biosynthesis reactions, create new molecules that form new cells and tissues, and revitalize organs.

Hormonal Regulation of Metabolism

Catabolic and anabolic hormones in the body help regulate metabolic processes. Catabolic hormones stimulate the breakdown of molecules and the production of energy. These include cortisol, glucagon, adrenaline/epinephrine, and cytokines. All of these hormones are mobilized at specific times to meet the needs of the body. Anabolic hormones are required for the synthesis of molecules and include growth hormone, insulin-like growth factor, insulin, testosterone, and estrogen. Table 1 summarizes the function of each of the catabolic hormones and Table 2 summarizes the functions of the anabolic hormones.
Catabolic Hormones (Table 1)
Hormone Function
Cortisol Released from the adrenal gland in response to stress; its main role is to increase blood glucose levels by gluconeogenesis (breaking down fats and proteins)
Glucagon Released from alpha cells in the pancreas either when starving or when the body needs to generate additional energy; it stimulates the breakdown of glycogen in the liver to increase blood glucose levels; its effect is the opposite of insulin; glucagon and insulin are a part of a negative-feedback system that stabilizes blood glucose levels
Adrenaline/epinephrine Released in response to the activation of the sympathetic nervous system; increases heart rate and heart contractility, constricts blood vessels, is a bronchodilator that opens (dilates) the bronchi of the lungs to increase air volume in the lungs, and stimulates gluconeogenesis
Anabolic Hormones (Table 2)
Hormone Function
Growth hormone (GH) Synthesized and released from the pituitary gland; stimulates the growth of cells, tissues, and bones
Insulin-like growth factor (IGF) Stimulates the growth of muscle and bone while also inhibiting cell death (apoptosis)
Insulin Produced by the beta cells of the pancreas; plays an essential role in carbohydrate and fat metabolism, controls blood glucose levels, and promotes the uptake of glucose into body cells; causes cells in muscle, adipose tissue, and liver to take up glucose from the blood and store it in the liver and muscle as glucagon; its effect is the opposite of glucagon; glucagon and insulin are a part of a negative-feedback system that stabilizes blood glucose levels
Testosterone Produced by the testes in males and the ovaries in females; stimulates an increase in muscle mass and strength as well as the growth and strengthening of bone
Estrogen Produced primarily by the ovaries, it is also produced by the liver and adrenal glands; its anabolic functions include increasing metabolism and fat deposition
Disorders of the…

Metabolic Processes: Cushing Syndrome and Addison’s Disease As might be expected for a fundamental physiological process like metabolism, errors or malfunctions in metabolic processing lead to a pathophysiology or—if uncorrected—a disease state. Metabolic diseases are most commonly the result of malfunctioning proteins or enzymes that are critical to one or more metabolic pathways. Protein or enzyme malfunction can be the consequence of a genetic alteration or mutation. However, normally functioning proteins and enzymes can also have deleterious effects if their availability is not appropriately matched with metabolic need. For example, excessive production of the hormone cortisol (see Table 1) gives rise to Cushing syndrome. Clinically, Cushing syndrome is characterized by rapid weight gain, especially in the trunk and face region, depression, and anxiety. It is worth mentioning that tumors of the pituitary that produce adrenocorticotropic hormone (ACTH), which subsequently stimulates the adrenal cortex to release excessive cortisol, produce similar effects. This indirect mechanism of cortisol overproduction is referred to as Cushing disease.

Patients with Cushing syndrome can exhibit high blood glucose levels and are at an increased risk of becoming obese. They also show slow growth, accumulation of fat between the shoulders, weak muscles, bone pain (because cortisol causes proteins to be broken down to make glucose via gluconeogenesis), and fatigue. Other symptoms include excessive sweating (hyperhidrosis), capillary dilation, and thinning of the skin, which can lead to easy bruising. The treatments for Cushing syndrome are all focused on reducing excessive cortisol levels. Depending on the cause of the excess, treatment may be as simple as discontinuing the use of cortisol ointments. In cases of tumors, surgery is often used to remove the offending tumor. Where surgery is inappropriate, radiation therapy can be used to reduce the size of a tumor or ablate portions of the adrenal cortex. Finally, medications are available that can help to regulate the amounts of cortisol.

Insufficient cortisol production is equally problematic. Adrenal insufficiency, or Addison’s disease, is characterized by the reduced production of cortisol from the adrenal gland. It can result from malfunction of the adrenal glands—they do not produce enough cortisol—or it can be a consequence of decreased ACTH availability from the pituitary. Patients with Addison’s disease may have low blood pressure, paleness, extreme weakness, fatigue, slow or sluggish movements, lightheadedness, and salt cravings due to the loss of sodium and high blood potassium levels (hyperkalemia). Victims also may suffer from loss of appetite, chronic diarrhea, vomiting, mouth lesions, and patchy skin color. Diagnosis typically involves blood tests and imaging tests of the adrenal and pituitary glands. Treatment involves cortisol replacement therapy, which usually must be continued for life.

Oxidation-Reduction Reactions

The chemical reactions underlying metabolism involve the transfer of electrons from one compound to another by processes catalyzed by enzymes. The electrons in these reactions commonly come from hydrogen atoms, which consist of an electron and a proton. A molecule gives up a hydrogen atom, in the form of a hydrogen ion (H+) and an electron, breaking the molecule into smaller parts. The loss of an electron, or oxidation, releases a small amount of energy; both the electron and the energy are then passed to another molecule in the process of reduction, or the gaining of an electron. These two reactions always happen together in an oxidation-reduction reaction (also called a redox reaction)—when an electron is passed between molecules, the donor is oxidized and the recipient is reduced. Oxidation-reduction reactions often happen in a series, so that a molecule that is reduced is subsequently oxidized, passing on not only the electron it just received but also the energy it received. As the series of reactions progresses, energy accumulates that is used to combine Pi and ADP to form ATP, the high-energy molecule that the body uses for fuel.

Oxidation-reduction reactions are catalyzed by enzymes that trigger the removal of hydrogen atoms. Coenzymes work with enzymes and accept hydrogen atoms. The two most common coenzymes of oxidation-reduction reactions are nicotinamide adenine dinucleotide (NAD) and flavin adenine dinucleotide (FAD). Their respective reduced coenzymes are NADH and FADH2, which are energy-containing molecules used to transfer energy during the creation of ATP.

[caption id="attachment_3021" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of metabolism and nutrition.[/caption]]]> 2637 2634 1 0 23.1 Overview of the Digestive System https://pressbooks.bccampus.ca/apdouglas2017/chapter/23-1-overview-of-the-digestive-system/ Mon, 17 Jul 2017 19:18:34 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2670

Learning Objectives

By the end of this section, you will be able to:
  • Identify the organs of the alimentary canal from proximal to distal, and briefly state their function
  • Identify the accessory digestive organs and briefly state their function
  • Describe the four fundamental tissue layers of the alimentary canal
  • Contrast the contributions of the enteric and autonomic nervous systems to digestive system functioning
  • Explain how the peritoneum anchors the digestive organs

The function of the digestive system is to break down the foods you eat, release their nutrients, and absorb those nutrients into the body. Although the small intestine is the workhorse of the system, where the majority of digestion occurs, and where most of the released nutrients are absorbed into the blood or lymph, each of the digestive system organs makes a vital contribution to this process (Figure 1).

[caption id="" align="aligncenter" width="400"]This diagram shows the digestive system of a human being, with the major organs labeled. Figure 1. Components of the Digestive System. All digestive organs play integral roles in the life-sustaining process of digestion.[/caption]

As is the case with all body systems, the digestive system does not work in isolation; it functions cooperatively with the other systems of the body. Consider for example, the interrelationship between the digestive and cardiovascular systems. Arteries supply the digestive organs with oxygen and processed nutrients, and veins drain the digestive tract. These intestinal veins, constituting the hepatic portal system, are unique; they do not return blood directly to the heart. Rather, this blood is diverted to the liver where its nutrients are off-loaded for processing before blood completes its circuit back to the heart. At the same time, the digestive system provides nutrients to the heart muscle and vascular tissue to support their functioning. The interrelationship of the digestive and endocrine systems is also critical. Hormones secreted by several endocrine glands, as well as endocrine cells of the pancreas, the stomach, and the small intestine, contribute to the control of digestion and nutrient metabolism. In turn, the digestive system provides the nutrients to fuel endocrine function. Table 1 gives a quick glimpse at how these other systems contribute to the functioning of the digestive system.

Contribution of Other Body Systems to the Digestive System (Table 1)
Body system Benefits received by the digestive system
Cardiovascular Blood supplies digestive organs with oxygen and processed nutrients
Endocrine Endocrine hormones help regulate secretion in digestive glands and accessory organs
Integumentary Skin helps protect digestive organs and synthesizes vitamin D for calcium absorption
Lymphatic Mucosa-associated lymphoid tissue and other lymphatic tissue defend against entry of pathogens; lacteals absorb lipids; and lymphatic vessels transport lipids to bloodstream
Muscular Skeletal muscles support and protect abdominal organs
Nervous Sensory and motor neurons help regulate secretions and muscle contractions in the digestive tract
Respiratory Respiratory organs provide oxygen and remove carbon dioxide
Skeletal Bones help protect and support digestive organs
Urinary Kidneys convert vitamin D into its active form, allowing calcium absorption in the small intestine

Digestive System Organs

The easiest way to understand the digestive system is to divide its organs into two main categories. The first group is the organs that make up the alimentary canal. Accessory digestive organs comprise the second group and are critical for orchestrating the breakdown of food and the assimilation of its nutrients into the body. Accessory digestive organs, despite their name, are critical to the function of the digestive system.

Alimentary Canal Organs

Also called the gastrointestinal (GI) tract or gut, the alimentary canal (aliment- = “to nourish”) is a one-way tube about 7.62 meters (25 feet) in length during life and closer to 10.67 meters (35 feet) in length when measured after death, once smooth muscle tone is lost. The main function of the organs of the alimentary canal is to nourish the body. This tube begins at the mouth and terminates at the anus. Between those two points, the canal is modified as the pharynx, esophagus, stomach, and small and large intestines to fit the functional needs of the body. Both the mouth and anus are open to the external environment; thus, food and wastes within the alimentary canal are technically considered to be outside the body. Only through the process of absorption do the nutrients in food enter into and nourish the body’s “inner space.”

Accessory Structures

Each accessory digestive organ aids in the breakdown of food (Figure 2). Within the mouth, the teeth and tongue begin mechanical digestion, whereas the salivary glands begin chemical digestion. Once food products enter the small intestine, the gallbladder, liver, and pancreas release secretions—such as bile and enzymes—essential for digestion to continue. Together, these are called accessory organs because they sprout from the lining cells of the developing gut (mucosa) and augment its function; indeed, you could not live without their vital contributions, and many significant diseases result from their malfunction. Even after development is complete, they maintain a connection to the gut by way of ducts.

Histology of the Alimentary Canal

Throughout its length, the alimentary tract is composed of the same four tissue layers; the details of their structural arrangements vary to fit their specific functions. Starting from the lumen and moving outwards, these layers are the mucosa, submucosa, muscularis, and serosa, which is continuous with the mesentery (see Figure 2).

[caption id="" align="aligncenter" width="480"]This image shows the cross section of the alimentary canal. The different layers of the alimentary canal are shown as concentric cylinders with major muscles and veins labeled. Figure 2. Layers of the Alimentary Canal. The wall of the alimentary canal has four basic tissue layers: the mucosa, submucosa, muscularis, and serosa.[/caption]

The mucosa is referred to as a mucous membrane, because mucus production is a characteristic feature of gut epithelium. The membrane consists of epithelium, which is in direct contact with ingested food, and the lamina propria, a layer of connective tissue analogous to the dermis. In addition, the mucosa has a thin, smooth muscle layer, called the muscularis mucosa (not to be confused with the muscularis layer, described below).

Epithelium—In the mouth, pharynx, esophagus, and anal canal, the epithelium is primarily a non-keratinized, stratified squamous epithelium. In the stomach and intestines, it is a simple columnar epithelium. Notice that the epithelium is in direct contact with the lumen, the space inside the alimentary canal. Interspersed among its epithelial cells are goblet cells, which secrete mucus and fluid into the lumen, and enteroendocrine cells, which secrete hormones into the interstitial spaces between cells. Epithelial cells have a very brief lifespan, averaging from only a couple of days (in the mouth) to about a week (in the gut). This process of rapid renewal helps preserve the health of the alimentary canal, despite the wear and tear resulting from continued contact with foodstuffs.

Lamina propria—In addition to loose connective tissue, the lamina propria contains numerous blood and lymphatic vessels that transport nutrients absorbed through the alimentary canal to other parts of the body. The lamina propria also serves an immune function by housing clusters of lymphocytes, making up the mucosa-associated lymphoid tissue (MALT). These lymphocyte clusters are particularly substantial in the distal ileum where they are known as Peyer’s patches. When you consider that the alimentary canal is exposed to foodborne bacteria and other foreign matter, it is not hard to appreciate why the immune system has evolved a means of defending against the pathogens encountered within it.

Muscularis mucosa—This thin layer of smooth muscle is in a constant state of tension, pulling the mucosa of the stomach and small intestine into undulating folds. These folds dramatically increase the surface area available for digestion and absorption.

As its name implies, the submucosa lies immediately beneath the mucosa. A broad layer of dense connective tissue, it connects the overlying mucosa to the underlying muscularis. It includes blood and lymphatic vessels (which transport absorbed nutrients), and a scattering of submucosal glands that release digestive secretions. Additionally, it serves as a conduit for a dense branching network of nerves, the submucosal plexus, which functions as described below.

The third layer of the alimentary canal is the muscalaris (also called the muscularis externa). The muscularis in the small intestine is made up of a double layer of smooth muscle: an inner circular layer and an outer longitudinal layer. The contractions of these layers promote mechanical digestion, expose more of the food to digestive chemicals, and move the food along the canal. In the most proximal and distal regions of the alimentary canal, including the mouth, pharynx, anterior part of the esophagus, and external anal sphincter, the muscularis is made up of skeletal muscle, which gives you voluntary control over swallowing and defecation. The basic two-layer structure found in the small intestine is modified in the organs proximal and distal to it. The stomach is equipped for its churning function by the addition of a third layer, the oblique muscle. While the colon has two layers like the small intestine, its longitudinal layer is segregated into three narrow parallel bands, the tenia coli, which make it look like a series of pouches rather than a simple tube.

The serosa is the portion of the alimentary canal superficial to the muscularis. Present only in the region of the alimentary canal within the abdominal cavity, it consists of a layer of visceral peritoneum overlying a layer of loose connective tissue. Instead of serosa, the mouth, pharynx, and esophagus have a dense sheath of collagen fibers called the adventitia. These tissues serve to hold the alimentary canal in place near the ventral surface of the vertebral column.

Nerve Supply

As soon as food enters the mouth, it is detected by receptors that send impulses along the sensory neurons of cranial nerves. Without these nerves, not only would your food be without taste, but you would also be unable to feel either the food or the structures of your mouth, and you would be unable to avoid biting yourself as you chew, an action enabled by the motor branches of cranial nerves.

Intrinsic innervation of much of the alimentary canal is provided by the enteric nervous system, which runs from the esophagus to the anus, and contains approximately 100 million motor, sensory, and interneurons (unique to this system compared to all other parts of the peripheral nervous system). These enteric neurons are grouped into two plexuses. The myenteric plexus (plexus of Auerbach) lies in the muscularis layer of the alimentary canal and is responsible for motility, especially the rhythm and force of the contractions of the muscularis. The submucosal plexus (plexus of Meissner) lies in the submucosal layer and is responsible for regulating digestive secretions and reacting to the presence of food (see [link]).

Extrinsic innervations of the alimentary canal are provided by the autonomic nervous system, which includes both sympathetic and parasympathetic nerves. In general, sympathetic activation (the fight-or-flight response) restricts the activity of enteric neurons, thereby decreasing GI secretion and motility. In contrast, parasympathetic activation (the rest-and-digest response) increases GI secretion and motility by stimulating neurons of the enteric nervous system.

Blood Supply

The blood vessels serving the digestive system have two functions. They transport the protein and carbohydrate nutrients absorbed by mucosal cells after food is digested in the lumen. Lipids are absorbed via lacteals, tiny structures of the lymphatic system. The blood vessels’ second function is to supply the organs of the alimentary canal with the nutrients and oxygen needed to drive their cellular processes.

Specifically, the more anterior parts of the alimentary canal are supplied with blood by arteries branching off the aortic arch and thoracic aorta. Below this point, the alimentary canal is supplied with blood by arteries branching from the abdominal aorta. The celiac trunk services the liver, stomach, and duodenum, whereas the superior and inferior mesenteric arteries supply blood to the remaining small and large intestines.

The veins that collect nutrient-rich blood from the small intestine (where most absorption occurs) empty into the hepatic portal system. This venous network takes the blood into the liver where the nutrients are either processed or stored for later use. Only then does the blood drained from the alimentary canal viscera circulate back to the heart. To appreciate just how demanding the digestive process is on the cardiovascular system, consider that while you are “resting and digesting,” about one-fourth of the blood pumped with each heartbeat enters arteries serving the intestines.

The Peritoneum

The digestive organs within the abdominal cavity are held in place by the peritoneum, a broad serous membranous sac made up of squamous epithelial tissue surrounded by connective tissue. It is composed of two different regions: the parietal peritoneum, which lines the abdominal wall, and the visceral peritoneum, which envelopes the abdominal organs (Figure 3). The peritoneal cavity is the space bounded by the visceral and parietal peritoneal surfaces. A few milliliters of watery fluid act as a lubricant to minimize friction between the serosal surfaces of the peritoneum.
[caption id="" align="aligncenter" width="420"]This diagram shows the cross section of the abdomen. The peritoneum is made distinguishable from the abdominal organs through darker lines. Figure 3. The Peritoneum. A cross-section of the abdomen shows the relationship between abdominal organs and the peritoneum (darker lines).[/caption]
[caption id="attachment_3015" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of the digestive system![/caption]
]]> 2670 2664 2 0 23.2 Digestive System Processes and Regulation https://pressbooks.bccampus.ca/apdouglas2017/chapter/23-2-digestive-system-processes-and-regulation/ Mon, 17 Jul 2017 19:24:57 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2674

Learning Objectives

By the end of this section, you will be able to:
  • Discuss six fundamental activities of the digestive system, giving an example of each
  • Compare and contrast the neural and hormonal controls involved in digestion

The digestive system uses mechanical and chemical activities to break food down into absorbable substances during its journey through the digestive system. Table 3 provides an overview of the basic functions of the digestive organs.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit this site for an overview of digestion of food in different regions of the digestive tract.[/caption]
Functions of the Digestive Organs (Table 3)
Organ Major functions Other functions
Mouth
  • Ingests food
  • Chews and mixes food
  • Begins chemical breakdown of carbohydrates
  • Moves food into the pharynx
  • Begins breakdown of lipids via lingual lipase
  • Moistens and dissolves food, allowing you to taste it
  • Cleans and lubricates the teeth and oral cavity
  • Has some antimicrobial activity
Pharynx
  • Propels food from the oral cavity to the esophagus
  • Lubricates food and passageways
Esophagus
  • Propels food to the stomach
  • Lubricates food and passageways
Stomach
  • Mixes and churns food with gastric juices to form chyme
  • Begins chemical breakdown of proteins
  • Releases food into the duodenum as chyme
  • Absorbs some fat-soluble substances (for example, alcohol, aspirin)
  • Possesses antimicrobial functions
  • Stimulates protein-digesting enzymes
  • Secretes intrinsic factor required for vitamin B12 absorption in small intestine
Small intestine
  • Mixes chyme with digestive juices
  • Propels food at a rate slow enough for digestion and absorption
  • Absorbs breakdown products of carbohydrates, proteins, lipids, and nucleic acids, along with vitamins, minerals, and water
  • Performs physical digestion via segmentation
  • Provides optimal medium for enzymatic activity
Accessory organs
  • Liver: produces bile salts, which emulsify lipids, aiding their digestion and absorption
  • Gallbladder: stores, concentrates, and releases bile
  • Pancreas: produces digestive enzymes and bicarbonate
  • Bicarbonate-rich pancreatic juices help neutralize acidic chyme and provide optimal environment for enzymatic activity
Large intestine
  • Further breaks down food residues
  • Absorbs most residual water, electrolytes, and vitamins produced by enteric bacteria
  • Propels feces toward rectum
  • Eliminates feces
  • Food residue is concentrated and temporarily stored prior to defecation
  • Mucus eases passage of feces through colon

Digestive Processes

The processes of digestion include six activities: ingestion, propulsion, mechanical or physical digestion, chemical digestion, absorption, and defecation.

The first of these processes, ingestion, refers to the entry of food into the alimentary canal through the mouth. There, the food is chewed and mixed with saliva, which contains enzymes that begin breaking down the carbohydrates in the food plus some lipid digestion via lingual lipase. Chewing increases the surface area of the food and allows an appropriately sized bolus to be produced.

Food leaves the mouth when the tongue and pharyngeal muscles propel it into the esophagus. This act of swallowing, the last voluntary act until defecation, is an example of propulsion, which refers to the movement of food through the digestive tract. It includes both the voluntary process of swallowing and the involuntary process of peristalsis. Peristalsis consists of sequential, alternating waves of contraction and relaxation of alimentary wall smooth muscles, which act to propel food along (Figure 1). These waves also play a role in mixing food with digestive juices. Peristalsis is so powerful that foods and liquids you swallow enter your stomach even if you are standing on your head.

[caption id="" align="aligncenter" width="280"]This image shows the peristaltic movement of food. In the left image, the food bolus is towards the top of the esophagus and arrows pointing downward show the direction of movement of the peristaltic wave. In the center image, the food bolus and the wave movement are closer to the center of the esophagus and in the right image, the bolus and the wave are close to the bottom end of the esophagus. Figure 1. Peristalsis. Peristalsis moves food through the digestive tract with alternating waves of muscle contraction and relaxation.[/caption]

Digestion includes both mechanical and chemical processes. Mechanical digestion is a purely physical process that does not change the chemical nature of the food. Instead, it makes the food smaller to increase both surface area and mobility. It includes mastication, or chewing, as well as tongue movements that help break food into smaller bits and mix food with saliva. Although there may be a tendency to think that mechanical digestion is limited to the first steps of the digestive process, it occurs after the food leaves the mouth, as well. The mechanical churning of food in the stomach serves to further break it apart and expose more of its surface area to digestive juices, creating an acidic “soup” called chyme. Segmentation, which occurs mainly in the small intestine, consists of localized contractions of circular muscle of the muscularis layer of the alimentary canal. These contractions isolate small sections of the intestine, moving their contents back and forth while continuously subdividing, breaking up, and mixing the contents. By moving food back and forth in the intestinal lumen, segmentation mixes food with digestive juices and facilitates absorption.

In chemical digestion, starting in the mouth, digestive secretions break down complex food molecules into their chemical building blocks (for example, proteins into separate amino acids). These secretions vary in composition, but typically contain water, various enzymes, acids, and salts. The process is completed in the small intestine.

Food that has been broken down is of no value to the body unless it enters the bloodstream and its nutrients are put to work. This occurs through the process of absorption, which takes place primarily within the small intestine. There, most nutrients are absorbed from the lumen of the alimentary canal into the bloodstream through the epithelial cells that make up the mucosa. Lipids are absorbed into lacteals and are transported via the lymphatic vessels to the bloodstream (the subclavian veins near the heart). The details of these processes will be discussed later.

In defecation, the final step in digestion, undigested materials are removed from the body as feces.

Aging and the…
Digestive System: From Appetite Suppression to Constipation Age-related changes in the digestive system begin in the mouth and can affect virtually every aspect of the digestive system. Taste buds become less sensitive, so food isn’t as appetizing as it once was. A slice of pizza is a challenge, not a treat, when you have lost teeth, your gums are diseased, and your salivary glands aren’t producing enough saliva. Swallowing can be difficult, and ingested food moves slowly through the alimentary canal because of reduced strength and tone of muscular tissue. Neurosensory feedback is also dampened, slowing the transmission of messages that stimulate the release of enzymes and hormones.

Pathologies that affect the digestive organs—such as hiatal hernia, gastritis, and peptic ulcer disease—can occur at greater frequencies as you age. Problems in the small intestine may include duodenal ulcers, maldigestion, and malabsorption. Problems in the large intestine include hemorrhoids, diverticular disease, and constipation. Conditions that affect the function of accessory organs—and their abilities to deliver pancreatic enzymes and bile to the small intestine—include jaundice, acute pancreatitis, cirrhosis, and gallstones.

In some cases, a single organ is in charge of a digestive process. For example, ingestion occurs only in the mouth and defecation only in the anus. However, most digestive processes involve the interaction of several organs and occur gradually as food moves through the alimentary canal (Figure 2).

[caption id="" align="aligncenter" width="420"]This image shows the different processes involved in digestion. The image shows how food travels from the mouth through the major organs. Associated textboxes list the different processes such as propulsion, chemical and mechanical digestion and absorption near the organs where they take place. Figure 2. Digestive Processes. The digestive processes are ingestion, propulsion, mechanical digestion, chemical digestion, absorption, and defecation.[/caption]

Some chemical digestion occurs in the mouth. Some absorption can occur in the mouth and stomach, for example, alcohol and aspirin.

Regulatory Mechanisms

Neural and endocrine regulatory mechanisms work to maintain the optimal conditions in the lumen needed for digestion and absorption. These regulatory mechanisms, which stimulate digestive activity through mechanical and chemical activity, are controlled both extrinsically and intrinsically.

Neural Controls

The walls of the alimentary canal contain a variety of sensors that help regulate digestive functions. These include mechanoreceptors, chemoreceptors, and osmoreceptors, which are capable of detecting mechanical, chemical, and osmotic stimuli, respectively. For example, these receptors can sense when the presence of food has caused the stomach to expand, whether food particles have been sufficiently broken down, how much liquid is present, and the type of nutrients in the food (lipids, carbohydrates, and/or proteins). Stimulation of these receptors provokes an appropriate reflex that furthers the process of digestion. This may entail sending a message that activates the glands that secrete digestive juices into the lumen, or it may mean the stimulation of muscles within the alimentary canal, thereby activating peristalsis and segmentation that move food along the intestinal tract. The walls of the entire alimentary canal are embedded with nerve plexuses that interact with the central nervous system and other nerve plexuses—either within the same digestive organ or in different ones. These interactions prompt several types of reflexes. Extrinsic nerve plexuses orchestrate long reflexes, which involve the central and autonomic nervous systems and work in response to stimuli from outside the digestive system. Short reflexes, on the other hand, are orchestrated by intrinsic nerve plexuses within the alimentary canal wall. These two plexuses and their connections were introduced earlier as the enteric nervous system. Short reflexes regulate activities in one area of the digestive tract and may coordinate local peristaltic movements and stimulate digestive secretions. For example, the sight, smell, and taste of food initiate long reflexes that begin with a sensory neuron delivering a signal to the medulla oblongata. The response to the signal is to stimulate cells in the stomach to begin secreting digestive juices in preparation for incoming food. In contrast, food that distends the stomach initiates short reflexes that cause cells in the stomach wall to increase their secretion of digestive juices.

Hormonal Controls

A variety of hormones are involved in the digestive process. The main digestive hormone of the stomach is gastrin, which is secreted in response to the presence of food. Gastrin stimulates the secretion of gastric acid by the parietal cells of the stomach mucosa. Other GI hormones are produced and act upon the gut and its accessory organs. Hormones produced by the duodenum include secretin, which stimulates a watery secretion of bicarbonate by the pancreas; cholecystokinin (CCK), which stimulates the secretion of pancreatic enzymes and bile from the liver and release of bile from the gallbladder; and gastric inhibitory peptide, which inhibits gastric secretion and slows gastric emptying and motility. These GI hormones are secreted by specialized epithelial cells, called endocrinocytes, located in the mucosal epithelium of the stomach and small intestine. These hormones then enter the bloodstream, through which they can reach their target organs.

[caption id="attachment_3017" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about digestion![/caption]
]]> 2674 2664 3 0 23.5 The Small and Large Intestines https://pressbooks.bccampus.ca/apdouglas2017/chapter/23-5-the-small-and-large-intestines/ Mon, 17 Jul 2017 19:27:29 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2698

Learning Objectives

By the end of this section, you will be able to:
  • Compare and contrast the location and gross anatomy of the small and large intestines
  • Identify three main adaptations of the small intestine wall that increase its absorptive capacity
  • Describe the mechanical and chemical digestion of chyme upon its release into the small intestine
  • List three features unique to the wall of the large intestine and identify their contributions to its function
  • Identify the beneficial roles of the bacterial flora in digestive system functioning
  • Trace the pathway of food waste from its point of entry into the large intestine through its exit from the body as feces

The word intestine is derived from a Latin root meaning “internal,” and indeed, the two organs together nearly fill the interior of the abdominal cavity. In addition, called the small and large bowel, or colloquially the “guts,” they constitute the greatest mass and length of the alimentary canal and, with the exception of ingestion, perform all digestive system functions.

The Small Intestine

Chyme released from the stomach enters the small intestine, which is the primary digestive organ in the body. Not only is this where most digestion occurs, it is also where practically all absorption occurs. The longest part of the alimentary canal, the small intestine is about 3.05 meters (10 feet) long in a living person (but about twice as long in a cadaver due to the loss of muscle tone). Since this makes it about five times longer than the large intestine, you might wonder why it is called “small.” In fact, its name derives from its relatively smaller diameter of only about 2.54 cm (1 in), compared with 7.62 cm (3 in) for the large intestine. As we’ll see shortly, in addition to its length, the folds and projections of the lining of the small intestine work to give it an enormous surface area, which is approximately 200 m2, more than 100 times the surface area of your skin. This large surface area is necessary for complex processes of digestion and absorption that occur within it.

Structure

The coiled tube of the small intestine is subdivided into three regions. From proximal (at the stomach) to distal, these are the duodenum, jejunum, and ileum (Figure 1).

The shortest region is the 25.4-cm (10-in) duodenum, which begins at the pyloric sphincter. Just past the pyloric sphincter, it bends posteriorly behind the peritoneum, becoming retroperitoneal, and then makes a C-shaped curve around the head of the pancreas before ascending anteriorly again to return to the peritoneal cavity and join the jejunum. The duodenum can therefore be subdivided into four segments: the superior, descending, horizontal, and ascending duodenum.

Of particular interest is the hepatopancreatic ampulla (ampulla of Vater). Located in the duodenal wall, the ampulla marks the transition from the anterior portion of the alimentary canal to the mid-region, and is where the bile duct (through which bile passes from the liver) and the main pancreatic duct (through which pancreatic juice passes from the pancreas) join. This ampulla opens into the duodenum at a tiny volcano-shaped structure called the major duodenal papilla. The hepatopancreatic sphincter (sphincter of Oddi) regulates the flow of both bile and pancreatic juice from the ampulla into the duodenum.

[caption id="" align="aligncenter" width="420"]This diagram shows the small intestine. The different parts of the small intestine are labeled. Figure 1. Small Intestine. The three regions of the small intestine are the duodenum, jejunum, and ileum.[/caption]

The jejunum is about 0.9 meters (3 feet) long (in life) and runs from the duodenum to the ileum. Jejunum means “empty” in Latin and supposedly was so named by the ancient Greeks who noticed it was always empty at death. No clear demarcation exists between the jejunum and the final segment of the small intestine, the ileum.

The ileum is the longest part of the small intestine, measuring about 1.8 meters (6 feet) in length. It is thicker, more vascular, and has more developed mucosal folds than the jejunum. The ileum joins the cecum, the first portion of the large intestine, at the ileocecal sphincter (or valve). The jejunum and ileum are tethered to the posterior abdominal wall by the mesentery. The large intestine frames these three parts of the small intestine.

Parasympathetic nerve fibers from the vagus nerve and sympathetic nerve fibers from the thoracic splanchnic nerve provide extrinsic innervation to the small intestine. The superior mesenteric artery is its main arterial supply. Veins run parallel to the arteries and drain into the superior mesenteric vein. Nutrient-rich blood from the small intestine is then carried to the liver via the hepatic portal vein.

Histology

The wall of the small intestine is composed of the same four layers typically present in the alimentary system. However, three features of the mucosa and submucosa are unique. These features, which increase the absorptive surface area of the small intestine more than 600-fold, include circular folds, villi, and microvilli (Figure 2). These adaptations are most abundant in the proximal two-thirds of the small intestine, where the majority of absorption occurs.

[caption id="" align="aligncenter" width="550"]Illustration (a) shows the histological cross-section of the small intestine. The left panel shows a small region of the small intestine, along with the blood vessels and the muscle layers. The middle panel shows a magnified view of a small region of the small intestine, highlighting the absorptive cells, the lacteal and the goblet cells. The right panel shows a further magnified view of the epithelial cells including the microvilli. Illustrations (b) shows a micrograph of the circular folds, and illustration (c) shows a micrograph of the villi. Illustration (d) shows an electron micrograph of the microvilli. Figure 2. Histology of the Small Intestine. (a) The absorptive surface of the small intestine is vastly enlarged by the presence of circular folds, villi, and microvilli. (b) Micrograph of the circular folds. (c) Micrograph of the villi. (d) Electron micrograph of the microvilli. From left to right, LM x 56, LM x 508, EM x 196,000. (credit b-d: Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Circular folds

Also called a plica circulare, a circular fold is a deep ridge in the mucosa and submucosa. Beginning near the proximal part of the duodenum and ending near the middle of the ileum, these folds facilitate absorption. Their shape causes the chyme to spiral, rather than move in a straight line, through the small intestine. Spiraling slows the movement of chyme and provides the time needed for nutrients to be fully absorbed.

Villi

Within the circular folds are small (0.5–1 mm long) hairlike vascularized projections called villi (singular = villus) that give the mucosa a furry texture. There are about 20 to 40 villi per square millimeter, increasing the surface area of the epithelium tremendously. The mucosal epithelium, primarily composed of absorptive cells, covers the villi. In addition to muscle and connective tissue to support its structure, each villus contains a capillary bed composed of one arteriole and one venule, as well as a lymphatic capillary called a lacteal. The breakdown products of carbohydrates and proteins (sugars and amino acids) can enter the bloodstream directly, but lipid breakdown products are absorbed by the lacteals and transported to the bloodstream via the lymphatic system.

Microvilli

As their name suggests, microvilli (singular = microvillus) are much smaller (1 µm) than villi. They are cylindrical apical surface extensions of the plasma membrane of the mucosa’s epithelial cells, and are supported by microfilaments within those cells. Although their small size makes it difficult to see each microvillus, their combined microscopic appearance suggests a mass of bristles, which is termed the brush border. Fixed to the surface of the microvilli membranes are enzymes that finish digesting carbohydrates and proteins. There are an estimated 200 million microvilli per square millimeter of small intestine, greatly expanding the surface area of the plasma membrane and thus greatly enhancing absorption.

Intestinal Glands

In addition to the three specialized absorptive features just discussed, the mucosa between the villi is dotted with deep crevices that each lead into a tubular intestinal gland (crypt of Lieberkühn), which is formed by cells that line the crevices (see Figure 2). These produce intestinal juice, a slightly alkaline (pH 7.4 to 7.8) mixture of water and mucus. Each day, about 0.95 to 1.9 liters (1 to 2 quarts) are secreted in response to the distention of the small intestine or the irritating effects of chyme on the intestinal mucosa.

The submucosa of the duodenum is the only site of the complex mucus-secreting duodenal glands (Brunner’s glands), which produce a bicarbonate-rich alkaline mucus that buffers the acidic chyme as it enters from the stomach.

The roles of the cells in the small intestinal mucosa are detailed in Table 7.

Cells of the Small Intestinal Mucosa (Table 7)
Cell type Location in the mucosa Function
Absorptive Epithelium/intestinal glands Digestion and absorption of nutrients in chyme
Goblet Epithelium/intestinal glands Secretion of mucus
Paneth Intestinal glands Secretion of the bactericidal enzyme lysozyme; phagocytosis
G cells Intestinal glands of duodenum Secretion of the hormone intestinal gastrin
I cells Intestinal glands of duodenum Secretion of the hormone cholecystokinin, which stimulates release of pancreatic juices and bile
K cells Intestinal glands Secretion of the hormone glucose-dependent insulinotropic peptide, which stimulates the release of insulin
M cells Intestinal glands of duodenum and jejunum Secretion of the hormone motilin, which accelerates gastric emptying, stimulates intestinal peristalsis, and stimulates the production of pepsin
S cells Intestinal glands Secretion of the hormone secretin

Intestinal MALT

The lamina propria of the small intestine mucosa is studded with quite a bit of MALT. In addition to solitary lymphatic nodules, aggregations of intestinal MALT, which are typically referred to as Peyer’s patches, are concentrated in the distal ileum, and serve to keep bacteria from entering the bloodstream. Peyer’s patches are most prominent in young people and become less distinct as you age, which coincides with the general activity of our immune system.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this animation that depicts the structure of the small intestine, and, in particular, the villi.[/caption]

Mechanical Digestion in the Small Intestine

The movement of intestinal smooth muscles includes both segmentation and a form of peristalsis called migrating motility complexes. The kind of peristaltic mixing waves seen in the stomach are not observed here.

If you could see into the small intestine when it was going through segmentation, it would look as if the contents were being shoved incrementally back and forth, as the rings of smooth muscle repeatedly contract and then relax. Segmentation in the small intestine does not force chyme through the tract. Instead, it combines the chyme with digestive juices and pushes food particles against the mucosa to be absorbed. The duodenum is where the most rapid segmentation occurs, at a rate of about 12 times per minute. In the ileum, segmentations are only about eight times per minute (Figure 3).

[caption id="" align="aligncenter" width="250"]This diagram shows the process of segmentation in the intestines. The left panel shows the separation of chime, the middle panel shows the remixing of the chime by pushing it back together and the right panel indicates that the chime is being digested and absorbed. Figure 3. Segmentation. Segmentation separates chyme and then pushes it back together, mixing it and providing time for digestion and absorption.[/caption]

When most of the chyme has been absorbed, the small intestinal wall becomes less distended. At this point, the localized segmentation process is replaced by transport movements. The duodenal mucosa secretes the hormone motilin, which initiates peristalsis in the form of a migrating motility complex. These complexes, which begin in the duodenum, force chyme through a short section of the small intestine and then stop. The next contraction begins a little bit farther down than the first, forces chyme a bit farther through the small intestine, then stops. These complexes move slowly down the small intestine, forcing chyme on the way, taking around 90 to 120 minutes to finally reach the end of the ileum. At this point, the process is repeated, starting in the duodenum.

The ileocecal valve, a sphincter, is usually in a constricted state, but when motility in the ileum increases, this sphincter relaxes, allowing food residue to enter the first portion of the large intestine, the cecum. Relaxation of the ileocecal sphincter is controlled by both nerves and hormones. First, digestive activity in the stomach provokes the gastroileal reflex, which increases the force of ileal segmentation. Second, the stomach releases the hormone gastrin, which enhances ileal motility, thus relaxing the ileocecal sphincter. After chyme passes through, backward pressure helps close the sphincter, preventing backflow into the ileum. Because of this reflex, your lunch is completely emptied from your stomach and small intestine by the time you eat your dinner. It takes about 3 to 5 hours for all chyme to leave the small intestine.

Chemical Digestion in the Small Intestine

The digestion of proteins and carbohydrates, which partially occurs in the stomach, is completed in the small intestine with the aid of intestinal and pancreatic juices. Lipids arrive in the intestine largely undigested, so much of the focus here is on lipid digestion, which is facilitated by bile and the enzyme pancreatic lipase.

Moreover, intestinal juice combines with pancreatic juice to provide a liquid medium that facilitates absorption. The intestine is also where most water is absorbed, via osmosis. The small intestine’s absorptive cells also synthesize digestive enzymes and then place them in the plasma membranes of the microvilli. This distinguishes the small intestine from the stomach; that is, enzymatic digestion occurs not only in the lumen, but also on the luminal surfaces of the mucosal cells.

For optimal chemical digestion, chyme must be delivered from the stomach slowly and in small amounts. This is because chyme from the stomach is typically hypertonic, and if large quantities were forced all at once into the small intestine, the resulting osmotic water loss from the blood into the intestinal lumen would result in potentially life-threatening low blood volume. In addition, continued digestion requires an upward adjustment of the low pH of stomach chyme, along with rigorous mixing of the chyme with bile and pancreatic juices. Both processes take time, so the pumping action of the pylorus must be carefully controlled to prevent the duodenum from being overwhelmed with chyme.

The Large Intestine

The large intestine is the terminal part of the alimentary canal. The primary function of this organ is to finish absorption of nutrients and water, synthesize certain vitamins, form feces, and eliminate feces from the body.

Structure

The large intestine runs from the appendix to the anus. It frames the small intestine on three sides. Despite its being about one-half as long as the small intestine, it is called large because it is more than twice the diameter of the small intestine, about 3 inches.

Subdivisions

The large intestine is subdivided into four main regions: the cecum, the colon, the rectum, and the anus. The ileocecal valve, located at the opening between the ileum and the large intestine, controls the flow of chyme from the small intestine to the large intestine.

Cecum

The first part of the large intestine is the cecum, a sac-like structure that is suspended inferior to the ileocecal valve. It is about 6 cm (2.4 in) long, receives the contents of the ileum, and continues the absorption of water and salts. The appendix (or vermiform appendix) is a winding tube that attaches to the cecum. Although the 7.6-cm (3-in) long appendix contains lymphoid tissue, suggesting an immunologic function, this organ is generally considered vestigial. However, at least one recent report postulates a survival advantage conferred by the appendix: In diarrheal illness, the appendix may serve as a bacterial reservoir to repopulate the enteric bacteria for those surviving the initial phases of the illness. Moreover, its twisted anatomy provides a haven for the accumulation and multiplication of enteric bacteria. The mesoappendix, the mesentery of the appendix, tethers it to the mesentery of the ileum.

Colon

The cecum blends seamlessly with the colon. Upon entering the colon, the food residue first travels up the ascending colon on the right side of the abdomen. At the inferior surface of the liver, the colon bends to form the right colic flexure (hepatic flexure) and becomes the transverse colon. The region defined as hindgut begins with the last third of the transverse colon and continues on. Food residue passing through the transverse colon travels across to the left side of the abdomen, where the colon angles sharply immediately inferior to the spleen, at the left colic flexure (splenic flexure). From there, food residue passes through the descending colon, which runs down the left side of the posterior abdominal wall. After entering the pelvis inferiorly, it becomes the s-shaped sigmoid colon, which extends medially to the midline (Figure 4). The ascending and descending colon, and the rectum (discussed next) are located in the retroperitoneum. The transverse and sigmoid colon are tethered to the posterior abdominal wall by the mesocolon.

[caption id="" align="aligncenter" width="350"]This image shows the large intestine; the major parts of the large intestine are labeled. Figure 4. Large Intestine. The large intestine includes the cecum, colon, and rectum.[/caption]

Rectum

Food residue leaving the sigmoid colon enters the rectum in the pelvis, near the third sacral vertebra. The final 20.3 cm (8 in) of the alimentary canal, the rectum extends anterior to the sacrum and coccyx. Even though rectum is Latin for “straight,” this structure follows the curved contour of the sacrum and has three lateral bends that create a trio of internal transverse folds called the rectal valves. These valves help separate the feces from gas to prevent the simultaneous passage of feces and gas.

Anal Canal

Finally, food residue reaches the last part of the large intestine, the anal canal, which is located in the perineum, completely outside of the abdominopelvic cavity. This 3.8–5 cm (1.5–2 in) long structure opens to the exterior of the body at the anus. The anal canal includes two sphincters. The internal anal sphincter is made of smooth muscle, and its contractions are involuntary. The external anal sphincter is made of skeletal muscle, which is under voluntary control. Except when defecating, both usually remain closed.

Histology

There are several notable differences between the walls of the large and small intestines (Figure 5). For example, few enzyme-secreting cells are found in the wall of the large intestine, and there are no circular folds or villi. Other than in the anal canal, the mucosa of the colon is simple columnar epithelium made mostly of enterocytes (absorptive cells) and goblet cells. In addition, the wall of the large intestine has far more intestinal glands, which contain a vast population of enterocytes and goblet cells. These goblet cells secrete mucus that eases the movement of feces and protects the intestine from the effects of the acids and gases produced by enteric bacteria. The enterocytes absorb water and salts as well as vitamins produced by your intestinal bacteria.

[caption id="" align="aligncenter" width="480"]This image shows the histological cross section of the large intestine. The left panel shows a small region of the large intestine. The center panel shows a magnified view of this region, highlighting the openings of the intestinal glands. The right panel shows a further magnified view, with the microvilli and goblet cells. Figure 5. Histology of the large Intestine. (a) The histologies of the large intestine and small intestine (not shown) are adapted for the digestive functions of each organ. (b) This micrograph shows the colon’s simple columnar epithelium and goblet cells. LM x 464. (credit b: Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Anatomy

Three features are unique to the large intestine: teniae coli, haustra, and epiploic appendages (Figure 6). The teniae coli are three bands of smooth muscle that make up the longitudinal muscle layer of the muscularis of the large intestine, except at its terminal end. Tonic contractions of the teniae coli bunch up the colon into a succession of pouches called haustra (singular = hostrum), which are responsible for the wrinkled appearance of the colon. Attached to the teniae coli are small, fat-filled sacs of visceral peritoneum called epiploic appendages. The purpose of these is unknown. Although the rectum and anal canal have neither teniae coli nor haustra, they do have well-developed layers of muscularis that create the strong contractions needed for defecation.

[caption id="" align="aligncenter" width="320"]This image shows the Taenia Coli, haustra and epiploic appendages, which are parts of the large intestine. Figure 6. Teniae Coli, Haustra, and Epiploic Appendages[/caption]
The stratified squamous epithelial mucosa of the anal canal connects to the skin on the outside of the anus. This mucosa varies considerably from that of the rest of the colon to accommodate the high level of abrasion as feces pass through. The anal canal’s mucous membrane is organized into longitudinal folds, each called an anal column, which house a grid of arteries and veins. Two superficial venous plexuses are found in the anal canal: one within the anal columns and one at the anus.

Depressions between the anal columns, each called an anal sinus, secrete mucus that facilitates defecation. The pectinate line (or dentate line) is a horizontal, jagged band that runs circumferentially just below the level of the anal sinuses, and represents the junction between the hindgut and external skin. The mucosa above this line is fairly insensitive, whereas the area below is very sensitive. The resulting difference in pain threshold is due to the fact that the upper region is innervated by visceral sensory fibers, and the lower region is innervated by somatic sensory fibers.

Bacterial Flora

Most bacteria that enter the alimentary canal are killed by lysozyme, defensins, HCl, or protein-digesting enzymes. However, trillions of bacteria live within the large intestine and are referred to as the bacterial flora. Most of the more than 700 species of these bacteria are nonpathogenic commensal organisms that cause no harm as long as they stay in the gut lumen. In fact, many facilitate chemical digestion and absorption, and some synthesize certain vitamins, mainly biotin, pantothenic acid, and vitamin K. Some are linked to increased immune response. A refined system prevents these bacteria from crossing the mucosal barrier. First, peptidoglycan, a component of bacterial cell walls, activates the release of chemicals by the mucosa’s epithelial cells, which draft immune cells, especially dendritic cells, into the mucosa. Dendritic cells open the tight junctions between epithelial cells and extend probes into the lumen to evaluate the microbial antigens. The dendritic cells with antigens then travel to neighboring lymphoid follicles in the mucosa where T cells inspect for antigens. This process triggers an IgA-mediated response, if warranted, in the lumen that blocks the commensal organisms from infiltrating the mucosa and setting off a far greater, widespread systematic reaction.

Digestive Functions of the Large Intestine

The residue of chyme that enters the large intestine contains few nutrients except water, which is reabsorbed as the residue lingers in the large intestine, typically for 12 to 24 hours. Thus, it may not surprise you that the large intestine can be completely removed without significantly affecting digestive functioning. For example, in severe cases of inflammatory bowel disease, the large intestine can be removed by a procedure known as a colectomy. Often, a new fecal pouch can be crafted from the small intestine and sutured to the anus, but if not, an ileostomy can be created by bringing the distal ileum through the abdominal wall, allowing the watery chyme to be collected in a bag-like adhesive appliance.

Mechanical Digestion

In the large intestine, mechanical digestion begins when chyme moves from the ileum into the cecum, an activity regulated by the ileocecal sphincter. Right after you eat, peristalsis in the ileum forces chyme into the cecum. When the cecum is distended with chyme, contractions of the ileocecal sphincter strengthen. Once chyme enters the cecum, colon movements begin.

Mechanical digestion in the large intestine includes a combination of three types of movements. The presence of food residues in the colon stimulates a slow-moving haustral contraction. This type of movement involves sluggish segmentation, primarily in the transverse and descending colons. When a haustrum is distended with chyme, its muscle contracts, pushing the residue into the next haustrum. These contractions occur about every 30 minutes, and each last about 1 minute. These movements also mix the food residue, which helps the large intestine absorb water. The second type of movement is peristalsis, which, in the large intestine, is slower than in the more proximal portions of the alimentary canal. The third type is a mass movement. These strong waves start midway through the transverse colon and quickly force the contents toward the rectum. Mass movements usually occur three or four times per day, either while you eat or immediately afterward. Distension in the stomach and the breakdown products of digestion in the small intestine provoke the gastrocolic reflex, which increases motility, including mass movements, in the colon. Fiber in the diet both softens the stool and increases the power of colonic contractions, optimizing the activities of the colon.

Chemical Digestion

Although the glands of the large intestine secrete mucus, they do not secrete digestive enzymes. Therefore, chemical digestion in the large intestine occurs exclusively because of bacteria in the lumen of the colon. Through the process of saccharolytic fermentation, bacteria break down some of the remaining carbohydrates. This results in the discharge of hydrogen, carbon dioxide, and methane gases that create flatus (gas) in the colon; flatulence is excessive flatus. Each day, up to 1500 mL of flatus is produced in the colon. More is produced when you eat foods such as beans, which are rich in otherwise indigestible sugars and complex carbohydrates like soluble dietary fiber.

Absorption, Feces Formation, and Defecation

The small intestine absorbs about 90 percent of the water you ingest (either as liquid or within solid food). The large intestine absorbs most of the remaining water, a process that converts the liquid chyme residue into semisolid feces (“stool”). Feces is composed of undigested food residues, unabsorbed digested substances, millions of bacteria, old epithelial cells from the GI mucosa, inorganic salts, and enough water to let it pass smoothly out of the body. Of every 500 mL (17 ounces) of food residue that enters the cecum each day, about 150 mL (5 ounces) become feces.

Feces are eliminated through contractions of the rectal muscles. You help this process by a voluntary procedure called Valsalva’s maneuver, in which you increase intra-abdominal pressure by contracting your diaphragm and abdominal wall muscles, and closing your glottis.

The process of defecation begins when mass movements force feces from the colon into the rectum, stretching the rectal wall and provoking the defecation reflex, which eliminates feces from the rectum. This parasympathetic reflex is mediated by the spinal cord. It contracts the sigmoid colon and rectum, relaxes the internal anal sphincter, and initially contracts the external anal sphincter. The presence of feces in the anal canal sends a signal to the brain, which gives you the choice of voluntarily opening the external anal sphincter (defecating) or keeping it temporarily closed. If you decide to delay defecation, it takes a few seconds for the reflex contractions to stop and the rectal walls to relax. The next mass movement will trigger additional defecation reflexes until you defecate.

If defecation is delayed for an extended time, additional water is absorbed, making the feces firmer and potentially leading to constipation. On the other hand, if the waste matter moves too quickly through the intestines, not enough water is absorbed, and diarrhea can result. This can be caused by the ingestion of foodborne pathogens. In general, diet, health, and stress determine the frequency of bowel movements. The number of bowel movements varies greatly between individuals, ranging from two or three per day to three or four per week.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL By watching this animation you will see that for the various food groups—proteins, fats, and carbohydrates—digestion begins in different parts of the digestion system, though all end in the same place.[/caption]
[caption id="attachment_3019" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about the role of the intestines in digestion![/caption]
]]> 2698 2664 6 0 25.1 Physical Characteristics of Urine https://pressbooks.bccampus.ca/apdouglas2017/chapter/25-1-physical-characteristics-of-urine/ Mon, 17 Jul 2017 19:37:59 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2713

Learning Objectives

By the end of this section, you will be able to:
  • Compare and contrast blood plasma, glomerular filtrate, and urine characteristics
  • Describe the characteristics of a normal urine sample, including normal range of pH, osmolarity, and volume

The urinary system’s ability to filter the blood resides in about 2 to 3 million tufts of specialized capillaries—the glomeruli—distributed more or less equally between the two kidneys. Because the glomeruli filter the blood based mostly on particle size, large elements like blood cells, platelets, antibodies, and albumen are excluded. The glomerulus is the first part of the nephron, which then continues as a highly specialized tubular structure responsible for creating the final urine composition. All other solutes, such as ions, amino acids, vitamins, and wastes, are filtered to create a filtrate composition very similar to plasma. The glomeruli create about 200 liters (189 quarts) of this filtrate every day, yet you excrete less than two liters of waste you call urine.

Characteristics of the urine change, depending on influences such as water intake, exercise, environmental temperature, nutrient intake, and other factors (Table 1). Some of the characteristics such as color and odor are rough descriptors of your state of hydration. For example, if you exercise or work outside, and sweat a great deal, your urine will turn darker and produce a slight odor, even if you drink plenty of water. Athletes are often advised to consume water until their urine is clear. This is good advice; however, it takes time for the kidneys to process body fluids and store it in the bladder. Another way of looking at this is that the quality of the urine produced is an average over the time it takes to make that urine. Producing clear urine may take only a few minutes if you are drinking a lot of water or several hours if you are working outside and not drinking much.

Normal Urine Characteristics (Table 1)
Characteristic Normal values
Color Pale yellow to deep amber
Odor Odorless
Volume 750–2000 mL/24 hour
pH 4.5–8.0
Specific gravity 1.003–1.032
Osmolarity 40–1350 mOsmol/kg
Urobilinogen 0.2–1.0 mg/100 mL
White blood cells 0–2 HPF (per high-power field of microscope)
Leukocyte esterase None
Protein None or trace
Bilirubin <0.3 mg/100 mL
Ketones None
Nitrites None
Blood None
Glucose None

Urinalysis (urine analysis) often provides clues to renal disease. Normally, only traces of protein are found in urine, and when higher amounts are found, damage to the glomeruli is the likely basis. Unusually large quantities of urine may point to diseases like diabetes mellitus or hypothalamic tumors that cause diabetes insipidus. The color of urine is determined mostly by the breakdown products of red blood cell destruction (Figure 1). The “heme” of hemoglobin is converted by the liver into water-soluble forms that can be excreted into the bile and indirectly into the urine. This yellow pigment is urochrome. Urine color may also be affected by certain foods like beets, berries, and fava beans. A kidney stone or a cancer of the urinary system may produce sufficient bleeding to manifest as pink or even bright red urine. Diseases of the liver or obstructions of bile drainage from the liver impart a dark “tea” or “cola” hue to the urine. Dehydration produces darker, concentrated urine that may also possess the slight odor of ammonia. Most of the ammonia produced from protein breakdown is converted into urea by the liver, so ammonia is rarely detected in fresh urine. The strong ammonia odor you may detect in bathrooms or alleys is due to the breakdown of urea into ammonia by bacteria in the environment. About one in five people detect a distinctive odor in their urine after consuming asparagus; other foods such as onions, garlic, and fish can impart their own aromas! These food-caused odors are harmless.

[caption id="" align="aligncenter" width="230"]This color chart shows different shades of yellow and associates each shade with hydration or dehydration. Figure 1. Urine Color.[/caption]
Urine volume varies considerably. The normal range is one to two liters per day (Table 2). The kidneys must produce a minimum urine volume of about 500 mL/day to rid the body of wastes. Output below this level may be caused by severe dehydration or renal disease and is termed oliguria. The virtual absence of urine production is termed anuria. Excessive urine production is polyuria, which may be due to diabetes mellitus or diabetes insipidus. In diabetes mellitus, blood glucose levels exceed the number of available sodium-glucose transporters in the kidney, and glucose appears in the urine. The osmotic nature of glucose attracts water, leading to its loss in the urine. In the case of diabetes insipidus, insufficient pituitary antidiuretic hormone (ADH) release or insufficient numbers of ADH receptors in the collecting ducts means that too few water channels are inserted into the cell membranes that line the collecting ducts of the kidney. Insufficient numbers of water channels (aquaporins) reduce water absorption, resulting in high volumes of very dilute urine.
Urine Volumes (Table 2)
Volume condition Volume Causes
Normal 1–2 L/day
Polyuria >2.5 L/day Diabetes mellitus; diabetes insipidus; excess caffeine or alcohol; kidney disease; certain drugs, such as diuretics; sickle cell anemia; excessive water intake
Oliguria 300–500 mL/day Dehydration; blood loss; diarrhea; cardiogenic shock; kidney disease; enlarged prostate
Anuria <50 mL/day Kidney failure; obstruction, such as kidney stone or tumor; enlarged prostate

The pH (hydrogen ion concentration) of the urine can vary more than 1000-fold, from a normal low of 4.5 to a maximum of 8.0. Diet can influence pH; meats lower the pH, whereas citrus fruits, vegetables, and dairy products raise the pH. Chronically high or low pH can lead to disorders, such as the development of kidney stones or osteomalacia.

Specific gravity is a measure of the quantity of solutes per unit volume of a solution and is traditionally easier to measure than osmolarity. Urine will always have a specific gravity greater than pure water (water = 1.0) due to the presence of solutes. Laboratories can now measure urine osmolarity directly, which is a more accurate indicator of urinary solutes than specific gravity. Remember that osmolarity is the number of osmoles or milliosmoles per liter of fluid (mOsmol/L). Urine osmolarity ranges from a low of 50–100 mOsmol/L to as high as 1200 mOsmol/L H2O.

Cells are not normally found in the urine. The presence of leukocytes may indicate a urinary tract infection. Leukocyte esterase is released by leukocytes; if detected in the urine, it can be taken as indirect evidence of a urinary tract infection (UTI).

Protein does not normally leave the glomerular capillaries, so only trace amounts of protein should be found in the urine, approximately 10 mg/100 mL in a random sample. If excessive protein is detected in the urine, it usually means that the glomerulus is damaged and is allowing protein to “leak” into the filtrate.

Ketones are byproducts of fat metabolism. Finding ketones in the urine suggests that the body is using fat as an energy source in preference to glucose. In diabetes mellitus when there is not enough insulin (type I diabetes mellitus) or because of insulin resistance (type II diabetes mellitus), there is plenty of glucose, but without the action of insulin, the cells cannot take it up, so it remains in the bloodstream. Instead, the cells are forced to use fat as their energy source, and fat consumed at such a level produces excessive ketones as byproducts. These excess ketones will appear in the urine. Ketones may also appear if there is a severe deficiency of proteins or carbohydrates in the diet.

Nitrates (NO3–) occur normally in the urine. Gram-negative bacteria metabolize nitrate into nitrite (NO2–), and its presence in the urine is indirect evidence of infection.

There should be no blood found in the urine. It may sometimes appear in urine samples as a result of menstrual contamination, but this is not an abnormal condition. Now that you understand what the normal characteristics of urine are, the next section will introduce you to how you store and dispose of this waste product and how you make it.

[caption id="attachment_3023" align="aligncenter" width="150"] Watch this CrashCourse video to learn about the urinary system and the characteristics of urine.[/caption]]]> 2713 2711 1 0 25.5 Physiology of Urine Formation https://pressbooks.bccampus.ca/apdouglas2017/chapter/25-5-physiology-of-urine-formation/ Mon, 17 Jul 2017 19:40:07 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2732

Learning Objectives

By the end of this section, you will be able to:
  • Describe the hydrostatic and colloid osmotic forces that favor and oppose filtration
  • Describe glomerular filtration rate (GFR), state the average value of GFR, and explain how clearance rate can be used to measure GFR
  • Predict specific factors that will increase or decrease GFR
  • State the percent of the filtrate that is normally reabsorbed and explain why the process of reabsorption is so important
  • Calculate daily urine production
  • List common symptoms of kidney failure
Having reviewed the anatomy and microanatomy of the urinary system, now is the time to focus on the physiology. You will discover that different parts of the nephron utilize specific processes to produce urine: filtration, reabsorption, and secretion. You will learn how each of these processes works and where they occur along the nephron and collecting ducts. The physiologic goal is to modify the composition of the plasma and, in doing so, produce the waste product urine. Failure of the renal anatomy and/or physiology can lead suddenly or gradually to renal failure. In this event, a number of symptoms, signs, or laboratory findings point to the diagnosis (Table 3).
Symptoms of Kidney Failure (Table 3)
Weakness
Lethargy
Shortness of breath
Widespread edema
Anemia
Metabolic acidosis
Metabolic alkalosis
Heart arrhythmias
Uremia (high urea level in the blood)
Loss of appetite
Fatigue
Excessive urination
Oliguria (too little urine output)

Glomerular Filtration Rate (GFR)

The volume of filtrate formed by both kidneys per minute is termed the glomerular filtration rate (GFR). The heart pumps about 5 L blood per min under resting conditions. Approximately 20 percent or one liter enters the kidneys to be filtered. On average, this liter results in the production of about 125 mL/min filtrate produced in men (range of 90 to 140 mL/min) and 105 mL/min filtrate produced in women (range of 80 to 125 mL/min). This amount equates to a volume of about 180 L/day in men and 150 L/day in women. Ninety-nine percent of this filtrate is returned to the circulation by reabsorption so that only about 1–2 liters of urine are produced per day (Table 4).
Calculating Urine Formation per Day (Table 4)
Flow per minute (mL) Calculation
Renal blood flow 1050 Cardiac output is about 5000 mL/minute, of which 21 percent flows through the kidney.
5000*0.21 = 1050 mL blood/min
Renal plasma flow 578 Renal plasma flow equals the blood flow per minute times the hematocrit. If a person has a hematocrit of 45, then the renal plasma flow is 55 percent.
1050*0.55 = 578 mL plasma/min
Glomerular filtration rate 110 The GFR is the amount of plasma entering Bowman’s capsule per minute. It is the renal plasma flow times the fraction that enters the renal capsule (19 percent).
578*0.19 = 110 mL filtrate/min
Urine 1296 ml/day The filtrate not recovered by the kidney is the urine that will be eliminated. It is the GFR times the fraction of the filtrate that is not reabsorbed (0.8 percent).
110*.008 = 0.9 mL urine /min
Multiply urine/min times 60 minutes times 24 hours to get daily urine production.
0.9*60*24 = 1296 mL/day urine

GFR is influenced by the hydrostatic pressure and colloid osmotic pressure on either side of the capillary membrane of the glomerulus. Recall that filtration occurs as pressure forces fluid and solutes through a semipermeable barrier with the solute movement constrained by particle size. Hydrostatic pressure is the pressure produced by a fluid against a surface. If you have a fluid on both sides of a barrier, both fluids exert a pressure in opposing directions. Net fluid movement will be in the direction of the lower pressure. Osmosis is the movement of solvent (water) across a membrane that is impermeable to a solute in the solution. This creates a pressure, osmotic pressure, which will exist until the solute concentration is the same on both sides of a semipermeable membrane. As long as the concentration differs, water will move. Glomerular filtration occurs when glomerular hydrostatic pressure exceeds the luminal hydrostatic pressure of Bowman’s capsule. There is also an opposing force, the osmotic pressure, which is typically higher in the glomerular capillary.

To understand why this is so, look more closely at the microenvironment on either side of the filtration membrane. You will find osmotic pressure exerted by the solutes inside the lumen of the capillary as well as inside of Bowman’s capsule. Since the filtration membrane limits the size of particles crossing the membrane, the osmotic pressure inside the glomerular capillary is higher than the osmotic pressure in Bowman’s capsule. Recall that cells and the medium-to-large proteins cannot pass between the podocyte processes or through the fenestrations of the capillary endothelial cells. This means that red and white blood cells, platelets, albumins, and other proteins too large to pass through the filter remain in the capillary, creating an average colloid osmotic pressure of 30 mm Hg within the capillary. The absence of proteins in Bowman’s space (the lumen within Bowman’s capsule) results in an osmotic pressure near zero. Thus, the only pressure moving fluid across the capillary wall into the lumen of Bowman’s space is hydrostatic pressure. Hydrostatic (fluid) pressure is sufficient to push water through the membrane despite the osmotic pressure working against it. The sum of all of the influences, both osmotic and hydrostatic, results in a net filtration pressure (NFP) of about 10 mm Hg (Figure 1).

[caption id="" align="aligncenter" width="350"]This figure shows the different pressures acting across the glomerulus. Figure 1. Net Filtration Pressure. The NFP is the sum of osmotic and hydrostatic pressures.[/caption]
A proper concentration of solutes in the blood is important in maintaining osmotic pressure both in the glomerulus and systemically. There are disorders in which too much protein passes through the filtration slits into the kidney filtrate. This excess protein in the filtrate leads to a deficiency of circulating plasma proteins. In turn, the presence of protein in the urine increases its osmolarity; this holds more water in the filtrate and results in an increase in urine volume. Because there is less circulating protein, principally albumin, the osmotic pressure of the blood falls. Less osmotic pressure pulling water into the capillaries tips the balance towards hydrostatic pressure, which tends to push it out of the capillaries. The net effect is that water is lost from the circulation to interstitial tissues and cells. This “plumps up” the tissues and cells, a condition termed systemic edema.

Net Filtration Pressure (NFP)

NFP determines filtration rates through the kidney. It is determined as follows:

NFP = Glomerular blood hydrostatic pressure (GBHP) – [capsular hydrostatic pressure (CHP) + blood colloid osmotic pressure (BCOP)] = 10 mm Hg

That is:

NFP = GBHP – [CHP + BCOP] = 10 mm Hg

Or:

NFP = 55 – [15 + 30] = 10 mm Hg As you can see, there is a low net pressure across the filtration membrane. Intuitively, you should realize that minor changes in osmolarity of the blood or changes in capillary blood pressure result in major changes in the amount of filtrate formed at any given point in time. The kidney is able to cope with a wide range of blood pressures. In large part, this is due to the autoregulatory nature of smooth muscle. When you stretch it, it contracts. Thus, when blood pressure goes up, smooth muscle in the afferent capillaries contracts to limit any increase in blood flow and filtration rate. When blood pressure drops, the same capillaries relax to maintain blood flow and filtration rate. The net result is a relatively steady flow of blood into the glomerulus and a relatively steady filtration rate in spite of significant systemic blood pressure changes. Mean arterial blood pressure is calculated by adding 1/3 of the difference between the systolic and diastolic pressures to the diastolic pressure. Therefore, if the blood pressure is 110/80, the difference between systolic and diastolic pressure is 30. One third of this is 10, and when you add this to the diastolic pressure of 80, you arrive at a calculated mean arterial pressure of 90 mm Hg. Therefore, if you use mean arterial pressure for the GBHP in the formula for calculating NFP, you can determine that as long as mean arterial pressure is above approximately 60 mm Hg, the pressure will be adequate to maintain glomerular filtration. Blood pressures below this level will impair renal function and cause systemic disorders that are severe enough to threaten survival. This condition is called shock.

Determination of the GFR is one of the tools used to assess the kidney’s excretory function. This is more than just an academic exercise. Since many drugs are excreted in the urine, a decline in renal function can lead to toxic accumulations. Additionally, administration of appropriate drug dosages for those drugs primarily excreted by the kidney requires an accurate assessment of GFR. GFR can be estimated closely by intravenous administration of inulin. Inulin is a plant polysaccharide that is neither reabsorbed nor secreted by the kidney. Its appearance in the urine is directly proportional to the rate at which it is filtered by the renal corpuscle. However, since measuring inulin clearance is cumbersome in the clinical setting, most often, the GFR is estimated by measuring naturally occurring creatinine, a protein-derived molecule produced by muscle metabolism that is not reabsorbed and only slightly secreted by the nephron.

[caption id="attachment_3025" align="aligncenter" width="150"] Watch this CrashCourse video to learn about the production of urine.[/caption]
]]> 2732 2711 5 0 27.1 Anatomy and Physiology of the Male Reproductive System https://pressbooks.bccampus.ca/apdouglas2017/chapter/27-1-anatomy-and-physiology-of-the-male-reproductive-system/ Mon, 17 Jul 2017 19:43:15 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2773

Learning Objectives

By the end of this section, you will be able to:
  • Describe the structure and function of the organs of the male reproductive system
  • Describe the structure and function of the sperm cell
  • Explain the events during spermatogenesis that produce haploid sperm from diploid cells
  • Identify the importance of testosterone in male reproductive function
[caption id="attachment_3027" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of the male reproductive system.[/caption]

Unique for its role in human reproduction, a gamete is a specialized sex cell carrying 23 chromosomes—one half the number in body cells. At fertilization, the chromosomes in one male gamete, called a sperm (or spermatozoon), combine with the chromosomes in one female gamete, called an oocyte. The function of the male reproductive system (Figure 1) is to produce sperm and transfer them to the female reproductive tract. The paired testes are a crucial component in this process, as they produce both sperm and androgens, the hormones that support male reproductive physiology. In humans, the most important male androgen is testosterone. Several accessory organs and ducts aid the process of sperm maturation and transport the sperm and other seminal components to the penis, which delivers sperm to the female reproductive tract. In this section, we examine each of these different structures, and discuss the process of sperm production and transport.

[caption id="" align="aligncenter" width="550"]This figure shows the different organs in the male reproductive system. The top panel shows the side view of a man and an uncircumcised and a circumcised penis. The bottom panel shows the lateral view of the male reproductive system and the major parts are labeled. Figure 1. Male Reproductive System. The structures of the male reproductive system include the testes, the epididymides, the penis, and the ducts and glands that produce and carry semen. Sperm exit the scrotum through the ductus deferens, which is bundled in the spermatic cord. The seminal vesicles and prostate gland add fluids to the sperm to create semen.[/caption]

Scrotum

The testes are located in a skin-covered, highly pigmented, muscular sack called the scrotum that extends from the body behind the penis (see Figure 1). This location is important in sperm production, which occurs within the testes, and proceeds more efficiently when the testes are kept 2 to 4°C below core body temperature.

The dartos muscle makes up the subcutaneous muscle layer of the scrotum (Figure 2). It continues internally to make up the scrotal septum, a wall that divides the scrotum into two compartments, each housing one testis. Descending from the internal oblique muscle of the abdominal wall are the two cremaster muscles, which cover each testis like a muscular net. By contracting simultaneously, the dartos and cremaster muscles can elevate the testes in cold weather (or water), moving the testes closer to the body and decreasing the surface area of the scrotum to retain heat. Alternatively, as the environmental temperature increases, the scrotum relaxes, moving the testes farther from the body core and increasing scrotal surface area, which promotes heat loss. Externally, the scrotum has a raised medial thickening on the surface called the raphae.

[caption id="" align="aligncenter" width="550"]This figure shows the scrotum and testes. The left panel shows the external view of the scrotum, the middle panel shows the muscle layer and the right panel shows the deep tissues of the scrotum. Figure 2. The Scrotum and Testes. This anterior view shows the structures of the scrotum and testes.[/caption]

Testes

The testes (singular = testis) are the male gonads—that is, the male reproductive organs. They produce both sperm and androgens, such as testosterone, and are active throughout the reproductive lifespan of the male.

Paired ovals, the testes are each approximately 4 to 5 cm in length and are housed within the scrotum (see Figure 2). They are surrounded by two distinct layers of protective connective tissue (Figure 3). The outer tunica vaginalis is a serous membrane that has both a parietal and a thin visceral layer. Beneath the tunica vaginalis is the tunica albuginea, a tough, white, dense connective tissue layer covering the testis itself. Not only does the tunica albuginea cover the outside of the testis, it also invaginates to form septa that divide the testis into 300 to 400 structures called lobules. Within the lobules, sperm develop in structures called seminiferous tubules. During the seventh month of the developmental period of a male fetus, each testis moves through the abdominal musculature to descend into the scrotal cavity. This is called the “descent of the testis.” Cryptorchidism is the clinical term used when one or both of the testes fail to descend into the scrotum prior to birth. [caption id="" align="aligncenter" width="450"]This diagram shows the cross section of the testis. Figure 3. Anatomy of the Testis. This sagittal view shows the seminiferous tubules, the site of sperm production. Formed sperm are transferred to the epididymis, where they mature. They leave the epididymis during an ejaculation via the ductus deferens.[/caption]

The tightly coiled seminiferous tubules form the bulk of each testis. They are composed of developing sperm cells surrounding a lumen, the hollow center of the tubule, where formed sperm are released into the duct system of the testis. Specifically, from the lumens of the seminiferous tubules, sperm move into the straight tubules (or tubuli recti), and from there into a fine meshwork of tubules called the rete testes. Sperm leave the rete testes, and the testis itself, through the 15 to 20 efferent ductules that cross the tunica albuginea.

Inside the seminiferous tubules are six different cell types. These include supporting cells called sustentacular cells, as well as five types of developing sperm cells called germ cells. Germ cell development progresses from the basement membrane—at the perimeter of the tubule—toward the lumen. Let’s look more closely at these cell types.

Sertoli Cells

Surrounding all stages of the developing sperm cells are elongate, branching Sertoli cells. Sertoli cells are a type of supporting cell called a sustentacular cell, or sustenocyte, that are typically found in epithelial tissue. Sertoli cells secrete signaling molecules that promote sperm production and can control whether germ cells live or die. They extend physically around the germ cells from the peripheral basement membrane of the seminiferous tubules to the lumen. Tight junctions between these sustentacular cells create the blood–testis barrier, which keeps bloodborne substances from reaching the germ cells and, at the same time, keeps surface antigens on developing germ cells from escaping into the bloodstream and prompting an autoimmune response.

Germ Cells

The least mature cells, the spermatogonia (singular = spermatogonium), line the basement membrane inside the tubule. Spermatogonia are the stem cells of the testis, which means that they are still able to differentiate into a variety of different cell types throughout adulthood. Spermatogonia divide to produce primary and secondary spermatocytes, then spermatids, which finally produce formed sperm. The process that begins with spermatogonia and concludes with the production of sperm is called spermatogenesis.

Spermatogenesis

As just noted, spermatogenesis occurs in the seminiferous tubules that form the bulk of each testis (see Figure 3). The process begins at puberty, after which time sperm are produced constantly throughout a man’s life. One production cycle, from spermatogonia through formed sperm, takes approximately 64 days. A new cycle starts approximately every 16 days, although this timing is not synchronous across the seminiferous tubules. Sperm counts—the total number of sperm a man produces—slowly decline after age 35, and some studies suggest that smoking can lower sperm counts irrespective of age.

The process of spermatogenesis begins with mitosis of the diploid spermatogonia (Figure 4). Because these cells are diploid (2n), they each have a complete copy of the father’s genetic material, or 46 chromosomes. However, mature gametes are haploid (1n), containing 23 chromosomes—meaning that daughter cells of spermatogonia must undergo a second cellular division through the process of meiosis. [caption id="" align="aligncenter" width="550"]This figure shows the steps in spermatogenesis. The left panel shows a flow chart that outlines the different steps in the formation of sperm. The right panel shows a micrograph with the cross section of a seminiferous tubule. Figure 4. Spermatogenesis. (a) Mitosis of a spermatogonial stem cell involves a single cell division that results in two identical, diploid daughter cells (spermatogonia to primary spermatocyte). Meiosis has two rounds of cell division: primary spermatocyte to secondary spermatocyte, and then secondary spermatocyte to spermatid. This produces four haploid daughter cells (spermatids). (b) In this electron micrograph of a cross-section of a seminiferous tubule from a rat, the lumen is the light-shaded area in the center of the image. The location of the primary spermatocytes is near the basement membrane, and the early spermatids are approaching the lumen (tissue source: rat). EM × 900. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Two identical diploid cells result from spermatogonia mitosis. One of these cells remains a spermatogonium, and the other becomes a primary spermatocyte, the next stage in the process of spermatogenesis. As in mitosis, DNA is replicated in a primary spermatocyte, and the cell undergoes cell division to produce two cells with identical chromosomes. Each of these is a secondary spermatocyte. Now a second round of cell division occurs in both of the secondary spermatocytes, separating the chromosome pairs. This second meiotic division results in a total of four cells with only half of the number of chromosomes. Each of these new cells is a spermatid. Although haploid, early spermatids look very similar to cells in the earlier stages of spermatogenesis, with a round shape, central nucleus, and large amount of cytoplasm. A process called spermiogenesis transforms these early spermatids, reducing the cytoplasm, and beginning the formation of the parts of a true sperm. The fifth stage of germ cell formation—spermatozoa, or formed sperm—is the end result of this process, which occurs in the portion of the tubule nearest the lumen. Eventually, the sperm are released into the lumen and are moved along a series of ducts in the testis toward a structure called the epididymis for the next step of sperm maturation.

Structure of Formed Sperm

Sperm are smaller than most cells in the body; in fact, the volume of a sperm cell is 85,000 times less than that of the female gamete. Approximately 100 to 300 million sperm are produced each day, whereas women typically ovulate only one oocyte per month as is true for most cells in the body, the structure of sperm cells speaks to their function. Sperm have a distinctive head, mid-piece, and tail region (Figure 5). The head of the sperm contains the extremely compact haploid nucleus with very little cytoplasm. These qualities contribute to the overall small size of the sperm (the head is only 5 μm long). A structure called the acrosome covers most of the head of the sperm cell as a “cap” that is filled with lysosomal enzymes important for preparing sperm to participate in fertilization. Tightly packed mitochondria fill the mid-piece of the sperm. ATP produced by these mitochondria will power the flagellum, which extends from the neck and the mid-piece through the tail of the sperm, enabling it to move the entire sperm cell. The central strand of the flagellum, the axial filament, is formed from one centriole inside the maturing sperm cell during the final stages of spermatogenesis. [caption id="" align="aligncenter" width="550"]This diagram shows the structure of sperm; the major parts are labeled. Figure 5. Structure of Sperm. Sperm cells are divided into a head, containing DNA; a mid-piece, containing mitochondria; and a tail, providing motility. The acrosome is oval and somewhat flattened.[/caption]

Sperm Transport

To fertilize an egg, sperm must be moved from the seminiferous tubules in the testes, through the epididymis, and—later during ejaculation—along the length of the penis and out into the female reproductive tract.

Role of the Epididymis

From the lumen of the seminiferous tubules, the immotile sperm are surrounded by testicular fluid and moved to the epididymis (plural = epididymides), a coiled tube attached to the testis where newly formed sperm continue to mature (see Figure 3). Though the epididymis does not take up much room in its tightly coiled state, it would be approximately 6 m (20 feet) long if straightened. It takes an average of 12 days for sperm to move through the coils of the epididymis, with the shortest recorded transit time in humans being one day. Sperm enter the head of the epididymis and are moved along predominantly by the contraction of smooth muscles lining the epididymal tubes. As they are moved along the length of the epididymis, the sperm further mature and acquire the ability to move under their own power. Once inside the female reproductive tract, they will use this ability to move independently toward the unfertilized egg. The more mature sperm are then stored in the tail of the epididymis (the final section) until ejaculation occurs.

Duct System

During ejaculation, sperm exit the tail of the epididymis and are pushed by smooth muscle contraction to the ductus deferens (also called the vas deferens). The ductus deferens is a thick, muscular tube that is bundled together inside the scrotum with connective tissue, blood vessels, and nerves into a structure called the spermatic cord (see Figure 1 and Figure 2). Because the ductus deferens is physically accessible within the scrotum, surgical sterilization to interrupt sperm delivery can be performed by cutting and sealing a small section of the ductus (vas) deferens. This procedure is called a vasectomy, and it is an effective form of male birth control. Although it may be possible to reverse a vasectomy, clinicians consider the procedure permanent, and advise men to undergo it only if they are certain they no longer wish to father children.

Interactive Link Feature
[caption id="" align="aligncenter" width="130"]QR Code representing a URL Watch this video to learn about a vasectomy.[/caption]

From each epididymis, each ductus deferens extends superiorly into the abdominal cavity through the inguinal canal in the abdominal wall. From here, the ductus deferens continues posteriorly to the pelvic cavity, ending posterior to the bladder where it dilates in a region called the ampulla (meaning “flask”).

Sperm make up only 5 percent of the final volume of semen, the thick, milky fluid that the male ejaculates. The bulk of semen is produced by three critical accessory glands of the male reproductive system: the seminal vesicles, the prostate, and the bulbourethral glands.

Seminal Vesicles

As sperm pass through the ampulla of the ductus deferens at ejaculation, they mix with fluid from the associated seminal vesicle (see Figure 1). The paired seminal vesicles are glands that contribute approximately 60 percent of the semen volume. Seminal vesicle fluid contains large amounts of fructose, which is used by the sperm mitochondria to generate ATP to allow movement through the female reproductive tract.

The fluid, now containing both sperm and seminal vesicle secretions, next moves into the associated ejaculatory duct, a short structure formed from the ampulla of the ductus deferens and the duct of the seminal vesicle. The paired ejaculatory ducts transport the seminal fluid into the next structure, the prostate gland.

Prostate Gland

As shown in Figure 1, the centrally located prostate gland sits anterior to the rectum at the base of the bladder surrounding the prostatic urethra (the portion of the urethra that runs within the prostate). About the size of a walnut, the prostate is formed of both muscular and glandular tissues. It excretes an alkaline, milky fluid to the passing seminal fluid—now called semen—that is critical to first coagulate and then decoagulate the semen following ejaculation. The temporary thickening of semen helps retain it within the female reproductive tract, providing time for sperm to utilize the fructose provided by seminal vesicle secretions. When the semen regains its fluid state, sperm can then pass farther into the female reproductive tract.

The prostate normally doubles in size during puberty. At approximately age 25, it gradually begins to enlarge again. This enlargement does not usually cause problems; however, abnormal growth of the prostate, or benign prostatic hyperplasia (BPH), can cause constriction of the urethra as it passes through the middle of the prostate gland, leading to a number of lower urinary tract symptoms, such as a frequent and intense urge to urinate, a weak stream, and a sensation that the bladder has not emptied completely. By age 60, approximately 40 percent of men have some degree of BPH. By age 80, the number of affected individuals has jumped to as many as 80 percent. Treatments for BPH attempt to relieve the pressure on the urethra so that urine can flow more normally. Mild to moderate symptoms are treated with medication, whereas severe enlargement of the prostate is treated by surgery in which a portion of the prostate tissue is removed.

Another common disorder involving the prostate is prostate cancer. According to the Centers for Disease Control and Prevention (CDC), prostate cancer is the second most common cancer in men. However, some forms of prostate cancer grow very slowly and thus may not ever require treatment. Aggressive forms of prostate cancer, in contrast, involve metastasis to vulnerable organs like the lungs and brain. There is no link between BPH and prostate cancer, but the symptoms are similar. Prostate cancer is detected by a medical history, a blood test, and a rectal exam that allows physicians to palpate the prostate and check for unusual masses. If a mass is detected, the cancer diagnosis is confirmed by biopsy of the cells.

Bulbourethral Glands

The final addition to semen is made by two bulbourethral glands (or Cowper’s glands) that release a thick, salty fluid that lubricates the end of the urethra and the vagina, and helps to clean urine residues from the penile urethra. The fluid from these accessory glands is released after the male becomes sexually aroused, and shortly before the release of the semen. It is therefore sometimes called pre-ejaculate. It is important to note that, in addition to the lubricating proteins, it is possible for bulbourethral fluid to pick up sperm already present in the urethra, and therefore it may be able to cause pregnancy.

Interactive Link Feature
[caption id="" align="aligncenter" width="130"]QR Code representing a URL Watch this video to explore the structures of the male reproductive system and the path of sperm, which starts in the testes and ends as the sperm leave the penis through the urethra.[/caption]

The Penis

The penis is the male organ of copulation (sexual intercourse). It is flaccid for non-sexual actions, such as urination, and turgid and rod-like with sexual arousal. When erect, the stiffness of the organ allows it to penetrate into the vagina and deposit semen into the female reproductive tract. [caption id="" align="aligncenter" width="550"]This multipart diagram shows the cross section of the penis. The top left panel shows the lateral view of the flaccid penis and the top right panel shows the transverse view. The bottom left panel shows the lateral view of the erect penis and the bottom right panel shows the transverse view. Figure 6. Cross-Sectional Anatomy of the Penis. Three columns of erectile tissue make up most of the volume of the penis.[/caption]

The shaft of the penis surrounds the urethra (Figure 6). The shaft is composed of three column-like chambers of erectile tissue that span the length of the shaft. Each of the two larger lateral chambers is called a corpus cavernosum (plural = corpora cavernosa). Together, these make up the bulk of the penis. The corpus spongiosum, which can be felt as a raised ridge on the erect penis, is a smaller chamber that surrounds the spongy, or penile, urethra. The end of the penis, called the glans penis, has a high concentration of nerve endings, resulting in very sensitive skin that influences the likelihood of ejaculation (see Figure 1). The skin from the shaft extends down over the glans and forms a collar called the prepuce (or foreskin). The foreskin also contains a dense concentration of nerve endings, and both lubricate and protect the sensitive skin of the glans penis. A surgical procedure called circumcision, often performed for religious or social reasons, removes the prepuce, typically within days of birth.

Both sexual arousal and REM sleep (during which dreaming occurs) can induce an erection. Penile erections are the result of vasocongestion, or engorgement of the tissues because of more arterial blood flowing into the penis than is leaving in the veins. During sexual arousal, nitric oxide (NO) is released from nerve endings near blood vessels within the corpora cavernosa and spongiosum. Release of NO activates a signaling pathway that results in relaxation of the smooth muscles that surround the penile arteries, causing them to dilate. This dilation increases the amount of blood that can enter the penis and induces the endothelial cells in the penile arterial walls to also secrete NO and perpetuate the vasodilation. The rapid increase in blood volume fills the erectile chambers, and the increased pressure of the filled chambers compresses the thin-walled penile venules, preventing venous drainage of the penis. The result of this increased blood flow to the penis and reduced blood return from the penis is erection. Depending on the flaccid dimensions of a penis, it can increase in size slightly or greatly during erection, with the average length of an erect penis measuring approximately 15 cm.

Disorders of the… Feature

Male Reproductive SystemErectile dysfunction (ED) is a condition in which a man has difficulty either initiating or maintaining an erection. The combined prevalence of minimal, moderate, and complete ED is approximately 40 percent in men at age 40, and reaches nearly 70 percent by 70 years of age. In addition to aging, ED is associated with diabetes, vascular disease, psychiatric disorders, prostate disorders, the use of some drugs such as certain antidepressants, and problems with the testes resulting in low testosterone concentrations. These physical and emotional conditions can lead to interruptions in the vasodilation pathway and result in an inability to achieve an erection.

Recall that the release of NO induces relaxation of the smooth muscles that surround the penile arteries, leading to the vasodilation necessary to achieve an erection. To reverse the process of vasodilation, an enzyme called phosphodiesterase (PDE) degrades a key component of the NO signaling pathway called cGMP. There are several different forms of this enzyme, and PDE type 5 is the type of PDE found in the tissues of the penis. Scientists discovered that inhibiting PDE5 increases blood flow, and allows vasodilation of the penis to occur.

PDEs and the vasodilation signaling pathway are found in the vasculature in other parts of the body. In the 1990s, clinical trials of a PDE5 inhibitor called sildenafil were initiated to treat hypertension and angina pectoris (chest pain caused by poor blood flow through the heart). The trial showed that the drug was not effective at treating heart conditions, but many men experienced erection and priapism (erection lasting longer than 4 hours). Because of this, a clinical trial was started to investigate the ability of sildenafil to promote erections in men suffering from ED. In 1998, the FDA approved the drug, marketed as Viagra®. Since approval of the drug, sildenafil and similar PDE inhibitors now generate over a billion dollars a year in sales, and are reported to be effective in treating approximately 70 to 85 percent of cases of ED. Importantly, men with health problems—especially those with cardiac disease taking nitrates—should avoid Viagra or talk to their physician to find out if they are a candidate for the use of this drug, as deaths have been reported for at-risk users.

Testosterone

Testosterone, an androgen, is a steroid hormone produced by Leydig cells. The alternate term for Leydig cells, interstitial cells, reflects their location between the seminiferous tubules in the testes. In male embryos, testosterone is secreted by Leydig cells by the seventh week of development, with peak concentrations reached in the second trimester. This early release of testosterone results in the anatomical differentiation of the male sexual organs. In childhood, testosterone concentrations are low. They increase during puberty, activating characteristic physical changes and initiating spermatogenesis.

Functions of Testosterone

The continued presence of testosterone is necessary to keep the male reproductive system working properly, and Leydig cells produce approximately 6 to 7 mg of testosterone per day. Testicular steroidogenesis (the manufacture of androgens, including testosterone) results in testosterone concentrations that are 100 times higher in the testes than in the circulation. Maintaining these normal concentrations of testosterone promotes spermatogenesis, whereas low levels of testosterone can lead to infertility. In addition to intratesticular secretion, testosterone is also released into the systemic circulation and plays an important role in muscle development, bone growth, the development of secondary sex characteristics, and maintaining libido (sex drive) in both males and females. In females, the ovaries secrete small amounts of testosterone, although most is converted to estradiol. A small amount of testosterone is also secreted by the adrenal glands in both sexes.

Control of Testosterone

The regulation of testosterone concentrations throughout the body is critical for male reproductive function. The intricate interplay between the endocrine system and the reproductive system is shown in Figure 7. [caption id="" align="aligncenter" width="550"]This figure shows the steps in the regulation of testosterone production. The top panel shows the hypothalamus and the bottom panel shows two micrographs. The left micrograph is that of sertoli cells and the right micrograph is that of Leydig cells. Figure 7. Regulation of Testosterone Production. The hypothalamus and pituitary gland regulate the production of testosterone and the cells that assist in spermatogenesis. GnRH activates the anterior pituitary to produce LH and FSH, which in turn stimulate Leydig cells and Sertoli cells, respectively. The system is a negative feedback loop because the end products of the pathway, testosterone and inhibin, interact with the activity of GnRH to inhibit their own production.[/caption]

The regulation of Leydig cell production of testosterone begins outside of the testes. The hypothalamus and the pituitary gland in the brain integrate external and internal signals to control testosterone synthesis and secretion. The regulation begins in the hypothalamus. Pulsatile release of a hormone called gonadotropin-releasing hormone (GnRH) from the hypothalamus stimulates the endocrine release of hormones from the pituitary gland. Binding of GnRH to its receptors on the anterior pituitary gland stimulates release of the two gonadotropins: luteinizing hormone (LH) and follicle-stimulating hormone (FSH). These two hormones are critical for reproductive function in both men and women. In men, FSH binds predominantly to the Sertoli cells within the seminiferous tubules to promote spermatogenesis. FSH also stimulates the Sertoli cells to produce hormones called inhibins, which function to inhibit FSH release from the pituitary, thus reducing testosterone secretion. These polypeptide hormones correlate directly with Sertoli cell function and sperm number; inhibin B can be used as a marker of spermatogenic activity. In men, LH binds to receptors on Leydig cells in the testes and upregulates the production of testosterone.

A negative feedback loop predominantly controls the synthesis and secretion of both FSH and LH. Low blood concentrations of testosterone stimulate the hypothalamic release of GnRH. GnRH then stimulates the anterior pituitary to secrete LH into the bloodstream. In the testis, LH binds to LH receptors on Leydig cells and stimulates the release of testosterone. When concentrations of testosterone in the blood reach a critical threshold, testosterone itself will bind to androgen receptors on both the hypothalamus and the anterior pituitary, inhibiting the synthesis and secretion of GnRH and LH, respectively. When the blood concentrations of testosterone once again decline, testosterone no longer interacts with the receptors to the same degree and GnRH and LH are once again secreted, stimulating more testosterone production. This same process occurs with FSH and inhibin to control spermatogenesis.

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]]> 2773 2763 1 0 27.2 Anatomy and Physiology of the Female Reproductive System https://pressbooks.bccampus.ca/apdouglas2017/chapter/27-2-anatomy-and-physiology-of-the-female-reproductive-system/ Mon, 17 Jul 2017 19:44:09 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2783

Learning Objectives

By the end of this section, you will be able to:
  • Describe the structure and function of the organs of the female reproductive system
  • List the steps of oogenesis
  • Describe the hormonal changes that occur during the ovarian and menstrual cycles
  • Trace the path of an oocyte from ovary to fertilization
[caption id="attachment_3029" align="aligncenter" width="150"] Watch this CrashCourse video for an overview of the female reproductive system.[/caption] The female reproductive system functions to produce gametes and reproductive hormones, just like the male reproductive system; however, it also has the additional task of supporting the developing fetus and delivering it to the outside world. Unlike its male counterpart, the female reproductive system is located primarily inside the pelvic cavity (Figure 1). Recall that the ovaries are the female gonads. The gamete they produce is called an oocyte. We’ll discuss the production of oocytes in detail shortly. First, let’s look at some of the structures of the female reproductive system. [caption id="" align="aligncenter" width="555"]This figure shows the structure and the different organs in the female reproductive system. The top panel shows the lateral view and the bottom panel shows the anterior view. Figure 1. Female Reproductive System. The major organs of the female reproductive system are located inside the pelvic cavity.[/caption]

External Female Genitals

The external female reproductive structures are referred to collectively as the vulva (Figure 2). The mons pubis is a pad of fat that is located at the anterior, over the pubic bone. After puberty, it becomes covered in pubic hair. The labia majora (labia = “lips”; majora = “larger”) are folds of hair-covered skin that begin just posterior to the mons pubis. The thinner and more pigmented labia minora (labia = “lips”; minora = “smaller”) extend medial to the labia majora. Although they naturally vary in shape and size from woman to woman, the labia minora serve to protect the female urethra and the entrance to the female reproductive tract.

The superior, anterior portions of the labia minora come together to encircle the clitoris (or glans clitoris), an organ that originates from the same cells as the glans penis and has abundant nerves that make it important in sexual sensation and orgasm. The hymen is a thin membrane that sometimes partially covers the entrance to the vagina. An intact hymen cannot be used as an indication of “virginity”; even at birth, this is only a partial membrane, as menstrual fluid and other secretions must be able to exit the body, regardless of penile–vaginal intercourse. The vaginal opening is located between the opening of the urethra and the anus. It is flanked by outlets to the Bartholin’s glands (or greater vestibular glands). [caption id="" align="aligncenter" width="550"]This figure shows the parts of the vulva. The right panel shows the external anterior view and the left panel shows the internal anteriolateral view. The major parts are labeled. Figure 2. The Vulva. The external female genitalia are referred to collectively as the vulva.[/caption]

Vagina

The vagina, shown at the bottom of Figure 1 and Figure 1, is a muscular canal (approximately 10 cm long) that serves as the entrance to the reproductive tract. It also serves as the exit from the uterus during menses and childbirth. The outer walls of the anterior and posterior vagina are formed into longitudinal columns, or ridges, and the superior portion of the vagina—called the fornix—meets the protruding uterine cervix. The walls of the vagina are lined with an outer, fibrous adventitia; a middle layer of smooth muscle; and an inner mucous membrane with transverse folds called rugae. Together, the middle and inner layers allow the expansion of the vagina to accommodate intercourse and childbirth. The thin, perforated hymen can partially surround the opening to the vaginal orifice. The hymen can be ruptured with strenuous physical exercise, penile–vaginal intercourse, and childbirth. The Bartholin’s glands and the lesser vestibular glands (located near the clitoris) secrete mucus, which keeps the vestibular area moist.

The vagina is home to a normal population of microorganisms that help to protect against infection by pathogenic bacteria, yeast, or other organisms that can enter the vagina. In a healthy woman, the most predominant type of vaginal bacteria is from the genus Lactobacillus. This family of beneficial bacterial flora secretes lactic acid, and thus protects the vagina by maintaining an acidic pH (below 4.5). Potential pathogens are less likely to survive in these acidic conditions. Lactic acid, in combination with other vaginal secretions, makes the vagina a self-cleansing organ. However, douching—or washing out the vagina with fluid—can disrupt the normal balance of healthy microorganisms, and actually increase a woman’s risk for infections and irritation. Indeed, the American College of Obstetricians and Gynecologists recommend that women do not douche, and that they allow the vagina to maintain its normal healthy population of protective microbial flora.

Ovaries

The ovaries are the female gonads (see Figure 1). Paired ovals, they are each about 2 to 3 cm in length, about the size of an almond. The ovaries are located within the pelvic cavity, and are supported by the mesovarium, an extension of the peritoneum that connects the ovaries to the broad ligament. Extending from the mesovarium itself is the suspensory ligament that contains the ovarian blood and lymph vessels. Finally, the ovary itself is attached to the uterus via the ovarian ligament.

The ovary comprises an outer covering of cuboidal epithelium called the ovarian surface epithelium that is superficial to a dense connective tissue covering called the tunica albuginea. Beneath the tunica albuginea is the cortex, or outer portion, of the organ. The cortex is composed of a tissue framework called the ovarian stroma that forms the bulk of the adult ovary. Oocytes develop within the outer layer of this stroma, each surrounded by supporting cells. This grouping of an oocyte and its supporting cells is called a follicle. The growth and development of ovarian follicles will be described shortly. Beneath the cortex lies the inner ovarian medulla, the site of blood vessels, lymph vessels, and the nerves of the ovary. You will learn more about the overall anatomy of the female reproductive system at the end of this section.

The Ovarian Cycle

The ovarian cycle is a set of predictable changes in a female’s oocytes and ovarian follicles. During a woman’s reproductive years, it is a roughly 28-day cycle that can be correlated with, but is not the same as, the menstrual cycle (discussed shortly). The cycle includes two interrelated processes: oogenesis (the production of female gametes) and folliculogenesis (the growth and development of ovarian follicles).

Oogenesis

Gametogenesis in females is called oogenesis. The process begins with the ovarian stem cells, or oogonia (Figure 3). Oogonia are formed during fetal development, and divide via mitosis, much like spermatogonia in the testis. Unlike spermatogonia, however, oogonia form primary oocytes in the fetal ovary prior to birth. These primary oocytes are then arrested in this stage of meiosis I, only to resume it years later, beginning at puberty and continuing until the woman is near menopause (the cessation of a woman’s reproductive functions). The number of primary oocytes present in the ovaries declines from one to two million in an infant, to approximately 400,000 at puberty, to zero by the end of menopause.

The initiation of ovulation—the release of an oocyte from the ovary—marks the transition from puberty into reproductive maturity for women. From then on, throughout a woman’s reproductive years, ovulation occurs approximately once every 28 days. Just prior to ovulation, a surge of luteinizing hormone triggers the resumption of meiosis in a primary oocyte. This initiates the transition from primary to secondary oocyte. However, as you can see in Figure 3, this cell division does not result in two identical cells. Instead, the cytoplasm is divided unequally, and one daughter cell is much larger than the other. This larger cell, the secondary oocyte, eventually leaves the ovary during ovulation. The smaller cell, called the first polar body, may or may not complete meiosis and produce second polar bodies; in either case, it eventually disintegrates. Therefore, even though oogenesis produces up to four cells, only one survives. [caption id="" align="aligncenter" width="550"]This flowchart shows the formation of oocytes in the female. The top half of the flowchart is before birth and the bottom half is after puberty. A callout to the left also shows the eggs before and after sperm penetration. Figure 3. Oogenesis. The unequal cell division of oogenesis produces one to three polar bodies that later degrade, as well as a single haploid ovum, which is produced only if there is penetration of the secondary oocyte by a sperm cell.[/caption]

How does the diploid secondary oocyte become an ovum—the haploid female gamete? Meiosis of a secondary oocyte is completed only if a sperm succeeds in penetrating its barriers. Meiosis II then resumes, producing one haploid ovum that, at the instant of fertilization by a (haploid) sperm, becomes the first diploid cell of the new offspring (a zygote). Thus, the ovum can be thought of as a brief, transitional, haploid stage between the diploid oocyte and diploid zygote.

The larger amount of cytoplasm contained in the female gamete is used to supply the developing zygote with nutrients during the period between fertilization and implantation into the uterus. Interestingly, sperm contribute only DNA at fertilization —not cytoplasm. Therefore, the cytoplasm and all of the cytoplasmic organelles in the developing embryo are of maternal origin. This includes mitochondria, which contain their own DNA. Scientific research in the 1980s determined that mitochondrial DNA was maternally inherited, meaning that you can trace your mitochondrial DNA directly to your mother, her mother, and so on back through your female ancestors.

Everyday Connections Feature

Mapping Human History with Mitochondrial DNAWhen we talk about human DNA, we’re usually referring to nuclear DNA; that is, the DNA coiled into chromosomal bundles in the nucleus of our cells. We inherit half of our nuclear DNA from our father, and half from our mother. However, mitochondrial DNA (mtDNA) comes only from the mitochondria in the cytoplasm of the fat ovum we inherit from our mother. She received her mtDNA from her mother, who got it from her mother, and so on. Each of our cells contains approximately 1700 mitochondria, with each mitochondrion packed with mtDNA containing approximately 37 genes.

Mutations (changes) in mtDNA occur spontaneously in a somewhat organized pattern at regular intervals in human history. By analyzing these mutational relationships, researchers have been able to determine that we can all trace our ancestry back to one woman who lived in Africa about 200,000 years ago. Scientists have given this woman the biblical name Eve, although she is not, of course, the first Homo sapiens female. More precisely, she is our most recent common ancestor through matrilineal descent.

This doesn’t mean that everyone’s mtDNA today looks exactly like that of our ancestral Eve. Because of the spontaneous mutations in mtDNA that have occurred over the centuries, researchers can map different “branches” off of the “main trunk” of our mtDNA family tree. Your mtDNA might have a pattern of mutations that aligns more closely with one branch, and your neighbor’s may align with another branch. Still, all branches eventually lead back to Eve.

But what happened to the mtDNA of all of the other Homo sapiens females who were living at the time of Eve? Researchers explain that, over the centuries, their female descendants died childless or with only male children, and thus, their maternal line—and its mtDNA—ended.

Folliculogenesis

Again, ovarian follicles are oocytes and their supporting cells. They grow and develop in a process called folliculogenesis, which typically leads to ovulation of one follicle approximately every 28 days, along with death to multiple other follicles. The death of ovarian follicles is called atresia, and can occur at any point during follicular development. Recall that, a female infant at birth will have one to two million oocytes within her ovarian follicles, and that this number declines throughout life until menopause, when no follicles remain. As you’ll see next, follicles progress from primordial, to primary, to secondary and tertiary stages prior to ovulation—with the oocyte inside the follicle remaining as a primary oocyte until right before ovulation.

Folliculogenesis begins with follicles in a resting state. These small primordial follicles are present in newborn females and are the prevailing follicle type in the adult ovary (Figure 4). Primordial follicles have only a single flat layer of support cells, called granulosa cells, that surround the oocyte, and they can stay in this resting state for years—some until right before menopause.

After puberty, a few primordial follicles will respond to a recruitment signal each day, and will join a pool of immature growing follicles called primary follicles. Primary follicles start with a single layer of granulosa cells, but the granulosa cells then become active and transition from a flat or squamous shape to a rounded, cuboidal shape as they increase in size and proliferate. As the granulosa cells divide, the follicles—now called secondary follicles (see Figure 4)—increase in diameter, adding a new outer layer of connective tissue, blood vessels, and theca cells—cells that work with the granulosa cells to produce estrogens.

Within the growing secondary follicle, the primary oocyte now secretes a thin acellular membrane called the zona pellucida that will play a critical role in fertilization. A thick fluid, called follicular fluid, that has formed between the granulosa cells also begins to collect into one large pool, or antrum. Follicles in which the antrum has become large and fully formed are considered tertiary follicles (or antral follicles). Several follicles reach the tertiary stage at the same time, and most of these will undergo atresia. The one that does not die will continue to grow and develop until ovulation, when it will expel its secondary oocyte surrounded by several layers of granulosa cells from the ovary. Keep in mind that most follicles don’t make it to this point. In fact, roughly 99 percent of the follicles in the ovary will undergo atresia, which can occur at any stage of folliculogenesis. [caption id="" align="aligncenter" width="550"]This multipart figure shows how follicles are generated. The top panel shows the six stages of folliculogenesis. In each stage, the major cell types are labeled. The bottom part shows a micrograph of a secondary follicle and the major parts are labeled. Figure 4. Folliculogenesis. (a) The maturation of a follicle is shown in a clockwise direction proceeding from the primordial follicles. FSH stimulates the growth of a tertiary follicle, and LH stimulates the production of estrogen by granulosa and theca cells. Once the follicle is mature, it ruptures and releases the oocyte. Cells remaining in the follicle then develop into the corpus luteum. (b) In this electron micrograph of a secondary follicle, the oocyte, theca cells (thecae folliculi), and developing antrum are clearly visible. EM × 1100. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Hormonal Control of the Ovarian Cycle

The process of development that we have just described, from primordial follicle to early tertiary follicle, takes approximately two months in humans. The final stages of development of a small cohort of tertiary follicles, ending with ovulation of a secondary oocyte, occur over a course of approximately 28 days. These changes are regulated by many of the same hormones that regulate the male reproductive system, including GnRH, LH, and FSH.

As in men, the hypothalamus produces GnRH, a hormone that signals the anterior pituitary gland to produce the gonadotropins FSH and LH (Figure 5). These gonadotropins leave the pituitary and travel through the bloodstream to the ovaries, where they bind to receptors on the granulosa and theca cells of the follicles. FSH stimulates the follicles to grow (hence its name of follicle-stimulating hormone), and the five or six tertiary follicles expand in diameter. The release of LH also stimulates the granulosa and theca cells of the follicles to produce the sex steroid hormone estradiol, a type of estrogen. This phase of the ovarian cycle, when the tertiary follicles are growing and secreting estrogen, is known as the follicular phase.

The more granulosa and theca cells a follicle has (that is, the larger and more developed it is), the more estrogen it will produce in response to LH stimulation. As a result of these large follicles producing large amounts of estrogen, systemic plasma estrogen concentrations increase. Following a classic negative feedback loop, the high concentrations of estrogen will stimulate the hypothalamus and pituitary to reduce the production of GnRH, LH, and FSH. Because the large tertiary follicles require FSH to grow and survive at this point, this decline in FSH caused by negative feedback leads most of them to die (atresia). Typically only one follicle, now called the dominant follicle, will survive this reduction in FSH, and this follicle will be the one that releases an oocyte. Scientists have studied many factors that lead to a particular follicle becoming dominant: size, the number of granulosa cells, and the number of FSH receptors on those granulosa cells all contribute to a follicle becoming the one surviving dominant follicle. [caption id="" align="aligncenter" width="550"]This figure shows three flowcharts. The flowchart on the top left shows the hormonal regulation of the follicular phase. The flowchart on the top right shows the hormonal regulation of the ovulation phase. The bottom flowchart shows the hormonal regulation of luteal phase. Figure 5. Hormonal Regulation of Ovulation. The hypothalamus and pituitary gland regulate the ovarian cycle and ovulation. GnRH activates the anterior pituitary to produce LH and FSH, which stimulate the production of estrogen and progesterone by the ovaries.[/caption]

When only the one dominant follicle remains in the ovary, it again begins to secrete estrogen. It produces more estrogen than all of the developing follicles did together before the negative feedback occurred. It produces so much estrogen that the normal negative feedback doesn’t occur. Instead, these extremely high concentrations of systemic plasma estrogen trigger a regulatory switch in the anterior pituitary that responds by secreting large amounts of LH and FSH into the bloodstream (see Figure 5). The positive feedback loop by which more estrogen triggers release of more LH and FSH only occurs at this point in the cycle.

It is this large burst of LH (called the LH surge) that leads to ovulation of the dominant follicle. The LH surge induces many changes in the dominant follicle, including stimulating the resumption of meiosis of the primary oocyte to a secondary oocyte. As noted earlier, the polar body that results from unequal cell division simply degrades. The LH surge also triggers proteases (enzymes that cleave proteins) to break down structural proteins in the ovary wall on the surface of the bulging dominant follicle. This degradation of the wall, combined with pressure from the large, fluid-filled antrum, results in the expulsion of the oocyte surrounded by granulosa cells into the peritoneal cavity. This release is ovulation.

In the next section, you will follow the ovulated oocyte as it travels toward the uterus, but there is one more important event that occurs in the ovarian cycle. The surge of LH also stimulates a change in the granulosa and theca cells that remain in the follicle after the oocyte has been ovulated. This change is called luteinization (recall that the full name of LH is luteinizing hormone), and it transforms the collapsed follicle into a new endocrine structure called the corpus luteum, a term meaning “yellowish body” (see Figure 4). Instead of estrogen, the luteinized granulosa and theca cells of the corpus luteum begin to produce large amounts of the sex steroid hormone progesterone, a hormone that is critical for the establishment and maintenance of pregnancy. Progesterone triggers negative feedback at the hypothalamus and pituitary, which keeps GnRH, LH, and FSH secretions low, so no new dominant follicles develop at this time.

The post-ovulatory phase of progesterone secretion is known as the luteal phase of the ovarian cycle. If pregnancy does not occur within 10 to 12 days, the corpus luteum will stop secreting progesterone and degrade into the corpus albicans, a nonfunctional “whitish body” that will disintegrate in the ovary over a period of several months. During this time of reduced progesterone secretion, FSH and LH are once again stimulated, and the follicular phase begins again with a new cohort of early tertiary follicles beginning to grow and secrete estrogen.

The Uterine Tubes

The uterine tubes (also called fallopian tubes or oviducts) serve as the conduit of the oocyte from the ovary to the uterus (Figure 6). Each of the two uterine tubes is close to, but not directly connected to, the ovary and divided into sections. The isthmus is the narrow medial end of each uterine tube that is connected to the uterus. The wide distal infundibulum flares out with slender, finger-like projections called fimbriae. The middle region of the tube, called the ampulla, is where fertilization often occurs. The uterine tubes also have three layers: an outer serosa, a middle smooth muscle layer, and an inner mucosal layer. In addition to its mucus-secreting cells, the inner mucosa contains ciliated cells that beat in the direction of the uterus, producing a current that will be critical to move the oocyte.

Following ovulation, the secondary oocyte surrounded by a few granulosa cells is released into the peritoneal cavity. The nearby uterine tube, either left or right, receives the oocyte. Unlike sperm, oocytes lack flagella, and therefore cannot move on their own. So how do they travel into the uterine tube and toward the uterus? High concentrations of estrogen that occur around the time of ovulation induce contractions of the smooth muscle along the length of the uterine tube. These contractions occur every 4 to 8 seconds, and the result is a coordinated movement that sweeps the surface of the ovary and the pelvic cavity. Current flowing toward the uterus is generated by coordinated beating of the cilia that line the outside and lumen of the length of the uterine tube. These cilia beat more strongly in response to the high estrogen concentrations that occur around the time of ovulation. As a result of these mechanisms, the oocyte–granulosa cell complex is pulled into the interior of the tube. Once inside, the muscular contractions and beating cilia move the oocyte slowly toward the uterus. When fertilization does occur, sperm typically meet the egg while it is still moving through the ampulla.

If the oocyte is successfully fertilized, the resulting zygote will begin to divide into two cells, then four, and so on, as it makes its way through the uterine tube and into the uterus. There, it will implant and continue to grow. If the egg is not fertilized, it will simply degrade—either in the uterine tube or in the uterus, where it may be shed with the next menstrual period. [caption id="" align="aligncenter" width="550"]This diagram shows the uterus and ovaries in the center. To the left is a micrograph showing the ultrastructure of the ovaries and to the right is a micrograph showing the ultrastructure of the uterus. Figure 6. Ovaries, Uterine Tubes, and Uterus. This anterior view shows the relationship of the ovaries, uterine tubes (oviducts), and uterus. Sperm enter through the vagina, and fertilization of an ovulated oocyte usually occurs in the distal uterine tube. From left to right, LM × 400, LM × 20. (Micrographs provided by the Regents of University of Michigan Medical School © 2012)[/caption]

The open-ended structure of the uterine tubes can have significant health consequences if bacteria or other contagions enter through the vagina and move through the uterus, into the tubes, and then into the pelvic cavity. If this is left unchecked, a bacterial infection (sepsis) could quickly become life-threatening. The spread of an infection in this manner is of special concern when unskilled practitioners perform abortions in non-sterile conditions. Sepsis is also associated with sexually transmitted bacterial infections, especially gonorrhea and chlamydia. These increase a woman’s risk for pelvic inflammatory disease (PID), infection of the uterine tubes or other reproductive organs. Even when resolved, PID can leave scar tissue in the tubes, leading to infertility.

The Uterus and Cervix

The uterus is the muscular organ that nourishes and supports the growing embryo (see Figure 6). Its average size is approximately 5 cm wide by 7 cm long (approximately 2 in by 3 in) when a female is not pregnant. It has three sections. The portion of the uterus superior to the opening of the uterine tubes is called the fundus. The middle section of the uterus is called the body of uterus (or corpus). The cervix is the narrow inferior portion of the uterus that projects into the vagina. The cervix produces mucus secretions that become thin and stringy under the influence of high systemic plasma estrogen concentrations, and these secretions can facilitate sperm movement through the reproductive tract.

Several ligaments maintain the position of the uterus within the abdominopelvic cavity. The broad ligament is a fold of peritoneum that serves as a primary support for the uterus, extending laterally from both sides of the uterus and attaching it to the pelvic wall. The round ligament attaches to the uterus near the uterine tubes, and extends to the labia majora. Finally, the uterosacral ligament stabilizes the uterus posteriorly by its connection from the cervix to the pelvic wall.

The wall of the uterus is made up of three layers. The most superficial layer is the serous membrane, or perimetrium, which consists of epithelial tissue that covers the exterior portion of the uterus. The middle layer, or myometrium, is a thick layer of smooth muscle responsible for uterine contractions. Most of the uterus is myometrial tissue, and the muscle fibers run horizontally, vertically, and diagonally, allowing the powerful contractions that occur during labor and the less powerful contractions (or cramps) that help to expel menstrual blood during a woman’s period. Anteriorly directed myometrial contractions also occur near the time of ovulation, and are thought to possibly facilitate the transport of sperm through the female reproductive tract.

The innermost layer of the uterus is called the endometrium. The endometrium contains a connective tissue lining, the lamina propria, which is covered by epithelial tissue that lines the lumen. Structurally, the endometrium consists of two layers: the stratum basalis and the stratum functionalis (the basal and functional layers). The stratum basalis layer is part of the lamina propria and is adjacent to the myometrium; this layer does not shed during menses. In contrast, the thicker stratum functionalis layer contains the glandular portion of the lamina propria and the endothelial tissue that lines the uterine lumen. It is the stratum functionalis that grows and thickens in response to increased levels of estrogen and progesterone. In the luteal phase of the menstrual cycle, special branches off of the uterine artery called spiral arteries supply the thickened stratum functionalis. This inner functional layer provides the proper site of implantation for the fertilized egg, and—should fertilization not occur—it is only the stratum functionalis layer of the endometrium that sheds during menstruation.

Recall that during the follicular phase of the ovarian cycle, the tertiary follicles are growing and secreting estrogen. At the same time, the stratum functionalis of the endometrium is thickening to prepare for a potential implantation. The post-ovulatory increase in progesterone, which characterizes the luteal phase, is key for maintaining a thick stratum functionalis. As long as a functional corpus luteum is present in the ovary, the endometrial lining is prepared for implantation. Indeed, if an embryo implants, signals are sent to the corpus luteum to continue secreting progesterone to maintain the endometrium, and thus maintain the pregnancy. If an embryo does not implant, no signal is sent to the corpus luteum and it degrades, ceasing progesterone production and ending the luteal phase. Without progesterone, the endometrium thins and, under the influence of prostaglandins, the spiral arteries of the endometrium constrict and rupture, preventing oxygenated blood from reaching the endometrial tissue. As a result, endometrial tissue dies and blood, pieces of the endometrial tissue, and white blood cells are shed through the vagina during menstruation, or the menses. The first menses after puberty, called menarche, can occur either before or after the first ovulation.

The Menstrual Cycle

Now that we have discussed the maturation of the cohort of tertiary follicles in the ovary, the build-up and then shedding of the endometrial lining in the uterus, and the function of the uterine tubes and vagina, we can put everything together to talk about the three phases of the menstrual cycle—the series of changes in which the uterine lining is shed, rebuilds, and prepares for implantation.

The timing of the menstrual cycle starts with the first day of menses, referred to as day one of a woman’s period. Cycle length is determined by counting the days between the onset of bleeding in two subsequent cycles. Because the average length of a woman’s menstrual cycle is 28 days, this is the time period used to identify the timing of events in the cycle. However, the length of the menstrual cycle varies among women, and even in the same woman from one cycle to the next, typically from 21 to 32 days.

Just as the hormones produced by the granulosa and theca cells of the ovary “drive” the follicular and luteal phases of the ovarian cycle, they also control the three distinct phases of the menstrual cycle. These are the menses phase, the proliferative phase, and the secretory phase.

Menses Phase

The menses phase of the menstrual cycle is the phase during which the lining is shed; that is, the days that the woman menstruates. Although it averages approximately five days, the menses phase can last from 2 to 7 days, or longer. As shown in Figure 7, the menses phase occurs during the early days of the follicular phase of the ovarian cycle, when progesterone, FSH, and LH levels are low. Recall that progesterone concentrations decline as a result of the degradation of the corpus luteum, marking the end of the luteal phase. This decline in progesterone triggers the shedding of the stratum functionalis of the endometrium. [caption id="" align="aligncenter" width="500"]The top panel of this image shows the stages in the follicular phase and how one follicle is selected at the end of this phase. The middle part of this image shows the ovarian cycle phases and the uterine cycle phases. The bottom panel shows the levels of different hormones as a function of time. Figure 7. Hormone Levels in Ovarian and Menstrual Cycles. The correlation of the hormone levels and their effects on the female reproductive system is shown in this timeline of the ovarian and menstrual cycles. The menstrual cycle begins at day one with the start of menses. Ovulation occurs around day 14 of a 28-day cycle, triggered by the LH surge.[/caption]

Proliferative Phase

Once menstrual flow ceases, the endometrium begins to proliferate again, marking the beginning of the proliferative phase of the menstrual cycle (see Figure 7). It occurs when the granulosa and theca cells of the tertiary follicles begin to produce increased amounts of estrogen. These rising estrogen concentrations stimulate the endometrial lining to rebuild.

Recall that the high estrogen concentrations will eventually lead to a decrease in FSH as a result of negative feedback, resulting in atresia of all but one of the developing tertiary follicles. The switch to positive feedback—which occurs with the elevated estrogen production from the dominant follicle—then stimulates the LH surge that will trigger ovulation. In a typical 28-day menstrual cycle, ovulation occurs on day 14. Ovulation marks the end of the proliferative phase as well as the end of the follicular phase.

Secretory Phase

In addition to prompting the LH surge, high estrogen levels increase the uterine tube contractions that facilitate the pick-up and transfer of the ovulated oocyte. High estrogen levels also slightly decrease the acidity of the vagina, making it more hospitable to sperm. In the ovary, the luteinization of the granulosa cells of the collapsed follicle forms the progesterone-producing corpus luteum, marking the beginning of the luteal phase of the ovarian cycle. In the uterus, progesterone from the corpus luteum begins the secretory phase of the menstrual cycle, in which the endometrial lining prepares for implantation (see Figure 7). Over the next 10 to 12 days, the endometrial glands secrete a fluid rich in glycogen. If fertilization has occurred, this fluid will nourish the ball of cells now developing from the zygote. At the same time, the spiral arteries develop to provide blood to the thickened stratum functionalis.

If no pregnancy occurs within approximately 10 to 12 days, the corpus luteum will degrade into the corpus albicans. Levels of both estrogen and progesterone will fall, and the endometrium will grow thinner. Prostaglandins will be secreted that cause constriction of the spiral arteries, reducing oxygen supply. The endometrial tissue will die, resulting in menses—or the first day of the next cycle.

Disorders of the… Feature

Female Reproductive System Research over many years has confirmed that cervical cancer is most often caused by a sexually transmitted infection with human papillomavirus (HPV). There are over 100 related viruses in the HPV family, and the characteristics of each strain determine the outcome of the infection. In all cases, the virus enters body cells and uses its own genetic material to take over the host cell’s metabolic machinery and produce more virus particles.

HPV infections are common in both men and women. Indeed, a recent study determined that 42.5 percent of females had HPV at the time of testing. These women ranged in age from 14 to 59 years and differed in race, ethnicity, and number of sexual partners. Of note, the prevalence of HPV infection was 53.8 percent among women aged 20 to 24 years, the age group with the highest infection rate.

HPV strains are classified as high or low risk according to their potential to cause cancer. Though most HPV infections do not cause disease, the disruption of normal cellular functions in the low-risk forms of HPV can cause the male or female human host to develop genital warts. Often, the body is able to clear an HPV infection by normal immune responses within 2 years. However, the more serious, high-risk infection by certain types of HPV can result in cancer of the cervix (Figure 8). Infection with either of the cancer-causing variants HPV 16 or HPV 18 has been linked to more than 70 percent of all cervical cancer diagnoses. Although even these high-risk HPV strains can be cleared from the body over time, infections persist in some individuals. If this happens, the HPV infection can influence the cells of the cervix to develop precancerous changes.

Risk factors for cervical cancer include having unprotected sex; having multiple sexual partners; a first sexual experience at a younger age, when the cells of the cervix are not fully mature; failure to receive the HPV vaccine; a compromised immune system; and smoking. The risk of developing cervical cancer is doubled with cigarette smoking. [caption id="" align="aligncenter" width="550"]The left panel shows cell cycle. An arrow from the G2 phase leads to the right panel. The top half of the right panel describes the next steps in the absence of HPV and the bottom half describes the next steps in the presence of HPV. Figure 8. Development of Cervical Cancer. In most cases, cells infected with the HPV virus heal on their own. In some cases, however, the virus continues to spread and becomes an invasive cancer.[/caption]

When the high-risk types of HPV enter a cell, two viral proteins are used to neutralize proteins that the host cells use as checkpoints in the cell cycle. The best studied of these proteins is p53. In a normal cell, p53 detects DNA damage in the cell’s genome and either halts the progression of the cell cycle—allowing time for DNA repair to occur—or initiates apoptosis. Both of these processes prevent the accumulation of mutations in a cell’s genome. High-risk HPV can neutralize p53, keeping the cell in a state in which fast growth is possible and impairing apoptosis, allowing mutations to accumulate in the cellular DNA.

The prevalence of cervical cancer in the United States is very low because of regular screening exams called pap smears. Pap smears sample cells of the cervix, allowing the detection of abnormal cells. If pre-cancerous cells are detected, there are several highly effective techniques that are currently in use to remove them before they pose a danger. However, women in developing countries often do not have access to regular pap smears. As a result, these women account for as many as 80 percent of the cases of cervical cancer worldwide.

In 2006, the first vaccine against the high-risk types of HPV was approved. There are now two HPV vaccines available: Gardasil® and Cervarix®. Whereas these vaccines were initially only targeted for women, because HPV is sexually transmitted, both men and women require vaccination for this approach to achieve its maximum efficacy. A recent study suggests that the HPV vaccine has cut the rates of HPV infection by the four targeted strains at least in half. Unfortunately, the high cost of manufacturing the vaccine is currently limiting access to many women worldwide.

The Breasts

Whereas the breasts are located far from the other female reproductive organs, they are considered accessory organs of the female reproductive system. The function of the breasts is to supply milk to an infant in a process called lactation. The external features of the breast include a nipple surrounded by a pigmented areola (Figure 9), whose coloration may deepen during pregnancy. The areola is typically circular and can vary in size from 25 to 100 mm in diameter. The areolar region is characterized by small, raised areolar glands that secrete lubricating fluid during lactation to protect the nipple from chafing. When a baby nurses, or draws milk from the breast, the entire areolar region is taken into the mouth.

Breast milk is produced by the mammary glands, which are modified sweat glands. The milk itself exits the breast through the nipple via 15 to 20 lactiferous ducts that open on the surface of the nipple. These lactiferous ducts each extend to a lactiferous sinus that connects to a glandular lobe within the breast itself that contains groups of milk-secreting cells in clusters called alveoli (see Figure 9). The clusters can change in size depending on the amount of milk in the alveolar lumen. Once milk is made in the alveoli, stimulated myoepithelial cells that surround the alveoli contract to push the milk to the lactiferous sinuses. From here, the baby can draw milk through the lactiferous ducts by suckling. The lobes themselves are surrounded by fat tissue, which determines the size of the breast; breast size differs between individuals and does not affect the amount of milk produced. Supporting the breasts are multiple bands of connective tissue called suspensory ligaments that connect the breast tissue to the dermis of the overlying skin. [caption id="" align="aligncenter" width="450"]This figure shows the anatomy of the breast. The left panel shows the front view and the right panel shows the side view. The main parts are labeled. Figure 9. Anatomy of the Breast. During lactation, milk moves from the alveoli through the lactiferous ducts to the nipple.[/caption]

During the normal hormonal fluctuations in the menstrual cycle, breast tissue responds to changing levels of estrogen and progesterone, which can lead to swelling and breast tenderness in some individuals, especially during the secretory phase. If pregnancy occurs, the increase in hormones leads to further development of the mammary tissue and enlargement of the breasts.

Hormonal Birth Control

Birth control pills take advantage of the negative feedback system that regulates the ovarian and menstrual cycles to stop ovulation and prevent pregnancy. Typically they work by providing a constant level of both estrogen and progesterone, which negatively feeds back onto the hypothalamus and pituitary, thus preventing the release of FSH and LH. Without FSH, the follicles do not mature, and without the LH surge, ovulation does not occur. Although the estrogen in birth control pills does stimulate some thickening of the endometrial wall, it is reduced compared with a normal cycle and is less likely to support implantation.

Some birth control pills contain 21 active pills containing hormones, and 7 inactive pills (placebos). The decline in hormones during the week that the woman takes the placebo pills triggers menses, although it is typically lighter than a normal menstrual flow because of the reduced endometrial thickening. Newer types of birth control pills have been developed that deliver low-dose estrogens and progesterone for the entire cycle (these are meant to be taken 365 days a year), and menses never occurs. While some women prefer to have the proof of a lack of pregnancy that a monthly period provides, menstruation every 28 days is not required for health reasons, and there are no reported adverse effects of not having a menstrual period in an otherwise healthy individual.

Because birth control pills function by providing constant estrogen and progesterone levels and disrupting negative feedback, skipping even just one or two pills at certain points of the cycle (or even being several hours late taking the pill) can lead to an increase in FSH and LH and result in ovulation. It is important, therefore, that the woman follow the directions on the birth control pill package to successfully prevent pregnancy.

Aging and the… Feature

Female Reproductive System Female fertility (the ability to conceive) peaks when women are in their twenties, and is slowly reduced until a women reaches 35 years of age. After that time, fertility declines more rapidly, until it ends completely at the end of menopause. Menopause is the cessation of the menstrual cycle that occurs as a result of the loss of ovarian follicles and the hormones that they produce. A woman is considered to have completed menopause if she has not menstruated in a full year. After that point, she is considered postmenopausal. The average age for this change is consistent worldwide at between 50 and 52 years of age, but it can normally occur in a woman’s forties, or later in her fifties. Poor health, including smoking, can lead to earlier loss of fertility and earlier menopause.

As a woman reaches the age of menopause, depletion of the number of viable follicles in the ovaries due to atresia affects the hormonal regulation of the menstrual cycle. During the years leading up to menopause, there is a decrease in the levels of the hormone inhibin, which normally participates in a negative feedback loop to the pituitary to control the production of FSH. The menopausal decrease in inhibin leads to an increase in FSH. The presence of FSH stimulates more follicles to grow and secrete estrogen. Because small, secondary follicles also respond to increases in FSH levels, larger numbers of follicles are stimulated to grow; however, most undergo atresia and die. Eventually, this process leads to the depletion of all follicles in the ovaries, and the production of estrogen falls off dramatically. It is primarily the lack of estrogens that leads to the symptoms of menopause.

The earliest changes occur during the menopausal transition, often referred to as peri-menopause, when a women’s cycle becomes irregular but does not stop entirely. Although the levels of estrogen are still nearly the same as before the transition, the level of progesterone produced by the corpus luteum is reduced. This decline in progesterone can lead to abnormal growth, or hyperplasia, of the endometrium. This condition is a concern because it increases the risk of developing endometrial cancer. Two harmless conditions that can develop during the transition are uterine fibroids, which are benign masses of cells, and irregular bleeding. As estrogen levels change, other symptoms that occur are hot flashes and night sweats, trouble sleeping, vaginal dryness, mood swings, difficulty focusing, and thinning of hair on the head along with the growth of more hair on the face. Depending on the individual, these symptoms can be entirely absent, moderate, or severe.

After menopause, lower amounts of estrogens can lead to other changes. Cardiovascular disease becomes as prevalent in women as in men, possibly because estrogens reduce the amount of cholesterol in the blood vessels. When estrogen is lacking, many women find that they suddenly have problems with high cholesterol and the cardiovascular issues that accompany it. Osteoporosis is another problem because bone density decreases rapidly in the first years after menopause. The reduction in bone density leads to a higher incidence of fractures.

Hormone therapy (HT), which employs medication (synthetic estrogens and progestins) to increase estrogen and progestin levels, can alleviate some of the symptoms of menopause. In 2002, the Women’s Health Initiative began a study to observe women for the long-term outcomes of hormone replacement therapy over 8.5 years. However, the study was prematurely terminated after 5.2 years because of evidence of a higher than normal risk of breast cancer in patients taking estrogen-only HT. The potential positive effects on cardiovascular disease were also not realized in the estrogen-only patients. The results of other hormone replacement studies over the last 50 years, including a 2012 study that followed over 1,000 menopausal women for 10 years, have shown cardiovascular benefits from estrogen and no increased risk for cancer. Some researchers believe that the age group tested in the 2002 trial may have been too old to benefit from the therapy, thus skewing the results. In the meantime, intense debate and study of the benefits and risks of replacement therapy is ongoing. Current guidelines approve HT for the reduction of hot flashes or flushes, but this treatment is generally only considered when women first start showing signs of menopausal changes, is used in the lowest dose possible for the shortest time possible (5 years or less), and it is suggested that women on HT have regular pelvic and breast exams.

]]> 2783 2763 2 0 28.1 Fertilization https://pressbooks.bccampus.ca/apdouglas2017/chapter/28-1-fertilization/ Mon, 17 Jul 2017 19:44:56 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2789

Learning Objectives

By the end of this section, you will be able to:
  • Describe the obstacles that sperm must overcome to reach an oocyte
  • Explain capacitation and its importance in fertilization
  • Summarize the events that occur as a sperm fertilizes an oocyte

Fertilization occurs when a sperm and an oocyte (egg) combine and their nuclei fuse. Because each of these reproductive cells is a haploid cell containing half of the genetic material needed to form a human being, their combination forms a diploid cell. This new single cell, called a zygote, contains all of the genetic material needed to form a human—half from the mother and half from the father.

Transit of Sperm

Fertilization is a numbers game. During ejaculation, hundreds of millions of sperm (spermatozoa) are released into the vagina. Almost immediately, millions of these sperm are overcome by the acidity of the vagina (approximately pH 3.8), and millions more may be blocked from entering the uterus by thick cervical mucus. Of those that do enter, thousands are destroyed by phagocytic uterine leukocytes. Thus, the race into the uterine tubes, which is the most typical site for sperm to encounter the oocyte, is reduced to a few thousand contenders. Their journey—thought to be facilitated by uterine contractions—usually takes from 30 minutes to 2 hours. If the sperm do not encounter an oocyte immediately, they can survive in the uterine tubes for another 3–5 days. Thus, fertilization can still occur if intercourse takes place a few days before ovulation. In comparison, an oocyte can survive independently for only approximately 24 hours following ovulation. Intercourse more than a day after ovulation will therefore usually not result in fertilization.

During the journey, fluids in the female reproductive tract prepare the sperm for fertilization through a process called capacitation, or priming. The fluids improve the motility of the spermatozoa. They also deplete cholesterol molecules embedded in the membrane of the head of the sperm, thinning the membrane in such a way that will help facilitate the release of the lysosomal (digestive) enzymes needed for the sperm to penetrate the oocyte’s exterior once contact is made. Sperm must undergo the process of capacitation in order to have the “capacity” to fertilize an oocyte. If they reach the oocyte before capacitation is complete, they will be unable to penetrate the oocyte’s thick outer layer of cells.

Contact Between Sperm and Oocyte

Upon ovulation, the oocyte released by the ovary is swept into—and along—the uterine tube. Fertilization must occur in the distal uterine tube because an unfertilized oocyte cannot survive the 72-hour journey to the uterus. As you will recall from your study of the oogenesis, this oocyte (specifically a secondary oocyte) is surrounded by two protective layers. The corona radiata is an outer layer of follicular (granulosa) cells that form around a developing oocyte in the ovary and remain with it upon ovulation. The underlying zona pellucida (pellucid = “transparent”) is a transparent, but thick, glycoprotein membrane that surrounds the cell’s plasma membrane.

As it is swept along the distal uterine tube, the oocyte encounters the surviving capacitated sperm, which stream toward it in response to chemical attractants released by the cells of the corona radiata. To reach the oocyte itself, the sperm must penetrate the two protective layers. The sperm first burrow through the cells of the corona radiata. Then, upon contact with the zona pellucida, the sperm bind to receptors in the zona pellucida. This initiates a process called the acrosomal reaction in which the enzyme-filled “cap” of the sperm, called the acrosome, releases its stored digestive enzymes. These enzymes clear a path through the zona pellucida that allows sperm to reach the oocyte. Finally, a single sperm makes contact with sperm-binding receptors on the oocyte’s plasma membrane (Figure 1). The plasma membrane of that sperm then fuses with the oocyte’s plasma membrane, and the head and mid-piece of the “winning” sperm enter the oocyte interior.

How do sperm penetrate the corona radiata? Some sperm undergo a spontaneous acrosomal reaction, which is an acrosomal reaction not triggered by contact with the zona pellucida. The digestive enzymes released by this reaction digest the extracellular matrix of the corona radiata. As you can see, the first sperm to reach the oocyte is never the one to fertilize it. Rather, hundreds of sperm cells must undergo the acrosomal reaction, each helping to degrade the corona radiata and zona pellucida until a path is created to allow one sperm to contact and fuse with the plasma membrane of the oocyte. If you consider the loss of millions of sperm between entry into the vagina and degradation of the zona pellucida, you can understand why a low sperm count can cause male infertility.

[caption id="" align="aligncenter" width="520"]This figure shows the process of sperm fertilizing an egg. There are many sperm trying to attach to the egg. Figure 1. Sperm and the Process of Fertilization. Before fertilization, hundreds of capacitated sperm must break through the surrounding corona radiata and zona pellucida so that one can contact and fuse with the oocyte plasma membrane.[/caption]

When the first sperm fuses with the oocyte, the oocyte deploys two mechanisms to prevent polyspermy, which is penetration by more than one sperm. This is critical because if more than one sperm were to fertilize the oocyte, the resulting zygote would be a triploid organism with three sets of chromosomes. This is incompatible with life.

The first mechanism is the fast block, which involves a near instantaneous change in sodium ion permeability upon binding of the first sperm, depolarizing the oocyte plasma membrane and preventing the fusion of additional sperm cells. The fast block sets in almost immediately and lasts for about a minute, during which time an influx of calcium ions following sperm penetration triggers the second mechanism, the slow block. In this process, referred to as the cortical reaction, cortical granules sitting immediately below the oocyte plasma membrane fuse with the membrane and release zonal inhibiting proteins and mucopolysaccharides into the space between the plasma membrane and the zona pellucida. Zonal inhibiting proteins cause the release of any other attached sperm and destroy the oocyte’s sperm receptors, thus preventing any more sperm from binding. The mucopolysaccharides then coat the nascent zygote in an impenetrable barrier that, together with hardened zona pellucida, is called a fertilization membrane.

The Zygote

Recall that at the point of fertilization, the oocyte has not yet completed meiosis; all secondary oocytes remain arrested in metaphase of meiosis II until fertilization. Only upon fertilization does the oocyte complete meiosis. The unneeded complement of genetic material that results is stored in a second polar body that is eventually ejected. At this moment, the oocyte has become an ovum, the female haploid gamete. The two haploid nuclei derived from the sperm and oocyte and contained within the egg are referred to as pronuclei. They decondense, expand, and replicate their DNA in preparation for mitosis. The pronuclei then migrate toward each other, their nuclear envelopes disintegrate, and the male- and female-derived genetic material intermingles. This step completes the process of fertilization and results in a single-celled diploid zygote with all the genetic instructions it needs to develop into a human.

Most of the time, a woman releases a single egg during an ovulation cycle. However, in approximately 1 percent of ovulation cycles, two eggs are released and both are fertilized. Two zygotes form, implant, and develop, resulting in the birth of dizygotic (or fraternal) twins. Because dizygotic twins develop from two eggs fertilized by two sperm, they are no more identical than siblings born at different times.

Much less commonly, a zygote can divide into two separate offspring during early development. This results in the birth of monozygotic (or identical) twins. Although the zygote can split as early as the two-cell stage, splitting occurs most commonly during the early blastocyst stage, with roughly 70–100 cells present. These two scenarios are distinct from each other, in that the twin embryos that separated at the two-cell stage will have individual placentas, whereas twin embryos that form from separation at the blastocyst stage will share a placenta and a chorionic cavity.
Everyday Connections

In Vitro Fertilization IVF, which stands for in vitro fertilization, is an assisted reproductive technology. In vitro, which in Latin translates to “in glass,” refers to a procedure that takes place outside of the body. There are many different indications for IVF. For example, a woman may produce normal eggs, but the eggs cannot reach the uterus because the uterine tubes are blocked or otherwise compromised. A man may have a low sperm count, low sperm motility, sperm with an unusually high percentage of morphological abnormalities, or sperm that are incapable of penetrating the zona pellucida of an egg.

A typical IVF procedure begins with egg collection. A normal ovulation cycle produces only one oocyte, but the number can be boosted significantly (to 10–20 oocytes) by administering a short course of gonadotropins. The course begins with follicle-stimulating hormone (FSH) analogs, which support the development of multiple follicles, and ends with a luteinizing hormone (LH) analog that triggers ovulation. Right before the ova would be released from the ovary, they are harvested using ultrasound-guided oocyte retrieval. In this procedure, ultrasound allows a physician to visualize mature follicles. The ova are aspirated (sucked out) using a syringe.

In parallel, sperm are obtained from the male partner or from a sperm bank. The sperm are prepared by washing to remove seminal fluid because seminal fluid contains a peptide, FPP (or, fertilization promoting peptide), that—in high concentrations—prevents capacitation of the sperm. The sperm sample is also concentrated, to increase the sperm count per milliliter.

Next, the eggs and sperm are mixed in a petri dish. The ideal ratio is 75,000 sperm to one egg. If there are severe problems with the sperm—for example, the count is exceedingly low, or the sperm are completely nonmotile, or incapable of binding to or penetrating the zona pellucida—a sperm can be injected into an egg. This is called intracytoplasmic sperm injection (ICSI).

The embryos are then incubated until they either reach the eight-cell stage or the blastocyst stage. In the United States, fertilized eggs are typically cultured to the blastocyst stage because this results in a higher pregnancy rate. Finally, the embryos are transferred to a woman’s uterus using a plastic catheter (tube). Figure 2 illustrates the steps involved in IVF.

[caption id="" align="aligncenter" width="420"]This multi-part figure shows the different steps in in vitro fertilization. The top panel shows how the oocytes and the sperm are collected and prepared. The next panel shows the sperm and oocytes being mixed in a petri dish. The panel below that shows the fertilized zygote being prepared for implantation. The last panel shows the fertilized zygote being implanted into the uterus. Figure 2. IVF. In vitro fertilization involves egg collection from the ovaries, fertilization in a petri dish, and the transfer of embryos into the uterus.[/caption]

IVF is a relatively new and still evolving technology, and until recently it was necessary to transfer multiple embryos to achieve a good chance of a pregnancy. Today, however, transferred embryos are much more likely to implant successfully, so countries that regulate the IVF industry cap the number of embryos that can be transferred per cycle at two. This reduces the risk of multiple-birth pregnancies.

The rate of success for IVF is correlated with a woman’s age. More than 40 percent of women under 35 succeed in giving birth following IVF, but the rate drops to a little over 10 percent in women over 40.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Go to this site to view resources covering various aspects of fertilization, including movies and animations showing sperm structure and motility, ovulation, and fertilization.[/caption]
[caption id="attachment_3031" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about fertilization![/caption]
]]> 2789 2785 1 0 28.2 Embryonic Development https://pressbooks.bccampus.ca/apdouglas2017/chapter/28-2-embryonic-development/ Mon, 17 Jul 2017 19:45:48 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2803

Learning Objectives

By the end of this section, you will be able to:
  • Distinguish the stages of embryonic development that occur before implantation
  • Describe the process of implantation
  • List and describe four embryonic membranes
  • Explain gastrulation
  • Describe how the placenta is formed and identify its functions
  • Explain how an embryo transforms from a flat disc of cells into a three-dimensional shape resembling a human
  • Summarize the process of organogenesis

Throughout this chapter, we will express embryonic and fetal ages in terms of weeks from fertilization, commonly called conception. The period of time required for full development of a fetus in utero is referred to as gestation (gestare = “to carry” or “to bear”). It can be subdivided into distinct gestational periods. The first 2 weeks of prenatal development are referred to as the pre-embryonic stage. A developing human is referred to as an embryo during weeks 3–8, and a fetus from the ninth week of gestation until birth. In this section, we’ll cover the pre-embryonic and embryonic stages of development, which are characterized by cell division, migration, and differentiation. By the end of the embryonic period, all of the organ systems are structured in rudimentary form, although the organs themselves are either nonfunctional or only semi-functional.

Pre-implantation Embryonic Development

Following fertilization, the zygote and its associated membranes, together referred to as the conceptus, continue to be projected toward the uterus by peristalsis and beating cilia. During its journey to the uterus, the zygote undergoes five or six rapid mitotic cell divisions. Although each cleavage results in more cells, it does not increase the total volume of the conceptus (Figure 1). Each daughter cell produced by cleavage is called a blastomere (blastos = “germ,” in the sense of a seed or sprout).

Approximately 3 days after fertilization, a 16-cell conceptus reaches the uterus. The cells that had been loosely grouped are now compacted and look more like a solid mass. The name given to this structure is the morula (morula = “little mulberry”). Once inside the uterus, the conceptus floats freely for several more days. It continues to divide, creating a ball of approximately 100 cells, and consuming nutritive endometrial secretions called uterine milk while the uterine lining thickens. The ball of now tightly bound cells starts to secrete fluid and organize themselves around a fluid-filled cavity, the blastocoel. At this developmental stage, the conceptus is referred to as a blastocyst. Within this structure, a group of cells forms into an inner cell mass, which is fated to become the embryo. The cells that form the outer shell are called trophoblasts (trophe = “to feed” or “to nourish”). These cells will develop into the chorionic sac and the fetal portion of the placenta (the organ of nutrient, waste, and gas exchange between mother and the developing offspring).

The inner mass of embryonic cells is totipotent during this stage, meaning that each cell has the potential to differentiate into any cell type in the human body. Totipotency lasts for only a few days before the cells’ fates are set as being the precursors to a specific lineage of cells.

[caption id="" align="aligncenter" width="380"]This figure shows the different stages of cell divisions taking place before the embryo is formed. The top panel shows the cell divisions occurring in the uterine tube and the bottom panel shows the cell divisions occurring in the uterus. Figure 1. Pre-Embryonic Cleavages. Pre-embryonic cleavages make use of the abundant cytoplasm of the conceptus as the cells rapidly divide without changing the total volume.[/caption]
As the blastocyst forms, the trophoblast excretes enzymes that begin to degrade the zona pellucida. In a process called “hatching,” the conceptus breaks free of the zona pellucida in preparation for implantation.
[caption id="" align="aligncenter" width="120"]QR Code representing a URL View this time-lapse movie of a conceptus starting at day 3.[/caption]

Implantation

At the end of the first week, the blastocyst comes in contact with the uterine wall and adheres to it, embedding itself in the uterine lining via the trophoblast cells. Thus begins the process of implantation, which signals the end of the pre-embryonic stage of development (Figure 2). Implantation can be accompanied by minor bleeding. The blastocyst typically implants in the fundus of the uterus or on the posterior wall. However, if the endometrium is not fully developed and ready to receive the blastocyst, the blastocyst will detach and find a better spot. A significant percentage (50–75 percent) of blastocysts fail to implant; when this occurs, the blastocyst is shed with the endometrium during menses. The high rate of implantation failure is one reason why pregnancy typically requires several ovulation cycles to achieve.

[caption id="" align="aligncenter" width="480"]This figure shows the different stages in pre-embryonic development. A diagram of the uterus is shown and from this image, eight callouts show the different stages of development. Figure 2. Pre-Embryonic Development. Ovulation, fertilization, pre-embryonic development, and implantation occur at specific locations within the female reproductive system in a time span of approximately 1 week.[/caption]

When implantation succeeds and the blastocyst adheres to the endometrium, the superficial cells of the trophoblast fuse with each other, forming the syncytiotrophoblast, a multinucleated body that digests endometrial cells to firmly secure the blastocyst to the uterine wall. In response, the uterine mucosa rebuilds itself and envelops the blastocyst (Figure 3). The trophoblast secretes human chorionic gonadotropin (hCG), a hormone that directs the corpus luteum to survive, enlarge, and continue producing progesterone and estrogen to suppress menses. These functions of hCG are necessary for creating an environment suitable for the developing embryo. As a result of this increased production, hCG accumulates in the maternal bloodstream and is excreted in the urine. Implantation is complete by the middle of the second week. Just a few days after implantation, the trophoblast has secreted enough hCG for an at-home urine pregnancy test to give a positive result.

[caption id="" align="aligncenter" width="520"]This figure shows the different steps during implantation. The top panel shows how the blastocyst burrows into the endometrium. The middle panel shows the blastocyst completely surrounded by the endometrium. The bottom panel shows the implanted embryo growing in the uterus. Figure 3. Implantation. During implantation, the trophoblast cells of the blastocyst adhere to the endometrium and digest endometrial cells until it is attached securely.[/caption]

Most of the time an embryo implants within the body of the uterus in a location that can support growth and development. However, in one to two percent of cases, the embryo implants either outside the uterus (an ectopic pregnancy) or in a region of uterus that can create complications for the pregnancy. If the embryo implants in the inferior portion of the uterus, the placenta can potentially grow over the opening of the cervix, a condition call placenta previa.

Disorders of the…

Development of the Embryo In the vast majority of ectopic pregnancies, the embryo does not complete its journey to the uterus and implants in the uterine tube, referred to as a tubal pregnancy. However, there are also ovarian ectopic pregnancies (in which the egg never left the ovary) and abdominal ectopic pregnancies (in which an egg was “lost” to the abdominal cavity during the transfer from ovary to uterine tube, or in which an embryo from a tubal pregnancy re-implanted in the abdomen). Once in the abdominal cavity, an embryo can implant into any well-vascularized structure—the rectouterine cavity (Douglas’ pouch), the mesentery of the intestines, and the greater omentum are some common sites.

Tubal pregnancies can be caused by scar tissue within the tube following a sexually transmitted bacterial infection. The scar tissue impedes the progress of the embryo into the uterus—in some cases “snagging” the embryo and, in other cases, blocking the tube completely. Approximately one half of tubal pregnancies resolve spontaneously. Implantation in a uterine tube causes bleeding, which appears to stimulate smooth muscle contractions and expulsion of the embryo. In the remaining cases, medical or surgical intervention is necessary. If an ectopic pregnancy is detected early, the embryo’s development can be arrested by the administration of the cytotoxic drug methotrexate, which inhibits the metabolism of folic acid. If diagnosis is late and the uterine tube is already ruptured, surgical repair is essential.

Even if the embryo has successfully found its way to the uterus, it does not always implant in an optimal location (the fundus or the posterior wall of the uterus). Placenta previa can result if an embryo implants close to the internal os of the uterus (the internal opening of the cervix). As the fetus grows, the placenta can partially or completely cover the opening of the cervix (Figure 4). Although it occurs in only 0.5 percent of pregnancies, placenta previa is the leading cause of antepartum hemorrhage (profuse vaginal bleeding after week 24 of pregnancy but prior to childbirth).

[caption id="" align="aligncenter" width="380"]The left panel of this image shows the normal location of the placenta and the right panel shows the location of the placenta in placenta previa. Figure 4. Placenta Previa. An embryo that implants too close to the opening of the cervix can lead to placenta previa, a condition in which the placenta partially or completely covers the cervix.[/caption]

Embryonic Membranes

During the second week of development, with the embryo implanted in the uterus, cells within the blastocyst start to organize into layers. Some grow to form the extra-embryonic membranes needed to support and protect the growing embryo: the amnion, the yolk sac, the allantois, and the chorion.

At the beginning of the second week, the cells of the inner cell mass form into a two-layered disc of embryonic cells, and a space—the amniotic cavity—opens up between it and the trophoblast (Figure 5). Cells from the upper layer of the disc (the epiblast) extend around the amniotic cavity, creating a membranous sac that forms into the amnion by the end of the second week. The amnion fills with amniotic fluid and eventually grows to surround the embryo. Early in development, amniotic fluid consists almost entirely of a filtrate of maternal plasma, but as the kidneys of the fetus begin to function at approximately the eighth week, they add urine to the volume of amniotic fluid. Floating within the amniotic fluid, the embryo—and later, the fetus—is protected from trauma and rapid temperature changes. It can move freely within the fluid and can prepare for swallowing and breathing out of the uterus.

[caption id="" align="aligncenter" width="380"]This image shows the development of the amniotic cavity and the location of the embryonic disc. Figure 5. Development of the Embryonic Disc. Formation of the embryonic disc leaves spaces on either side that develop into the amniotic cavity and the yolk sac.[/caption]

On the ventral side of the embryonic disc, opposite the amnion, cells in the lower layer of the embryonic disk (the hypoblast) extend into the blastocyst cavity and form a yolk sac. The yolk sac supplies some nutrients absorbed from the trophoblast and also provides primitive blood circulation to the developing embryo for the second and third week of development. When the placenta takes over nourishing the embryo at approximately week 4, the yolk sac has been greatly reduced in size and its main function is to serve as the source of blood cells and germ cells (cells that will give rise to gametes). During week 3, a finger-like outpocketing of the yolk sac develops into the allantois, a primitive excretory duct of the embryo that will become part of the urinary bladder. Together, the stalks of the yolk sac and allantois establish the outer structure of the umbilical cord.

The last of the extra-embryonic membranes is the chorion, which is the one membrane that surrounds all others. The development of the chorion will be discussed in more detail shortly, as it relates to the growth and development of the placenta.

Embryogenesis

As the third week of development begins, the two-layered disc of cells becomes a three-layered disc through the process of gastrulation, during which the cells transition from totipotency to multipotency. The embryo, which takes the shape of an oval-shaped disc, forms an indentation called the primitive streak along the dorsal surface of the epiblast. A node at the caudal or “tail” end of the primitive streak emits growth factors that direct cells to multiply and migrate. Cells migrate toward and through the primitive streak and then move laterally to create two new layers of cells. The first layer is the endoderm, a sheet of cells that displaces the hypoblast and lies adjacent to the yolk sac. The second layer of cells fills in as the middle layer, or mesoderm. The cells of the epiblast that remain (not having migrated through the primitive streak) become the ectoderm (Figure 6).

[caption id="" align="aligncenter" width="480"]This image shows the different germ layers. The top panel shows the epiblast and trophoblast cells in the early stages of development. The bottom panel shows the three germ layers: the endoderm, ectoderm, and mesoderm. All the other major parts are also labeled. Figure 6. Germ Layers. Formation of the three primary germ layers occurs during the first 2 weeks of development. The embryo at this stage is only a few millimeters in length.[/caption]

Each of these germ layers will develop into specific structures in the embryo. Whereas the ectoderm and endoderm form tightly connected epithelial sheets, the mesodermal cells are less organized and exist as a loosely connected cell community. The ectoderm gives rise to cell lineages that differentiate to become the central and peripheral nervous systems, sensory organs, epidermis, hair, and nails. Mesodermal cells ultimately become the skeleton, muscles, connective tissue, heart, blood vessels, and kidneys. The endoderm goes on to form the epithelial lining of the gastrointestinal tract, liver, and pancreas, as well as the lungs (Figure 7).

[caption id="" align="aligncenter" width="320"]This image shows the structure of the embryo in the third week of development. Under the image, three callouts list the different organ systems into which each germ layer develops. Figure 7. Fates of Germ Layers in Embryo. Following gastrulation of the embryo in the third week, embryonic cells of the ectoderm, mesoderm, and endoderm begin to migrate and differentiate into the cell lineages that will give rise to mature organs and organ systems in the infant.[/caption]

Development of the Placenta

During the first several weeks of development, the cells of the endometrium—referred to as decidual cells—nourish the nascent embryo. During prenatal weeks 4–12, the developing placenta gradually takes over the role of feeding the embryo, and the decidual cells are no longer needed. The mature placenta is composed of tissues derived from the embryo, as well as maternal tissues of the endometrium. The placenta connects to the conceptus via the umbilical cord, which carries deoxygenated blood and wastes from the fetus through two umbilical arteries; nutrients and oxygen are carried from the mother to the fetus through the single umbilical vein. The umbilical cord is surrounded by the amnion, and the spaces within the cord around the blood vessels are filled with Wharton’s jelly, a mucous connective tissue.

The maternal portion of the placenta develops from the deepest layer of the endometrium, the decidua basalis. To form the embryonic portion of the placenta, the syncytiotrophoblast and the underlying cells of the trophoblast (cytotrophoblast cells) begin to proliferate along with a layer of extraembryonic mesoderm cells. These form the chorionic membrane, which envelops the entire conceptus as the chorion. The chorionic membrane forms finger-like structures called chorionic villi that burrow into the endometrium like tree roots, making up the fetal portion of the placenta. The cytotrophoblast cells perforate the chorionic villi, burrow farther into the endometrium, and remodel maternal blood vessels to augment maternal blood flow surrounding the villi. Meanwhile, fetal mesenchymal cells derived from the mesoderm fill the villi and differentiate into blood vessels, including the three umbilical blood vessels that connect the embryo to the developing placenta (Figure 8).

[caption id="" align="aligncenter" width="520"]This figure shows the location and structure of the placenta. The left panel shows a fetus in the womb. The right panel shows a magnified view of a small region including the placenta and the blood vessels. Figure 8. Cross-Section of the Placenta. In the placenta, maternal and fetal blood components are conducted through the surface of the chorionic villi, but maternal and fetal bloodstreams never mix directly.[/caption]

The placenta develops throughout the embryonic period and during the first several weeks of the fetal period; placentation is complete by weeks 14–16. As a fully developed organ, the placenta provides nutrition and excretion, respiration, and endocrine function (Table 1 and Figure 9). It receives blood from the fetus through the umbilical arteries. Capillaries in the chorionic villi filter fetal wastes out of the blood and return clean, oxygenated blood to the fetus through the umbilical vein. Nutrients and oxygen are transferred from maternal blood surrounding the villi through the capillaries and into the fetal bloodstream. Some substances move across the placenta by simple diffusion. Oxygen, carbon dioxide, and any other lipid-soluble substances take this route. Other substances move across by facilitated diffusion. This includes water-soluble glucose. The fetus has a high demand for amino acids and iron, and those substances are moved across the placenta by active transport.

Maternal and fetal blood does not commingle because blood cells cannot move across the placenta. This separation prevents the mother’s cytotoxic T cells from reaching and subsequently destroying the fetus, which bears “non-self” antigens. Further, it ensures the fetal red blood cells do not enter the mother’s circulation and trigger antibody development (if they carry “non-self” antigens)—at least until the final stages of pregnancy or birth. This is the reason that, even in the absence of preventive treatment, an Rh− mother doesn’t develop antibodies that could cause hemolytic disease in her first Rh+ fetus.

Although blood cells are not exchanged, the chorionic villi provide ample surface area for the two-way exchange of substances between maternal and fetal blood. The rate of exchange increases throughout gestation as the villi become thinner and increasingly branched. The placenta is permeable to lipid-soluble fetotoxic substances: alcohol, nicotine, barbiturates, antibiotics, certain pathogens, and many other substances that can be dangerous or fatal to the developing embryo or fetus. For these reasons, pregnant women should avoid fetotoxic substances. Alcohol consumption by pregnant women, for example, can result in a range of abnormalities referred to as fetal alcohol spectrum disorders (FASD). These include organ and facial malformations, as well as cognitive and behavioral disorders.

Functions of the Placenta (Table 1)
Nutrition and digestion Respiration Endocrine function
  • Mediates diffusion of maternal glucose, amino acids, fatty acids, vitamins, and minerals
  • Stores nutrients during early pregnancy to accommodate increased fetal demand later in pregnancy
  • Excretes and filters fetal nitrogenous wastes into maternal blood
  • Mediates maternal-to-fetal oxygen transport and fetal-to-maternal carbon dioxide transport
  • Secretes several hormones, including hCG, estrogens, and progesterone, to maintain the pregnancy and stimulate maternal and fetal development
  • Mediates the transmission of maternal hormones into fetal blood and vice versa
[caption id="" align="aligncenter" width="320"]This is a photo of a placenta and umbilical cord post-expulsion. Figure 9. Placenta. This post-expulsion placenta and umbilical cord (white) are viewed from the fetal side.[/caption]

Organogenesis

Following gastrulation, rudiments of the central nervous system develop from the ectoderm in the process of neurulation (Figure 10). Specialized neuroectodermal tissues along the length of the embryo thicken into the neural plate. During the fourth week, tissues on either side of the plate fold upward into a neural fold. The two folds converge to form the neural tube. The tube lies atop a rod-shaped, mesoderm-derived notochord, which eventually becomes the nucleus pulposus of intervertebral discs. Block-like structures called somites form on either side of the tube, eventually differentiating into the axial skeleton, skeletal muscle, and dermis. During the fourth and fifth weeks, the anterior neural tube dilates and subdivides to form vesicles that will become the brain structures.

Folate, one of the B vitamins, is important to the healthy development of the neural tube. A deficiency of maternal folate in the first weeks of pregnancy can result in neural tube defects, including spina bifida—a birth defect in which spinal tissue protrudes through the newborn’s vertebral column, which has failed to completely close. A more severe neural tube defect is anencephaly, a partial or complete absence of brain tissue.

[caption id="" align="aligncenter" width="500"]This multi-part image shows the formation of the neural tube and the notochord. The top panel shows the ectoderm and mesoderm. The second panel shows the neural plate starting to fold over and the third panel shows the closed neural plate forming the neural tube. The fourth panel shows the mesoderm-derived notochord under the neural tube. Figure 10. Neurulation. The embryonic process of neurulation establishes the rudiments of the future central nervous system and skeleton.[/caption]

The embryo, which begins as a flat sheet of cells, begins to acquire a cylindrical shape through the process of embryonic folding (Figure 11). The embryo folds laterally and again at either end, forming a C-shape with distinct head and tail ends. The embryo envelops a portion of the yolk sac, which protrudes with the umbilical cord from what will become the abdomen. The folding essentially creates a tube, called the primitive gut, that is lined by the endoderm. The amniotic sac, which was sitting on top of the flat embryo, envelops the embryo as it folds.

[caption id="" align="aligncenter" width="520"]This multipart image shows the folding of the embryo. Each of the six panels shows a progression of steps in which the embryo folds on itself. Figure 11. Embryonic Folding. Embryonic folding converts a flat sheet of cells into a hollow, tube-like structure.[/caption]

Within the first 8 weeks of gestation, a developing embryo establishes the rudimentary structures of all of its organs and tissues from the ectoderm, mesoderm, and endoderm. This process is called organogenesis.

Like the central nervous system, the heart also begins its development in the embryo as a tube-like structure, connected via capillaries to the chorionic villi. Cells of the primitive tube-shaped heart are capable of electrical conduction and contraction. The heart begins beating in the beginning of the fourth week, although it does not actually pump embryonic blood until a week later, when the oversized liver has begun producing red blood cells. (This is a temporary responsibility of the embryonic liver that the bone marrow will assume during fetal development.) During weeks 4–5, the eye pits form, limb buds become apparent, and the rudiments of the pulmonary system are formed.

During the sixth week, uncontrolled fetal limb movements begin to occur. The gastrointestinal system develops too rapidly for the embryonic abdomen to accommodate it, and the intestines temporarily loop into the umbilical cord. Paddle-shaped hands and feet develop fingers and toes by the process of apoptosis (programmed cell death), which causes the tissues between the fingers to disintegrate. By week 7, the facial structure is more complex and includes nostrils, outer ears, and lenses (Figure 12). By the eighth week, the head is nearly as large as the rest of the embryo’s body, and all major brain structures are in place. The external genitalia are apparent, but at this point, male and female embryos are indistinguishable. Bone begins to replace cartilage in the embryonic skeleton through the process of ossification. By the end of the embryonic period, the embryo is approximately 3 cm (1.2 in) from crown to rump and weighs approximately 8 g (0.25 oz).

[caption id="" align="aligncenter" width="320"]A photograph of an embryo derived from an ectopic pregnancy is shown. Figure 12. Embryo at 7 Weeks. An embryo at the end of 7 weeks of development is only 10 mm in length, but its developing eyes, limb buds, and tail are already visible. (This embryo was derived from an ectopic pregnancy.) (credit: Ed Uthman)[/caption]
[caption id="attachment_3033" align="aligncenter" width="150"] Watch this CrashCourse video to learn about the stages of embryonic development![/caption]
]]> 2803 2785 2 0 1.1 Overview of Anatomy and Physiology https://pressbooks.bccampus.ca/apdouglas2017/chapter/1-1-overview-of-anatomy-and-physiology/ Wed, 02 Aug 2017 18:38:31 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1836

Learning Objectives

  • Compare and contrast anatomy and physiology, including their specializations and methods of study
  • Discuss the fundamental relationship between anatomy and physiology

Human anatomy is the scientific study of the body’s structures. Some of these structures are very small and can only be observed and analyzed with the assistance of a microscope. Other larger structures can readily be seen, manipulated, measured, and weighed. The word “anatomy” comes from a Greek root that means “to cut apart.” Human anatomy was first studied by observing the exterior of the body and observing the wounds of soldiers and other injuries. Later, physicians were allowed to dissect bodies of the dead to augment their knowledge. When a body is dissected, its structures are cut apart in order to observe their physical attributes and their relationships to one another. Dissection is still used in medical schools, anatomy courses, and in pathology labs. In order to observe structures in living people, however, a number of imaging techniques have been developed. These techniques allow clinicians to visualize structures inside the living body such as a cancerous tumor or a fractured bone.

Like most scientific disciplines, anatomy has areas of specialization. Gross anatomy is the study of the larger structures of the body, those visible without the aid of magnification (Figure 1a). Macro- means “large,” thus, gross anatomy is also referred to as macroscopic anatomy. In contrast, micro- means “small,” and microscopic anatomy is the study of structures that can be observed only with the use of a microscope or other magnification devices (Figure 1b). Microscopic anatomy includes cytology, the study of cells and histology, the study of tissues. As the technology of microscopes has advanced, anatomists have been able to observe smaller and smaller structures of the body, from slices of large structures like the heart, to the three-dimensional structures of large molecules in the body.

[caption id="" align="aligncenter" width="480"]Photo A shows an entire human brain which has a lumpy and deeply striated appearance. Photo B is a micrograph of neural tissue. It contains two roughly diamond-shaped cells with dark nuclei. The cells are embedded in a light colored tissue containing smaller cells and fiber strands. Figure 1. Gross and Microscopic Anatomy. (a) Gross anatomy considers large structures such as the brain. (b) Microscopic anatomy can deal with the same structures, though at a different scale. This is a micrograph of nerve cells from the brain. LM × 1600. (credit a: “WriterHound”/Wikimedia Commons; credit b: Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Anatomists take two general approaches to the study of the body’s structures: regional and systemic. Regional anatomy is the study of the interrelationships of all of the structures in a specific body region, such as the abdomen. Studying regional anatomy helps us appreciate the interrelationships of body structures, such as how muscles, nerves, blood vessels, and other structures work together to serve a particular body region. In contrast, systemic anatomy is the study of the structures that make up a discrete body system—that is, a group of structures that work together to perform a unique body function. For example, a systemic anatomical study of the muscular system would consider all of the skeletal muscles of the body.

Whereas anatomy is about structure, physiology is about function. Human physiology is the scientific study of the chemistry and physics of the structures of the body and the ways in which they work together to support the functions of life. Much of the study of physiology centers on the body’s tendency toward homeostasis. Homeostasis is the state of steady internal conditions maintained by living things. The study of physiology certainly includes observation, both with the naked eye and with microscopes, as well as manipulations and measurements. However, current advances in physiology usually depend on carefully designed laboratory experiments that reveal the functions of the many structures and chemical compounds that make up the human body.

Like anatomists, physiologists typically specialize in a particular branch of physiology. For example, neurophysiology is the study of the brain, spinal cord, and nerves and how these work together to perform functions as complex and diverse as vision, movement, and thinking. Physiologists may work from the organ level (exploring, for example, what different parts of the brain do) to the molecular level (such as exploring how an electrochemical signal travels along nerves).

Form is closely related to function in all living things. For example, the thin flap of your eyelid can snap down to clear away dust particles and almost instantaneously slide back up to allow you to see again. At the microscopic level, the arrangement and function of the nerves and muscles that serve the eyelid allow for its quick action and retreat. At a smaller level of analysis, the function of these nerves and muscles likewise relies on the interactions of specific molecules and ions. Even the three-dimensional structure of certain molecules is essential to their function.

Your study of anatomy and physiology will make more sense if you continually relate the form of the structures you are studying to their function. In fact, it can be somewhat frustrating to attempt to study anatomy without an understanding of the physiology that a body structure supports. Imagine, for example, trying to appreciate the unique arrangement of the bones of the human hand if you had no conception of the function of the hand. Fortunately, your understanding of how the human hand manipulates tools—from pens to cell phones—helps you appreciate the unique alignment of the thumb in opposition to the four fingers, making your hand a structure that allows you to pinch and grasp objects and type text messages

]]> 1836 1834 1 0 2.5 Organic Compounds Essential to Human Functioning https://pressbooks.bccampus.ca/apdouglas2017/chapter/2-5-organic-compounds-essential-to-human-functioning/ Wed, 02 Aug 2017 21:57:15 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1935

By the end of this section, you will be able to:

  • Identify four types of organic molecules essential to human functioning
  • Explain the chemistry behind carbon’s affinity for covalently bonding in organic compounds
  • Provide examples of three types of carbohydrates, and identify the primary functions of carbohydrates in the body
  • Discuss four types of lipids important in human functioning
  • Describe the structure of proteins, and discuss their importance to human functioning
  • Identify the building blocks of nucleic acids, and the roles of DNA, RNA, and ATP in human functioning

Organic compounds typically consist of groups of carbon atoms covalently bonded to hydrogen, usually oxygen, and often other elements as well. Created by living things, they are found throughout the world, in soils and seas, commercial products, and every cell of the human body. The four types most important to human structure and function are carbohydrates, lipids, proteins, and nucleotides. Before exploring these compounds, you need to first understand the chemistry of carbon.

The Chemistry of Carbon

What makes organic compounds ubiquitous is the chemistry of their carbon core. Recall that carbon atoms have four electrons in their valence shell, and that the octet rule dictates that atoms tend to react in such a way as to complete their valence shell with eight electrons. Carbon atoms do not complete their valence shells by donating or accepting four electrons. Instead, they readily share electrons via covalent bonds.

Commonly, carbon atoms share with other carbon atoms, often forming a long carbon chain referred to as a carbon skeleton. When they do share, however, they do not share all their electrons exclusively with each other. Rather, carbon atoms tend to share electrons with a variety of other elements, one of which is always hydrogen. Carbon and hydrogen groupings are called hydrocarbons. If you study the figures of organic compounds in the remainder of this chapter, you will see several with chains of hydrocarbons in one region of the compound.

Many combinations are possible to fill carbon’s four “vacancies.” Carbon may share electrons with oxygen or nitrogen or other atoms in a particular region of an organic compound. Moreover, the atoms to which carbon atoms bond may also be part of a functional group. A functional group is a group of atoms linked by strong covalent bonds and tending to function in chemical reactions as a single unit. You can think of functional groups as tightly knit “cliques” whose members are unlikely to be parted. Five functional groups are important in human physiology; these are the hydroxyl, carboxyl, amino, methyl and phosphate groups (Table 1).
Functional Groups Important in Human Physiology
Functional group Structural formula Importance
Hydroxyl —O—H Hydroxyl groups are polar. They are components of all four types of organic compounds discussed in this chapter. They are involved in dehydration synthesis and hydrolysis reactions.
Carboxyl O—C—OH Carboxyl groups are found within fatty acids, amino acids, and many other acids.
Amino —N—H2 Amino groups are found within amino acids, the building blocks of proteins.
Methyl —C—H3 Methyl groups are found within amino acids.
Phosphate —P—O42– Phosphate groups are found within phospholipids and nucleotides.

Carbon’s affinity for covalent bonding means that many distinct and relatively stable organic molecules nevertheless readily form larger, more complex molecules. Any large molecule is referred to as macromolecule (macro- = “large”), and the organic compounds in this section all fit this description. However, some macromolecules are made up of several “copies” of single units called monomer (mono- = “one”; -mer = “part”). Like beads in a long necklace, these monomers link by covalent bonds to form long polymers (poly- = “many”). There are many examples of monomers and polymers among the organic compounds.

Monomers form polymers by engaging in dehydration synthesis (see Chapter 2.4 Figure 1). As was noted earlier, this reaction results in the release of a molecule of water. Each monomer contributes: One gives up a hydrogen atom and the other gives up a hydroxyl group. Polymers are split into monomers by hydrolysis (-lysis = “rupture”). The bonds between their monomers are broken, via the donation of a molecule of water, which contributes a hydrogen atom to one monomer and a hydroxyl group to the other.

Carbohydrates

The term carbohydrate means “hydrated carbon.” Recall that the root hydro- indicates water. A carbohydrate is a molecule composed of carbon, hydrogen, and oxygen; in most carbohydrates, hydrogen and oxygen are found in the same two-to-one relative proportions they have in water. In fact, the chemical formula for a “generic” molecule of carbohydrate is (CH2O)n.

Carbohydrates are referred to as saccharides, a word meaning “sugars.” Three forms are important in the body. Monosaccharides are the monomers of carbohydrates. Disaccharides (di- = “two”) are made up of two monomers. Polysaccharides are the polymers, and can consist of hundreds to thousands of monomers.
[caption id="" align="alignleft" width="420"]This figure shows the structure of glucose, fructose, galactose, deoxyribose, and ribose. Figure 1. Five Important Monosaccharides.[/caption]

Monosaccharides

A monosaccharide is a monomer of carbohydrates. Five monosaccharides are important in the body. Three of these are the hexose sugars, so called because they each contain six atoms of carbon. These are glucose, fructose, and galactose, shown in Figure 1a. The remaining monosaccharides are the two pentose sugars, each of which contains five atoms of carbon. They are ribose and deoxyribose, shown in Figure 2b.

Disaccharides

A disaccharide is a pair of monosaccharides. Disaccharides are formed via dehydration synthesis, and the bond linking them is referred to as a glycosidic bond (glyco- = “sugar”). Three disaccharides (shown in Figure 2) are important to humans. These are sucrose, commonly referred to as table sugar; lactose, or milk sugar; and maltose, or malt sugar. As you can tell from their common names, you consume these in your diet; however, your body cannot use them directly. Instead, in the digestive tract, they are split into their component monosaccharides via hydrolysis.

[caption id="" align="aligncenter" width="420"]This figure shows the structure of sucrose, lactose, and maltose. Figure 2. Three Important Disaccharides. All three important disaccharides form by dehydration synthesis.[/caption]

Polysaccharides

Polysaccharides can contain a few to a thousand or more monosaccharides. Three are important to the body (Figure 3):

  • Starches are polymers of glucose. They occur in long chains called amylose or branched chains called amylopectin, both of which are stored in plant-based foods and are relatively easy to digest.
  • Glycogen is also a polymer of glucose, but it is stored in the tissues of animals, especially in the muscles and liver. It is not considered a dietary carbohydrate because very little glycogen remains in animal tissues after slaughter; however, the human body stores excess glucose as glycogen, again, in the muscles and liver.
  • Cellulose, a polysaccharide that is the primary component of the cell wall of green plants, is the component of plant food referred to as “fiber”. In humans, cellulose/fiber is not digestible; however, dietary fiber has many health benefits. It helps you feel full so you eat less, it promotes a healthy digestive tract, and a diet high in fiber is thought to reduce the risk of heart disease and possibly some forms of cancer.
[caption id="" align="aligncenter" width="412"]This figure shows the structure of starch, glycogen, and cellulose. Figure 3. Three Important Polysaccharides. Three important polysaccharides are starches, glycogen, and fiber.[/caption]

The body obtains carbohydrates from plant-based foods. Grains, fruits, and legumes and other vegetables provide most of the carbohydrate in the human diet, although lactose is found in dairy products. Although most body cells can break down other organic compounds for fuel, all body cells can use glucose. Moreover, nerve cells (neurons) in the brain, spinal cord, and through the peripheral nervous system, as well as red blood cells, can use only glucose for fuel. In the breakdown of glucose for energy, molecules of adenosine triphosphate, better known as ATP, are produced. Adenosine triphosphate (ATP) is composed of a ribose sugar, an adenine base, and three phosphate groups. ATP releases free energy when its phosphate bonds are broken, and thus supplies ready energy to the cell. More ATP is produced in the presence of oxygen (O2) than in pathways that do not use oxygen. The overall reaction for the conversion of the energy in glucose to energy stored in ATP can be written:
C6H12O6 + 6 O2 → 6 CO2 + 6 H2O + ATP

In addition to being a critical fuel source, carbohydrates are present in very small amounts in cells’ structure. For instance, some carbohydrate molecules bind with proteins to produce glycoproteins, and others combine with lipids to produce glycolipids, both of which are found in the membrane that encloses the contents of body cells.

Lipids

A lipid is one of a highly diverse group of compounds made up mostly of hydrocarbons. The few oxygen atoms they contain are often at the periphery of the molecule. Their nonpolar hydrocarbons make all lipids hydrophobic. In water, lipids do not form a true solution, but they may form an emulsion, which is the term for a mixture of solutions that do not mix well.

Triglycerides

A triglyceride is one of the most common dietary lipid groups, and the type found most abundantly in body tissues. This compound, which is commonly referred to as a fat, is formed from the synthesis of two types of molecules (Figure 4):

  • A glycerol backbone at the core of triglycerides, consists of three carbon atoms.
  • Three fatty acids, long chains of hydrocarbons with a carboxyl group and a methyl group at opposite ends, extend from each of the carbons of the glycerol.
[caption id="" align="alignleft" width="550"]This image shows the reaction for the formation of triglycerides. Figure 4. Triglycerides. Triglycerides are composed of glycerol attached to three fatty acids via dehydration synthesis. Notice that glycerol gives up a hydrogen atom, and the carboxyl groups on the fatty acids each give up a hydroxyl group.[/caption]

Triglycerides form via dehydration synthesis. Glycerol gives up hydrogen atoms from its hydroxyl groups at each bond, and the carboxyl group on each fatty acid chain gives up a hydroxyl group. A total of three water molecules are thereby released.

Fatty acid chains that have no double carbon bonds anywhere along their length and therefore contain the maximum number of hydrogen atoms are called saturated fatty acids. These straight, rigid chains pack tightly together and are solid or semi-solid at room temperature (Figure 5a). Butter and lard are examples, as is the fat found on a steak or in your own body. In contrast, fatty acids with one double carbon bond are kinked at that bond (Figure 5b). These monounsaturated fatty acids are therefore unable to pack together tightly, and are liquid at room temperature. Polyunsaturated fatty acids contain two or more double carbon bonds, and are also liquid at room temperature. Plant oils such as olive oil typically contain both mono- and polyunsaturated fatty acids.

[caption id="" align="alignright" width="380"]This diagram shows the chain structures of a saturated and an unsaturated fatty acid. Figure 5. Fatty Acid Shapes. The level of saturation of a fatty acid affects its shape. (a) Saturated fatty acid chains are straight. (b) Unsaturated fatty acid chains are kinked.[/caption]

Whereas a diet high in saturated fatty acids increases the risk of heart disease, a diet high in unsaturated fatty acids is thought to reduce the risk. This is especially true for the omega-3 unsaturated fatty acids found in cold-water fish such as salmon. These fatty acids have their first double carbon bond at the third hydrocarbon from the methyl group (referred to as the omega end of the molecule).

Finally, trans fatty acids found in some processed foods, including some stick and tub margarines, are thought to be even more harmful to the heart and blood vessels than saturated fatty acids. Trans fats are created from unsaturated fatty acids (such as corn oil) when chemically treated to produce partially hydrogenated fats.

As a group, triglycerides are a major fuel source for the body. When you are resting or asleep, a majority of the energy used to keep you alive is derived from triglycerides stored in your fat (adipose) tissues. Triglycerides also fuel long, slow physical activity such as gardening or hiking, and contribute a modest percentage of energy for vigorous physical activity. Dietary fat also assists the absorption and transport of the nonpolar fat-soluble vitamins A, D, E, and K. Additionally, stored body fat protects and cushions the body’s bones and internal organs, and acts as insulation to retain body heat.

Fatty acids are also components of glycolipids, which are sugar-fat compounds found in the cell membrane. Lipoproteins are compounds in which the hydrophobic triglycerides are packaged in protein envelopes for transport in body fluids.

Phospholipids

As its name suggests, a phospholipid is a bond between the glycerol component of a lipid and a phosphorous molecule. In fact, phospholipids are similar in structure to triglycerides. However, instead of having three fatty acids, a phospholipid is generated from a diglyceride, a glycerol with just two fatty acid chains (Figure 6). The third binding site on the glycerol is taken up by the phosphate group, which in turn is attached to a polar “head” region of the molecule. Recall that triglycerides are nonpolar and hydrophobic. This still holds for the fatty acid portion of a phospholipid compound. However, the head of a phospholipid contains charges on the phosphate groups, as well as on the nitrogen atom. These charges make the phospholipid head hydrophilic. Therefore, phospholipids are said to have hydrophobic tails, containing the neutral fatty acids, and hydrophilic heads, containing the charged phosphate groups and nitrogen atom.

[caption id="" align="alignleft" width="600"]This figure shows the chemical structure of different lipids. Figure 6. Other Important Lipids. (a) Phospholipids are composed of two fatty acids, glycerol, and a phosphate group. (b) Sterols are ring-shaped lipids. Shown here is cholesterol. (c) Prostaglandins are derived from unsaturated fatty acids. Prostaglandin E2 (PGE2) includes hydroxyl and carboxyl groups.[/caption]

Steroids

A steroid compound (referred to as a sterol) has as its foundation a set of four hydrocarbon rings bonded to a variety of other atoms and molecules (see Figure 6b). Although both plants and animals synthesize sterols, the type that makes the most important contribution to human structure and function is cholesterol, which is synthesized by the liver in humans and animals and is also present in most animal-based foods. Like other lipids, cholesterol’s hydrocarbons make it hydrophobic; however, it has a polar hydroxyl head that is hydrophilic. Cholesterol is an important component of bile acids, compounds that help emulsify dietary fats. In fact, the word root chole- refers to bile. Cholesterol is also a building block of many hormones, signaling molecules that the body releases to regulate processes at distant sites. Finally, like phospholipids, cholesterol molecules are found in the cell membrane, where their hydrophobic and hydrophilic regions help regulate the flow of substances into and out of the cell.

Prostaglandins

Like a hormone, a prostaglandin is one of a group of signaling molecules, but prostaglandins are derived from unsaturated fatty acids (see Figure 6c). One reason that the omega-3 fatty acids found in fish are beneficial is that they stimulate the production of certain prostaglandins that help regulate aspects of blood pressure and inflammation, and thereby reduce the risk for heart disease. Prostaglandins also sensitize nerves to pain. One class of pain-relieving medications called nonsteroidal anti-inflammatory drugs (NSAIDs) works by reducing the effects of prostaglandins.

Proteins

You might associate proteins with muscle tissue, but in fact, proteins are critical components of all tissues and organs. A protein is an organic molecule composed of amino acids linked by peptide bonds. Proteins include the keratin in the epidermis of skin that protects underlying tissues, the collagen found in the dermis of skin, in bones, and in the meninges that cover the brain and spinal cord. Proteins are also components of many of the body’s functional chemicals, including digestive enzymes in the digestive tract, antibodies, the neurotransmitters that neurons use to communicate with other cells, and the peptide-based hormones that regulate certain body functions (for instance, growth hormone). While carbohydrates and lipids are composed of hydrocarbons and oxygen, all proteins also contain nitrogen (N), and many contain sulfur (S), in addition to carbon, hydrogen, and oxygen.

Microstructure of Proteins

Proteins are polymers made up of nitrogen-containing monomers called amino acids. An amino acid is a molecule composed of an amino group and a carboxyl group, together with a variable side chain. Just 20 different amino acids contribute to nearly all of the thousands of different proteins important in human structure and function. Body proteins contain a unique combination of a few dozen to a few hundred of these 20 amino acid monomers. All 20 of these amino acids share a similar structure (Figure 7). All consist of a central carbon atom to which the following are bonded:

[caption id="" align="alignright" width="320"]This figure shows the structure of an amino acid. Figure 8. Structure of an Amino Acid[/caption]
  • a hydrogen atom
  • an alkaline (basic) amino group NH2 (see Table 1)
  • an acidic carboxyl group COOH (see Table 1)
  • a variable group

Notice that all amino acids contain both an acid (the carboxyl group) and a base (the amino group) (amine = “nitrogen-containing”). For this reason, they make excellent buffers, helping the body regulate acid–base balance. What distinguishes the 20 amino acids from one another is their variable group, which is referred to as a side chain or an R-group. This group can vary in size and can be polar or nonpolar, giving each amino acid its unique characteristics. For example, the side chains of two amino acids—cysteine and methionine—contain sulfur. Sulfur does not readily participate in hydrogen bonds, whereas all other amino acids do. This variation influences the way that proteins containing cysteine and methionine are assembled.

Amino acids join via dehydration synthesis to form protein polymers (Figure 8). The unique bond holding amino acids together is called a peptide bond. A peptide bond is a covalent bond between two amino acids that forms by dehydration synthesis. A peptide, in fact, is a very short chain of amino acids. Strands containing fewer than about 100 amino acids are generally referred to as polypeptides rather than proteins. [caption id="" align="alignright" width="280"]This figure shows the formation of a peptide bond, highlighted in blue. Figure 8.Peptide Bond. Different amino acids join together to form peptides, polypeptides, or proteins via dehydration synthesis. The bonds between the amino acids are peptide bonds.[/caption]

The body is able to synthesize most of the amino acids from components of other molecules; however, nine cannot be synthesized and have to be consumed in the diet. These are known as the essential amino acids.

Free amino acids available for protein construction are said to reside in the amino acid pool within cells. Structures within cells use these amino acids when assembling proteins. If a particular essential amino acid is not available in sufficient quantities in the amino acid pool, however, synthesis of proteins containing it can slow or even cease.

Shape of Proteins

Just as a fork cannot be used to eat soup and a spoon cannot be used to spear meat, a protein’s shape is essential to its function. A protein’s shape is determined, most fundamentally, by the sequence of amino acids of which it is made (Figure 9a).

The sequence is called the primary structure of the protein. [caption id="" align="alignright" width="480"]This figure shows the secondary structure of peptides. The top panel shows a straight chain, the middle panel shows an alpha-helix and a beta sheet. The bottom panel shows the tertiary structure and fully folded protein. Figure 9. The Shape of Proteins. (a) The primary structure is the sequence of amino acids that make up the polypeptide chain. (b) The secondary structure, which can take the form of an alpha-helix or a beta-pleated sheet, is maintained by hydrogen bonds between amino acids in different regions of the original polypeptide strand. (c) The tertiary structure occurs as a result of further folding and bonding of the secondary structure. (d) The quaternary structure occurs as a result of interactions between two or more tertiary subunits. The example shown here is hemoglobin, a protein in red blood cells which transports oxygen to body tissues.[/caption]

Although some polypeptides exist as linear chains, most are twisted or folded into more complex secondary structures that form when bonding occurs between amino acids with different properties at different regions of the polypeptide. The most common secondary structure is a spiral called an alpha-helix. If you were to take a length of string and simply twist it into a spiral, it would not hold the shape. Similarly, a strand of amino acids could not maintain a stable spiral shape without the help of hydrogen bonds, which create bridges between different regions of the same strand (see Figure 9b). Less commonly, a polypeptide chain can form a beta-pleated sheet, in which hydrogen bonds form bridges between different regions of a single polypeptide that has folded back upon itself, or between two or more adjacent polypeptide chains.

The secondary structure of proteins further folds into a compact three-dimensional shape, referred to as the protein’s tertiary structure (see Figure 9c). In this configuration, amino acids that had been very distant in the primary chain can be brought quite close via hydrogen bonds or, in proteins containing cysteine, via disulfide bonds. A disulfide bond is a covalent bond between sulfur atoms in a polypeptide. Often, two or more separate polypeptides bond to form an even larger protein with a quaternary structure (see Figure 9d). The polypeptide subunits forming a quaternary structure can be identical or different. For instance, hemoglobin, the protein found in red blood cells is composed of four tertiary polypeptides, two of which are called alpha chains and two of which are called beta chains.

When they are exposed to extreme heat, acids, bases, and certain other substances, proteins will denature. Denaturation is a change in the structure of a molecule through physical or chemical means. Denatured proteins lose their functional shape and are no longer able to carry out their jobs. An everyday example of protein denaturation is the curdling of milk when acidic lemon juice is added.

The contribution of the shape of a protein to its function can hardly be exaggerated. For example, the long, slender shape of protein strands that make up muscle tissue is essential to their ability to contract (shorten) and relax (lengthen). As another example, bones contain long threads of a protein called collagen that acts as scaffolding upon which bone minerals are deposited. These elongated proteins, called fibrous proteins, are strong and durable and typically hydrophobic.

In contrast, globular proteins are globes or spheres that tend to be highly reactive and are hydrophilic. The hemoglobin proteins packed into red blood cells are an example (see Figure 9d); however, globular proteins are abundant throughout the body, playing critical roles in most body functions. Enzymes, introduced earlier as protein catalysts, are examples of this. The next section takes a closer look at the action of enzymes.

Proteins Function as Enzymes

If you were trying to type a paper, and every time you hit a key on your laptop there was a delay of six or seven minutes before you got a response, you would probably get a new laptop. In a similar way, without enzymes to catalyze chemical reactions, the human body would be nonfunctional. It functions only because enzymes function.

Enzymatic reactions—chemical reactions catalyzed by enzymes—begin when substrates bind to the enzyme. A substrate is a reactant in an enzymatic reaction. This occurs on regions of the enzyme known as active sites (Figure 10). Any given enzyme catalyzes just one type of chemical reaction. This characteristic, called specificity, is due to the fact that a substrate with a particular shape and electrical charge can bind only to an active site corresponding to that substrate.

[caption id="" align="alignleft" width="520"]This image shows the steps in which an enzyme can act. The substrate is shown binding to the enzyme, forming a product, and the detachment of the product. Figure 10. Steps in an Enzymatic Reaction. (a) Substrates approach active sites on enzyme. (b) Substrates bind to active sites, producing an enzyme–substrate complex. (c) Changes internal to the enzyme–substrate complex facilitate interaction of the substrates. (d) Products are released and the enzyme returns to its original form, ready to facilitate another enzymatic reaction.[/caption]

Binding of a substrate produces an enzyme–substrate complex. It is likely that enzymes speed up chemical reactions in part because the enzyme–substrate complex undergoes a set of temporary and reversible changes that cause the substrates to be oriented toward each other in an optimal position to facilitate their interaction. This promotes increased reaction speed. The enzyme then releases the product(s), and resumes its original shape. The enzyme is then free to engage in the process again, and will do so as long as substrate remains.

Other Functions of Proteins

Advertisements for protein bars, powders, and shakes all say that protein is important in building, repairing, and maintaining muscle tissue, but the truth is that proteins contribute to all body tissues, from the skin to the brain cells. Also, certain proteins act as hormones, chemical messengers that help regulate body functions, For example, growth hormone is important for skeletal growth, among other roles.

As was noted earlier, the basic and acidic components enable proteins to function as buffers in maintaining acid–base balance, but they also help regulate fluid–electrolyte balance. Proteins attract fluid, and a healthy concentration of proteins in the blood, the cells, and the spaces between cells helps ensure a balance of fluids in these various “compartments.” Moreover, proteins in the cell membrane help to transport electrolytes in and out of the cell, keeping these ions in a healthy balance. Like lipids, proteins can bind with carbohydrates. They can thereby produce glycoproteins or proteoglycans, both of which have many functions in the body.

The body can use proteins for energy when carbohydrate and fat intake is inadequate, and stores of glycogen and adipose tissue become depleted. However, since there is no storage site for protein except functional tissues, using protein for energy causes tissue breakdown, and results in body wasting.

Nucleotides

The fourth type of organic compound important to human structure and function are the nucleotides (Figure 11). A nucleotide is one of a class of organic compounds composed of three subunits:

  • one or more phosphate groups
  • a pentose sugar: either deoxyribose or ribose
  • a nitrogen-containing base: adenine, cytosine, guanine, thymine, or uracil

Nucleotides can be assembled into nucleic acids (DNA or RNA) or the energy compound adenosine triphosphate.

[caption id="" align="alignright" width="520"]This figure shows the structure of nucleotides. Figure 11. Nucleotides. (a) The building blocks of all nucleotides are one or more phosphate groups, a pentose sugar, and a nitrogen-containing base. (b) The nitrogen-containing bases of nucleotides. (c) The two pentose sugars of DNA and RNA.[/caption]

Nucleic Acids

The nucleic acids differ in their type of pentose sugar. Deoxyribonucleic acid (DNA) is nucleotide that stores genetic information. DNA contains deoxyribose (so-called because it has one less atom of oxygen than ribose) plus one phosphate group and one nitrogen-containing base. The “choices” of base for DNA are adenine, cytosine, guanine, and thymine. Ribonucleic acid (RNA) is a ribose-containing nucleotide that helps manifest the genetic code as protein. RNA contains ribose, one phosphate group, and one nitrogen-containing base, but the “choices” of base for RNA are adenine, cytosine, guanine, and uracil.

The nitrogen-containing bases adenine and guanine are classified as purines. A purine is a nitrogen-containing molecule with a double ring structure, which accommodates several nitrogen atoms. The bases cytosine, thymine (found in DNA only) and uracil (found in RNA only) are pyramidines. A pyramidine is a nitrogen-containing base with a single ring structure

[caption id="" align="alignleft" width="320"]This figure shows a double helix. Figure 12. DNA. In the DNA double helix, two strands attach via hydrogen bonds between the bases of the component nucleotides.[/caption]

Bonds formed by dehydration synthesis between the pentose sugar of one nucleic acid monomer and the phosphate group of another form a “backbone,” from which the components’ nitrogen-containing bases protrude. In DNA, two such backbones attach at their protruding bases via hydrogen bonds. These twist to form a shape known as a double helix (Figure 12). The sequence of nitrogen-containing bases within a strand of DNA form the genes that act as a molecular code instructing cells in the assembly of amino acids into proteins. Humans have almost 22,000 genes in their DNA, locked up in the 46 chromosomes inside the nucleus of each cell (except red blood cells which lose their nuclei during development). These genes carry the genetic code to build one’s body, and are unique for each individual except identical twins.

In contrast, RNA consists of a single strand of sugar-phosphate backbone studded with bases. Messenger RNA (mRNA) is created during protein synthesis to carry the genetic instructions from the DNA to the cell’s protein manufacturing plants in the cytoplasm, the ribosomes.

Adenosine Triphosphate

[caption id="" align="alignright" width="420"]This figure shows the structure of ATP. Figure 13. Structure of Adenosine Triphosphate (ATP).[/caption]

The nucleotide adenosine triphosphate (ATP), is composed of a ribose sugar, an adenine base, and three phosphate groups (Figure 13). ATP is classified as a high energy compound because the two covalent bonds linking its three phosphates store a significant amount of potential energy. In the body, the energy released from these high energy bonds helps fuel the body’s activities, from muscle contraction to the transport of substances in and out of cells to anabolic chemical reactions.

When a phosphate group is cleaved from ATP, the products are adenosine diphosphate (ADP) and inorganic phosphate (Pi). This hydrolysis reaction can be written:

ATP + H2O → ADP + Pi + energy

Removal of a second phosphate leaves adenosine monophosphate (AMP) and two phosphate groups. Again, these reactions also liberate the energy that had been stored in the phosphate-phosphate bonds. They are reversible, too, as when ADP undergoes phosphorylation. Phosphorylation is the addition of a phosphate group to an organic compound, in this case, resulting in ATP. In such cases, the same level of energy that had been released during hydrolysis must be reinvested to power dehydration synthesis.

Cells can also transfer a phosphate group from ATP to another organic compound. For example, when glucose first enters a cell, a phosphate group is transferred from ATP, forming glucose phosphate (C6H12O6—P) and ADP. Once glucose is phosphorylated in this way, it can be stored as glycogen or metabolized for immediate energy.

[caption id="attachment_3052" align="aligncenter" width="150"] Watch this amoeba sisters video to learn more about biomolecules![/caption]
]]> 1935 1901 5 0 Introduction https://pressbooks.bccampus.ca/apdouglas2017/chapter/introduction-2/ Wed, 02 Aug 2017 20:22:02 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1938 In this image, a fluorescently stained cell is shown undergoing mitosis. The cell membrane is stained red and the green stains show the mitotic spindles inside the cell. The chromosomes are shown in blue. Figure 1. Fluorescence-stained Cell Undergoing Mitosis. A lung cell from a newt, commonly studied for its similarity to human lung cells, is stained with fluorescent dyes. The green stain reveals mitotic spindles, red is the cell membrane and part of the cytoplasm, and the structures that appear light blue are chromosomes. This cell is in anaphase of mitosis. (credit: “Mortadelo2005”/Wikimedia Commons)[/caption]

Chapter Objectives

After studying this chapter, you will be able to:
  • Describe the structure and function of the cell membrane, including its regulation of materials into and out of the cell
  • Describe the functions of the various cytoplasmic organelles
  • Explain the structure and contents of the nucleus, as well as the process of DNA replication
  • Explain the process by which a cell builds proteins using the DNA code
  • List the stages of the cell cycle in order, including the steps of cell division in somatic cells
  • Discuss how a cell differentiates and becomes more specialized
  • List the morphological and physiological characteristics of some representative cell types in the human body
You developed from a single fertilized egg cell into the complex organism containing trillions of cells that you see when you look in a mirror. During this developmental process, early, undifferentiated cells differentiate and become specialized in their structure and function. These different cell types form specialized tissues that work in concert to perform all of the functions necessary for the living organism. Cellular and developmental biologists study how the continued division of a single cell leads to such complexity and differentiation. Consider the difference between a structural cell in the skin and a nerve cell. A structural skin cell may be shaped like a flat plate (squamous) and live only for a short time before it is shed and replaced. Packed tightly into rows and sheets, the squamous skin cells provide a protective barrier for the cells and tissues that lie beneath. A nerve cell, on the other hand, may be shaped something like a star, sending out long processes up to a meter in length and may live for the entire lifetime of the organism. With their long winding appendages, nerve cells can communicate with one another and with other types of body cells and send rapid signals that inform the organism about its environment and allow it to interact with that environment. These differences illustrate one very important theme that is consistent at all organizational levels of biology: the form of a structure is optimally suited to perform particular functions assigned to that structure. Keep this theme in mind as you tour the inside of a cell and are introduced to the various types of cells in the body. A primary responsibility of each cell is to contribute to homeostasis. Homeostasis is a term used in biology that refers to a dynamic state of balance within parameters that are compatible with life. For example, living cells require a water-based environment to survive in, and there are various physical (anatomical) and physiological mechanisms that keep all of the trillions of living cells in the human body moist. This is one aspect of homeostasis. When a particular parameter, such as blood pressure or blood oxygen content, moves far enough out of homeostasis (generally becoming too high or too low), illness or disease—and sometimes death—inevitably results. The concept of a cell started with microscopic observations of dead cork tissue by scientist Robert Hooke in 1665. Without realizing their function or importance, Hook coined the term “cell” based on the resemblance of the small subdivisions in the cork to the rooms that monks inhabited, called cells. About ten years later, Antonie van Leeuwenhoek became the first person to observe living and moving cells under a microscope. In the century that followed, the theory that cells represented the basic unit of life would develop. These tiny fluid-filled sacs house components responsible for the thousands of biochemical reactions necessary for an organism to grow and survive. In this chapter, you will learn about the major components and functions of a prototypical, generalized cell and discover some of the different types of cells in the human body. [caption id="attachment_3048" align="aligncenter" width="150"] Watch this amoeba sisters video for an introduction to the cell![/caption]]]> 1938 1936 1 0 3.6 Cellular Differentiation https://pressbooks.bccampus.ca/apdouglas2017/chapter/3-6-cellular-differentiation/ Wed, 02 Aug 2017 18:36:54 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=1976

Learning Objectives

By the end of this section, you will be able to:
  • Discuss how the generalized cells of a developing embryo or the stem cells of an adult organism become differentiated into specialized cells
  • Distinguish between the categories of stem cells
How does a complex organism such as a human develop from a single cell—a fertilized egg—into the vast array of cell types such as nerve cells, muscle cells, and epithelial cells that characterize the adult? Throughout development and adulthood, the process of cellular differentiation leads cells to assume their final morphology and physiology. Differentiation is the process by which unspecialized cells become specialized to carry out distinct functions.

Stem Cells

A stem cell is an unspecialized cell that can divide without limit as needed and can, under specific conditions, differentiate into specialized cells. Stem cells are divided into several categories according to their potential to differentiate.

The first embryonic cells that arise from the division of the zygote are the ultimate stem cells; these stems cells are described as totipotent because they have the potential to differentiate into any of the cells needed to enable an organism to grow and develop.

The embryonic cells that develop from totipotent stem cells and are precursors to the fundamental tissue layers of the embryo are classified as pluripotent. A pluripotent stem cell is one that has the potential to differentiate into any type of human tissue but cannot support the full development of an organism. These cells then become slightly more specialized, and are referred to as multipotent cells.

A multipotent stem cell has the potential to differentiate into different types of cells within a given cell lineage or small number of lineages, such as a red blood cell or white blood cell.

Finally, multipotent cells can become further specialized oligopotent cells. An oligopotent stem cell is limited to becoming one of a few different cell types. In contrast, a unipotent cell is fully specialized and can only reproduce to generate more of its own specific cell type.

Stem cells are unique in that they can also continually divide and regenerate new stem cells instead of further specializing. There are different stem cells present at different stages of a human’s life. They include the embryonic stem cells of the embryo, fetal stem cells of the fetus, and adult stem cells in the adult. One type of adult stem cell is the epithelial stem cell, which gives rise to the keratinocytes in the multiple layers of epithelial cells in the epidermis of skin. Adult bone marrow has three distinct types of stem cells: hematopoietic stem cells, which give rise to red blood cells, white blood cells, and platelets (Figure 1); endothelial stem cells, which give rise to the endothelial cell types that line blood and lymph vessels; and mesenchymal stem cells, which give rise to the different types of muscle cells.

[caption id="" align="aligncenter" width="600"]This flowchart shows the differentiation of a hemocytoblast, a stem cell, into the different types of cells found in blood. Figure 1. Hematopoiesis. The process of hematopoiesis involves the differentiation of multipotent cells into blood and immune cells. The multipotent hematopoietic stem cells give rise to many different cell types, including the cells of the immune system and red blood cells.[/caption]

Differentiation

When a cell differentiates (becomes more specialized), it may undertake major changes in its size, shape, metabolic activity, and overall function. Because all cells in the body, beginning with the fertilized egg, contain the same DNA, how do the different cell types come to be so different? The answer is analogous to a movie script. The different actors in a movie all read from the same script, however, they are each only reading their own part of the script. Similarly, all cells contain the same full complement of DNA, but each type of cell only “reads” the portions of DNA that are relevant to its own function. In biology, this is referred to as the unique genetic expression of each cell.

In order for a cell to differentiate into its specialized form and function, it need only manipulate those genes (and thus those proteins) that will be expressed, and not those that will remain silent. The primary mechanism by which genes are turned “on” or “off” is through transcription factors. A transcription factor is one of a class of proteins that bind to specific genes on the DNA molecule and either promote or inhibit their transcription (Figure 2).

[caption id="" align="aligncenter" width="525"]This diagram shows transcription factors and then RNA polymerase binding to a stretch of RNA to initiate transcription. Figure 2. Transcription Factors Regulate Gene Expression. While each body cell contains the organism’s entire genome, different cells regulate gene expression with the use of various transcription factors. Transcription factors are proteins that affect the binding of RNA polymerase to a particular gene on the DNA molecule.[/caption]
Everyday Connection

Stem Cell Research Stem cell research aims to find ways to use stem cells to regenerate and repair cellular damage. Over time, most adult cells undergo the wear and tear of aging and lose their ability to divide and repair themselves. Stem cells do not display a particular morphology or function. Adult stem cells, which exist as a small subset of cells in most tissues, keep dividing and can differentiate into a number of specialized cells generally formed by that tissue. These cells enable the body to renew and repair body tissues.

The mechanisms that induce a non-differentiated cell to become a specialized cell are poorly understood. In a laboratory setting, it is possible to induce stem cells to differentiate into specialized cells by changing the physical and chemical conditions of growth. Several sources of stem cells are used experimentally and are classified according to their origin and potential for differentiation. Human embryonic stem cells (hESCs) are extracted from embryos and are pluripotent. The adult stem cells that are present in many organs and differentiated tissues, such as bone marrow and skin, are multipotent, being limited in differentiation to the types of cells found in those tissues. The stem cells isolated from umbilical cord blood are also multipotent, as are cells from deciduous teeth (baby teeth). Researchers have recently developed induced pluripotent stem cells (iPSCs) from mouse and human adult stem cells. These cells are genetically reprogrammed multipotent adult cells that function like embryonic stem cells; they are capable of generating cells characteristic of all three germ layers.

Because of their capacity to divide and differentiate into specialized cells, stem cells offer a potential treatment for diseases such as diabetes and heart disease (Figure 3). Cell-based therapy refers to treatment in which stem cells induced to differentiate in a growth dish are injected into a patient to repair damaged or destroyed cells or tissues. Many obstacles must be overcome for the application of cell-based therapy. Although embryonic stem cells have a nearly unlimited range of differentiation potential, they are seen as foreign by the patient’s immune system and may trigger rejection. Also, the destruction of embryos to isolate embryonic stem cells raises considerable ethical and legal questions.

[caption id="" align="aligncenter" width="500"]This flow chart shows the differentiation of stem cells into different cell types. The top layer shows a totipotent stem cell, which becomes a pluripotent stem cell and then a multipotent stem cell. A multipotent stem cell can then differentiate into different cell types. Figure 3. Stem Cells. The capacity of stem cells to differentiate into specialized cells make them potentially valuable in therapeutic applications designed to replace damaged cells of different body tissues.[/caption]

In contrast, adult stem cells isolated from a patient are not seen as foreign by the body, but they have a limited range of differentiation. Some individuals bank the cord blood or deciduous teeth of their child, storing away those sources of stem cells for future use, should their child need it. Induced pluripotent stem cells are considered a promising advance in the field because using them avoids the legal, ethical, and immunological pitfalls of embryonic stem cells.

  [caption id="attachment_3042" align="aligncenter" width="150"] Watch this amoeba sisters video to learn more about how cells become specialized![/caption] [caption id="attachment_3043" align="aligncenter" width="150"] Watch this amoeba sisters video for some examples of specialized cells![/caption]

multipotent
describes the condition of being able to differentiate into different types of cells within a given cell lineage or small number of lineages, such as a red blood cell or white blood cell
oligopotent
describes the condition of being more specialized than multipotency; the condition of being able to differentiate into one of a few possible cell types
pluripotent
describes the condition of being able to differentiate into a large variety of cell types
stem cell
cell that is oligo-, multi-, or pleuripotent that has the ability to produce additional stem cells rather than becoming further specialized
totipotent
embryonic cells that have the ability to differentiate into any type of cell and organ in the body
transcription factor
one of the proteins that regulate the transcription of genes
unipotent
describes the condition of being committed to a single specialized cell type
]]> 1976 1936 7 0 6.3 Bone Structure https://pressbooks.bccampus.ca/apdouglas2017/chapter/6-3-bone-structure/ Wed, 02 Aug 2017 21:58:04 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2043

Learning Objectives

By the end of this section, you will be able to:
  • Identify the anatomical features of a bone
  • Define and list examples of bone markings
  • Describe the histology of bone tissue
  • Compare and contrast compact and spongy bone
  • Identify the structures that compose compact and spongy bone
  • Describe how bones are nourished and innervated

Bone tissue (osseous tissue) differs greatly from other tissues in the body. Bone is hard and many of its functions depend on that characteristic hardness. Later discussions in this chapter will show that bone is also dynamic in that its shape adjusts to accommodate stresses. This section will examine the gross anatomy of bone first and then move on to its histology.

Gross Anatomy of Bone

The structure of a long bone allows for the best visualization of all of the parts of a bone (Figure 1). A long bone has two parts: the diaphysis and the epiphysis. The diaphysis is the tubular shaft that runs between the proximal and distal ends of the bone. The hollow region in the diaphysis is called the medullary cavity, which is filled with yellow marrow. The walls of the diaphysis are composed of dense and hard compact bone.

[caption id="" align="aligncenter" width="320"]This illustration depicts an anterior view of the right femur, or thigh bone. The inferior end that connects to the knee is at the bottom of the diagram and the superior end that connects to the hip is at the top of the diagram. The bottom end of the bone contains a smaller lateral bulge and a larger medial bulge. A blue articular cartilage covers the inner half of each bulge as well as the small trench that runs between the bulges. This area of the inferior end of the bone is labeled the distal epiphysis. Above the distal epiphysis is the metaphysis, where the bone tapers from the wide epiphysis into the relatively thin shaft. The entire length of the shaft is the diaphysis. The superior half of the femur is cut away to show its internal contents. The bone is covered with an outer translucent sheet called the periosteum. At the midpoint of the diaphysis, a nutrient artery travels through the periosteum and into the inner layers of the bone. The periosteum surrounds a white cylinder of solid bone labeled compact bone. The cavity at the center of the compact bone is called the medullary cavity. The inner layer of the compact bone that lines the medullary cavity is called the endosteum. Within the diaphysis, the medullary cavity contains a cylinder of yellow bone marrow that is penetrated by the nutrient artery. The superior end of the femur is also connected to the diaphysis by a metaphysis. In this upper metaphysis, the bone gradually widens between the diaphysis and the proximal epiphysis. The proximal epiphysis of the femur is roughly hexagonal in shape. However, the upper right side of the hexagon has a large, protruding knob. The femur connects and rotates within the hip socket at this knob. The knob is covered with a blue colored articular cartilage. The internal anatomy of the upper metaphysis and proximal epiphysis are revealed. The medullary cavity in these regions is filled with the mesh like spongy bone. Red bone marrow occupies the many cavities within the spongy bone. There is a clear, white line separating the spongy bone of the upper metaphysis with that of the proximal epiphysis. This line is labeled the epiphyseal line. Figure 1. Anatomy of a Long Bone. A typical long bone shows the gross anatomical characteristics of bone.[/caption]

The wider section at each end of the bone is called the epiphysis (plural = epiphyses), which is filled with spongy bone. Red marrow fills the spaces in the spongy bone. Each epiphysis meets the diaphysis at the metaphysis, the narrow area that contains the epiphyseal plate (growth plate), a layer of hyaline (transparent) cartilage in a growing bone. When the bone stops growing in early adulthood (approximately 18–21 years), the cartilage is replaced by osseous tissue and the epiphyseal plate becomes an epiphyseal line.

The medullary cavity has a delicate membranous lining called the endosteum (end- = “inside”; oste- = “bone”), where bone growth, repair, and remodeling occur. The outer surface of the bone is covered with a fibrous membrane called the periosteum (peri- = “around” or “surrounding”). The periosteum contains blood vessels, nerves, and lymphatic vessels that nourish compact bone. Tendons and ligaments also attach to bones at the periosteum. The periosteum covers the entire outer surface except where the epiphyses meet other bones to form joints (Figure 2). In this region, the epiphyses are covered with articular cartilage, a thin layer of cartilage that reduces friction and acts as a shock absorber.

[caption id="" align="aligncenter" width="480"]The top of this illustration shows an anterior view of the proximal end of the femur. The top image has two zoom in boxes. The left box is situated on the border between the diaphysis and the metaphysis. Its callout magnifies the periosteum on the right side of the femur. The view shows that the periosteum contains an outer fibrous layer composed of yellow fibers. The inner layer of the periosteum is called the cellular layer, which is composed of irregularly shaped cells. The cellular layer gradually shrinks in width as it transitions from the metaphysis to the diaphysis. A small blood vessel runs through both layers and enters the bone. The right zoom in box magnifies the endosteum on the left side of the bone. The box is situated just inferior to the border between the diaphysis and the metaphysic. It calls out the inner edge of the compact bone layer. The magnified view shows concentric circles of dark colored bone matrix. Between the circles are small cavities containing orange, diamond-shaped cells labeled osteocytes. The left edge of the bone matrix is lined with a single layer of flattened cells called the endosteum. There is a large cell, labeled an osteoclast, between two of the endosteum cells. The osteoclast is cutting a depression into the bony matrix under the endosteum. At another part of the endosteum, three smaller osteoblasts are secreting a blue substance that builds up the outermost layer of the bony matrix. Figure 2. Periosteum and Endosteum. The periosteum forms the outer surface of bone, and the endosteum lines the medullary cavity.[/caption]
Flat bones, like those of the cranium, consist of a layer of diploë (spongy bone), lined on either side by a layer of compact bone (Figure 3). The two layers of compact bone and the interior spongy bone work together to protect the internal organs. If the outer layer of a cranial bone fractures, the brain is still protected by the intact inner layer.
[caption id="" align="aligncenter" width="420"]This illustration shows a cross section of a cranial bone, constructed somewhat like a sandwich. The topmost and bottommost layers are the thin, translucent, periosteum. The upper and lower periosteum cover an upper and lower layer of compact bone, respectively. The compact bone is solid, with each layer occupying about one tenth of the thickness of the cranial bone. The majority of the cross section is occupied by the spongy bone, or diploe, sandwiched between the upper and lower compact bone. The spongy bone contains many crisscrossing threads of bone. Dark air spaces occur between the threads, giving the bone a porous appearance, much like that of a sponge or Swiss cheese. Figure 3. Anatomy of a Flat Bone. This cross-section of a flat bone shows the spongy bone (diploë) lined on either side by a layer of compact bone.[/caption]

Bone Markings

The surface features of bones vary considerably, depending on the function and location in the body. Table 2 describes the bone markings, which are illustrated in (Figure 4). There are three general classes of bone markings: (1) articulations, (2) projections, and (3) holes. As the name implies, an articulation is where two bone surfaces come together (articulus = “joint”). These surfaces tend to conform to one another, such as one being rounded and the other cupped, to facilitate the function of the articulation. A projection is an area of a bone that projects above the surface of the bone. These are the attachment points for tendons and ligaments. In general, their size and shape is an indication of the forces exerted through the attachment to the bone. A hole is an opening or groove in the bone that allows blood vessels and nerves to enter the bone. As with the other markings, their size and shape reflect the size of the vessels and nerves that penetrate the bone at these points.

Bone Markings (Table 2)
Marking Description Example
Articulations Where two bones meet Knee joint
Head Prominent rounded surface Head of femur
Facet Flat surface Vertebrae
Condyle Rounded surface Occipital condyles
Projections Raised markings Spinous process of the vertebrae
Protuberance Protruding Chin
Process Prominence feature Transverse process of vertebra
Spine Sharp process Ischial spine
Tubercle Small, rounded process Tubercle of humerus
Tuberosity Rough surface Deltoid tuberosity
Line Slight, elongated ridge Temporal lines of the parietal bones
Crest Ridge Iliac crest
Holes Holes and depressions Foramen (holes through which blood vessels can pass through)
Fossa Elongated basin Mandibular fossa
Fovea Small pit Fovea capitis on the head of the femur
Sulcus Groove Sigmoid sulcus of the temporal bones
Canal Passage in bone Auditory canal
Fissure Slit through bone Auricular fissure
Foramen Hole through bone Foramen magnum in the occipital bone
Meatus Opening into canal External auditory meatus
Sinus Air-filled space in bone Nasal sinus
[caption id="" align="aligncenter" width="500"]This illustration contains three diagrams. The left diagram is titled examples of processes formed where tendons or ligaments attach. The image shows an anterior view of the femur and an anterior view of the humerus. For the femur, the distal epiphysis contains a smaller lateral bulge and a larger medial bulge. These are examples of condyles. The inner halves of the two condyles as well as the groove between them compose a facet. An oval-shaped ridge on the medial surface of the distal metaphysis is an example of a tubercle. On the proximal epiphysis of the femur, the large knob that attaches to the hip socket is an example of a head. The tip of the head contains a small depression, an example of a fovea called the fovea capitis. On the humerus, the distal epiphysis contains a central depression that is an example of a fossa. Two condyles are located on the right and left sides of the fossa. The diaphysis of the humerus contains a small ridge running up the shaft that is an example of a tuberosity. The proximal epiphysis of the humerus contains a lateral and a medial bulge that are both examples of tubercles. Finally, a narrow groove runs from the center of the proximal metaphysis in between the medial and lateral condyles. This is an example of a sulcus. The middle image is entitled elevations or depressions. It shows an anterior view of the hip bones. The hip bones are shaped like two wings that join at the bottom. The crest along the upper edge of each hip bones, at the tip of each “wing” is an example of an elevation. A depression on the inner surface of both hip bones just under the crest is called out as a fossa. The right image is entitled examples of openings and shows an anterior view of the skull. The bone underlying the chin is an example of a protuberance while two small holes above each eye socket are examples of foramen. Five green sinuses surround the nose cavity are colored green. These are sinuses because they are hollowed out cavities within the skull bones. A small channel leads into the corner of each eye where the tear ducts occur. These two channels are both examples of a canal. Finally, the bones that form the posterior wall of the eye socket have a small crack running diagonally away from the nose. These are examples of fissures. Figure 4. Bone Features. The surface features of bones depend on their function, location, attachment of ligaments and tendons, or the penetration of blood vessels and nerves.[/caption]

Bone Cells and Tissue

Bone contains a relatively small number of cells entrenched in a matrix of collagen fibers that provide a surface for inorganic salt crystals to adhere. These salt crystals form when calcium phosphate and calcium carbonate combine to create hydroxyapatite, which incorporates other inorganic salts like magnesium hydroxide, fluoride, and sulfate as it crystallizes, or calcifies, on the collagen fibers. The hydroxyapatite crystals give bones their hardness and strength, while the collagen fibers give them flexibility so that they are not brittle.

Although bone cells compose a small amount of the bone volume, they are crucial to the function of bones. Four types of cells are found within bone tissue: osteoblasts, osteocytes, osteogenic cells, and osteoclasts (Figure 5).

[caption id="" align="aligncenter" width="395"]The top of this diagram shows the cross section of a generic bone with three zoom in boxes. The first box is on the periosteum. The second box is on the middle of the compact bone layer. The third box is on the inner edge of the compact bone where it transitions into the spongy bone. The callout in the periosteum points to two images. In the first image, four osteoblast cells are sitting end to end on the periosteum. The osteoblasts are roughly square shaped, except for one of the cells which is developing small, finger like projections. The caption says, “Osteoblasts form the matrix of the bone.” The second image called out from the periosteum shows a large, amorphous osteogenic cell sitting on the periosteum. The osteogenic cell is surrounded on both sides by a row of much smaller osteoblasts. The cell is shaped like a mushroom cap and also has finger like projections. The cell is a stem cell that develops into other bone cells. The box in the middle of the compact bone layer is pointing to an osteocyte. The osteocyte is a thin cell, roughly diamond shaped, with many branching, finger-like projections. The osteoctyes maintain bone tissue. The box at the inner edge of the compact bone is pointing to an osteoclast. The osteoclast is a large, round cell with multiple nuclei. It also has rows of fine finger like projections on its lower surface where it is sitting on the compact bone. The osteoclast reabsorbs bone. Figure 5. Bone Cells. Four types of cells are found within bone tissue. Osteogenic cells are undifferentiated and develop into osteoblasts. When osteoblasts get trapped within the calcified matrix, their structure and function changes, and they become osteocytes. Osteoclasts develop from monocytes and macrophages and differ in appearance from other bone cells.[/caption]

The osteoblast is the bone cell responsible for forming new bone and is found in the growing portions of bone, including the periosteum and endosteum. Osteoblasts, which do not divide, synthesize and secrete the collagen matrix and calcium salts. As the secreted matrix surrounding the osteoblast calcifies, the osteoblast become trapped within it; as a result, it changes in structure and becomes an osteocyte, the primary cell of mature bone and the most common type of bone cell. Each osteocyte is located in a space called a lacuna and is surrounded by bone tissue. Osteocytes maintain the mineral concentration of the matrix via the secretion of enzymes. Like osteoblasts, osteocytes lack mitotic activity. They can communicate with each other and receive nutrients via long cytoplasmic processes that extend through canaliculi (singular = canaliculus), channels within the bone matrix.

If osteoblasts and osteocytes are incapable of mitosis, then how are they replenished when old ones die? The answer lies in the properties of a third category of bone cells—the osteogenic cell. These osteogenic cells are undifferentiated with high mitotic activity and they are the only bone cells that divide. Immature osteogenic cells are found in the deep layers of the periosteum and the marrow. They differentiate and develop into osteoblasts.

The dynamic nature of bone means that new tissue is constantly formed, and old, injured, or unnecessary bone is dissolved for repair or for calcium release. The cell responsible for bone resorption, or breakdown, is the osteoclast. They are found on bone surfaces, are multinucleated, and originate from monocytes and macrophages, two types of white blood cells, not from osteogenic cells. Osteoclasts are continually breaking down old bone while osteoblasts are continually forming new bone. The ongoing balance between osteoblasts and osteoclasts is responsible for the constant but subtle reshaping of bone. Table 3 reviews the bone cells, their functions, and locations.
Bone Cells (Table 3)
Cell type Function Location
Osteogenic cells Develop into osteoblasts Deep layers of the periosteum and the marrow
Osteoblasts Bone formation Growing portions of bone, including periosteum and endosteum
Osteocytes Maintain mineral concentration of matrix Entrapped in matrix
Osteoclasts Bone resorption Bone surfaces and at sites of old, injured, or unneeded bone

Compact and Spongy Bone

The differences between compact and spongy bone are best explored via their histology. Most bones contain compact and spongy osseous tissue, but their distribution and concentration vary based on the bone’s overall function. Compact bone is dense so that it can withstand compressive forces, while spongy (cancellous) bone has open spaces and supports shifts in weight distribution.

Compact Bone

Compact bone is the denser, stronger of the two types of bone tissue (Figure 6). It can be found under the periosteum and in the diaphyses of long bones, where it provides support and protection.
[caption id="" align="aligncenter" width="480"]A generic long bone is shown at the top of this illustration. The bone is split in half lengthwise to show its internal anatomy. The outer gray covering of the bone is labeled the periosteum. Within the periosteum is a thin layer of compact bone. The compact bone surrounds a central cavity called the medullary cavity. The medullary cavity is filled with spongy bone at the two epiphyses. A callout box shows that the main image is zooming in on the compact bone on the left side of the bone. On the main image, the periosteum is being peeled back to show its two layers. The outer layer of the periosteum is the outer fibrous layer. This layer has a periosteal artery and a periosteal vein running along its outside edge. The inner layer of the periosteum is labeled the inner osteogenic layer. The compact bone lies to the right of the periosteum and occupies the majority of the main image. Two flat layers of compact bone line the inner surface of the ostegenic periosteum. These sheets of compact bone are called the circumferential lamellae. The majority of the compact bone has lamellae running perpendicular to that of the circumferential lamellae. These concentric lamellae are arranged in a series of concentric tubes. There are small cavities between the layers of concentric lamellae called lacunae. The centermost concentric lamella surrounds a hollow central canal. A blue vein, a red artery, a yellow nerve and a green lymph vessel run vertically through the central canal. A set of concentric lamellae, its associated lacunae and the vessels and nerves of the central canal are collectively called an osteon. The front edge of the diagram shows a longitudinal cross section of one of the osteons. The vessels and nerve are visible running through the center of the osteon throughout its length. In addition, blood vessels can run from the periosteum through the sides of the osteons and connect with the vessels of the central canal. The blood vessels travel through the sides of the osteons via a perforating canal. The open areas between neighboring osteons are also filled with compact bone. This “filler” bone is referred to as the interstitial lamellae. At the far right of the compact bone, the edge of the spongy bone is visible. The spongy bone is a series of crisscrossing bony arches called trabeculae. There are many open spaces between the trabeculae, giving the spongy bone its sponge-like appearance. Figure 6. Diagram of Compact Bone. (a) This cross-sectional view of compact bone shows the basic structural unit, the osteon. (b) In this micrograph of the osteon, you can clearly see the concentric lamellae and central canals. LM × 40. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

The microscopic structural unit of compact bone is called an osteon, or Haversian system. Each osteon is composed of concentric rings of calcified matrix called lamellae (singular = lamella). Running down the center of each osteon is the central canal, or Haversian canal, which contains blood vessels, nerves, and lymphatic vessels. These vessels and nerves branch off at right angles through a perforating canal, also known as Volkmann’s canals, to extend to the periosteum and endosteum.

The osteocytes are located inside spaces called lacunae (singular = lacuna), found at the borders of adjacent lamellae. As described earlier, canaliculi connect with the canaliculi of other lacunae and eventually with the central canal. This system allows nutrients to be transported to the osteocytes and wastes to be removed from them.

Spongy (Cancellous) Bone

Like compact bone, spongy bone, also known as cancellous bone, contains osteocytes housed in lacunae, but they are not arranged in concentric circles. Instead, the lacunae and osteocytes are found in a lattice-like network of matrix spikes called trabeculae (singular = trabecula) (Figure 7). The trabeculae may appear to be a random network, but each trabecula forms along lines of stress to provide strength to the bone. The spaces of the trabeculated network provide balance to the dense and heavy compact bone by making bones lighter so that muscles can move them more easily. In addition, the spaces in some spongy bones contain red marrow, protected by the trabeculae, where hematopoiesis occurs.

[caption id="" align="aligncenter" width="480"]This illustration shows the spongy bone within the proximal epiphysis of the femur in two successively magnified images. The lower-magnification image shows two layers of crisscrossing trabeculae. The surface of each is dotted with small black holes which are the openings of the canaliculi. One of the trabeculae is in a cross section to show its internal layers. The outermost covering of the lamellae is called the endosteum. This endosteum surrounds several layers of concentric lamellae. The higher-magnification image shows the cross section of the trabeculae more clearly. Three concentric lamellae are shown in this view, each possessing perpendicular black lines. These lines are the canaliculi and are oriented on the round lamellae similar to the spokes of a wheel. In between the lamellae are small cavities called lacunae which house cells called osteocytes. In addition, two large osteoclasts are seated on the outer edge of the outermost lamellae. The outermost lamellae are also surrounded by groups of small, white, osteoblasts. Figure 7. Diagram of Spongy Bone. Spongy bone is composed of trabeculae that contain the osteocytes. Red marrow fills the spaces in some bones.[/caption]
Aging and the…

Skeletal System: Paget’s Disease Paget’s disease usually occurs in adults over age 40. It is a disorder of the bone remodeling process that begins with overactive osteoclasts. This means more bone is resorbed than is laid down. The osteoblasts try to compensate but the new bone they lay down is weak and brittle and therefore prone to fracture.

While some people with Paget’s disease have no symptoms, others experience pain, bone fractures, and bone deformities (Figure 8). Bones of the pelvis, skull, spine, and legs are the most commonly affected. When occurring in the skull, Paget’s disease can cause headaches and hearing loss.
[caption id="" align="aligncenter" width="300"]This illustration shows the normal skeletal structure of the legs from an anterior view. The flesh of the legs and feet are outlined around the skeleton for reference. A second illustration shows the legs of someone with Paget’s disease. The affected person’s left femur is curved outward, causing the left leg to be bowed and shorter than the right leg. Figure 9. Paget's Disease. Normal leg bones are relatively straight, but those affected by Paget’s disease are porous and curved.[/caption]
What causes the osteoclasts to become overactive? The answer is still unknown, but hereditary factors seem to play a role. Some scientists believe Paget’s disease is due to an as-yet-unidentified virus. Paget’s disease is diagnosed via imaging studies and lab tests. X-rays may show bone deformities or areas of bone resorption. Bone scans are also useful. In these studies, a dye containing a radioactive ion is injected into the body. Areas of bone resorption have an affinity for the ion, so they will light up on the scan if the ions are absorbed. In addition, blood levels of an enzyme called alkaline phosphatase are typically elevated in people with Paget’s disease. Bisphosphonates, drugs that decrease the activity of osteoclasts, are often used in the treatment of Paget’s disease. However, in a small percentage of cases, bisphosphonates themselves have been linked to an increased risk of fractures because the old bone that is left after bisphosphonates are administered becomes worn out and brittle. Still, most doctors feel that the benefits of bisphosphonates more than outweigh the risk; the medical professional has to weigh the benefits and risks on a case-by-case basis. Bisphosphonate treatment can reduce the overall risk of deformities or fractures, which in turn reduces the risk of surgical repair and its associated risks and complications.

Blood and Nerve Supply

The spongy bone and medullary cavity receive nourishment from arteries that pass through the compact bone. The arteries enter through the nutrient foramen (plural = foramina), small openings in the diaphysis ([link]). The osteocytes in spongy bone are nourished by blood vessels of the periosteum that penetrate spongy bone and blood that circulates in the marrow cavities. As the blood passes through the marrow cavities, it is collected by veins, which then pass out of the bone through the foramina.

In addition to the blood vessels, nerves follow the same paths into the bone where they tend to concentrate in the more metabolically active regions of the bone. The nerves sense pain, and it appears the nerves also play roles in regulating blood supplies and in bone growth, hence their concentrations in metabolically active sites of the bone. [caption id="" align="aligncenter" width="280"]This illustration shows an anterior view if the right femur. The femur is split in half lengthwise to show its internal anatomy. The outer covering of the femur is labeled the periosteum. Within it is a thin layer of compact bone that surrounds a central cavity called the medullary or marrow cavity. This cavity is filled with spongy bone at both epiphyses. A nutrient artery and vein travels through the periosteum and compact bone at the center of the diaphysis. After entering the bone, the nutrient arteries and veins spread throughout the marrow cavity in both directions. Some of the arteries and veins in the marrow cavity also spread into the spongy bone within the distal and proximal epiphyses. However, additional blood vessels called the metaphyseal arteries and the metaphyseal veins enter into the metaphysis from outside of the bone. Figure 9. Diagram of Blood and Nerve Supply to Bone.Blood vessels and nerves enter the bone through the nutrient foramen.[/caption]

articular cartilage
thin layer of cartilage covering an epiphysis; reduces friction and acts as a shock absorber
articulation
where two bone surfaces meet
canaliculi
(singular = canaliculus) channels within the bone matrix that house one of an osteocyte’s many cytoplasmic extensions that it uses to communicate and receive nutrients
central canal
longitudinal channel in the center of each osteon; contains blood vessels, nerves, and lymphatic vessels; also known as the Haversian canal
compact bone
dense osseous tissue that can withstand compressive forces
diaphysis
tubular shaft that runs between the proximal and distal ends of a long bone
diploë
layer of spongy bone, that is sandwiched between two the layers of compact bone found in flat bones
endosteum
delicate membranous lining of a bone’s medullary cavity
epiphyseal plate
(also, growth plate) sheet of hyaline cartilage in the metaphysis of an immature bone; replaced by bone tissue as the organ grows in length
epiphysis
wide section at each end of a long bone; filled with spongy bone and red marrow
hole
opening or depression in a bone
lacunae
(singular = lacuna) spaces in a bone that house an osteocyte
medullary cavity
hollow region of the diaphysis; filled with yellow marrow
nutrient foramen
small opening in the middle of the external surface of the diaphysis, through which an artery enters the bone to provide nourishment
osteoblast
cell responsible for forming new bone
osteoclast
cell responsible for resorbing bone
osteocyte
primary cell in mature bone; responsible for maintaining the matrix
osteogenic cell
undifferentiated cell with high mitotic activity; the only bone cells that divide; they differentiate and develop into osteoblasts
osteon
(also, Haversian system) basic structural unit of compact bone; made of concentric layers of calcified matrix
perforating canal
(also, Volkmann’s canal) channel that branches off from the central canal and houses vessels and nerves that extend to the periosteum and endosteum
periosteum
fibrous membrane covering the outer surface of bone and continuous with ligaments
projection
bone markings where part of the surface sticks out above the rest of the surface, where tendons and ligaments attach
spongy bone
(also, cancellous bone) trabeculated osseous tissue that supports shifts in weight distribution
trabeculae
(singular = trabecula) spikes or sections of the lattice-like matrix in spongy bone
]]> 2043 2025 4 0 9.2 Fibrous Joints https://pressbooks.bccampus.ca/apdouglas2017/chapter/9-2-fibrous-joints/ Wed, 02 Aug 2017 21:58:31 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2117

Learning Objectives

By the end of this section, you will be able to:
  • Describe the structural features of fibrous joints
  • Distinguish between a suture, syndesmosis, and gomphosis
  • Give an example of each type of fibrous joint

At a fibrous joint, the adjacent bones are directly connected to each other by fibrous connective tissue, and thus the bones do not have a joint cavity between them (Figure 1). The gap between the bones may be narrow or wide. There are three types of fibrous joints. A suture is the narrow fibrous joint found between most bones of the skull. At a syndesmosis joint, the bones are more widely separated but are held together by a narrow band of fibrous connective tissue called a ligament or a wide sheet of connective tissue called an interosseous membrane. This type of fibrous joint is found between the shaft regions of the long bones in the forearm and in the leg. Lastly, a gomphosis is the narrow fibrous joint between the roots of a tooth and the bony socket in the jaw into which the tooth fits.

[caption id="" align="aligncenter" width="520"]This figure shows the different types of fibrous joints. The right panel shows sutures, the middle panel shows an interosseous membrane, and the left panel shows a gomphosis. Figure 1. Fibrous Joints. Fibrous joints form strong connections between bones. (a) Sutures join most bones of the skull. (b) An interosseous membrane forms a syndesmosis between the radius and ulna bones of the forearm. (c) A gomphosis is a specialized fibrous joint that anchors a tooth to its socket in the jaw.[/caption]

Suture

All the bones of the skull, except for the mandible, are joined to each other by a fibrous joint called a suture. The fibrous connective tissue found at a suture (“to bind or sew”) strongly unites the adjacent skull bones and thus helps to protect the brain and form the face. In adults, the skull bones are closely opposed and fibrous connective tissue fills the narrow gap between the bones. The suture is frequently convoluted, forming a tight union that prevents most movement between the bones. (See Figure 1a.) Thus, skull sutures are functionally classified as a synarthrosis, although some sutures may allow for slight movements between the cranial bones.

In newborns and infants, the areas of connective tissue between the bones are much wider, especially in those areas on the top and sides of the skull that will become the sagittal, coronal, squamous, and lambdoid sutures. These broad areas of connective tissue are called fontanelles (Figure 2). During birth, the fontanelles provide flexibility to the skull, allowing the bones to push closer together or to overlap slightly, thus aiding movement of the infant’s head through the birth canal. After birth, these expanded regions of connective tissue allow for rapid growth of the skull and enlargement of the brain. The fontanelles greatly decrease in width during the first year after birth as the skull bones enlarge. When the connective tissue between the adjacent bones is reduced to a narrow layer, these fibrous joints are now called sutures. At some sutures, the connective tissue will ossify and be converted into bone, causing the adjacent bones to fuse to each other. This fusion between bones is called a synostosis (“joined by bone”). Examples of synostosis fusions between cranial bones are found both early and late in life. At the time of birth, the frontal and maxillary bones consist of right and left halves joined together by sutures, which disappear by the eighth year as the halves fuse together to form a single bone. Late in life, the sagittal, coronal, and lambdoid sutures of the skull will begin to ossify and fuse, causing the suture line to gradually disappear.
[caption id="" align="aligncenter" width="380"]This figure shows the lateral view of the newborn skull with the major parts labeled. Figure 2. The Newborn Skull. The fontanelles of a newborn’s skull are broad areas of fibrous connective tissue that form fibrous joints between the bones of the skull.[/caption]

Syndesmosis

A syndesmosis (“fastened with a band”) is a type of fibrous joint in which two parallel bones are united to each other by fibrous connective tissue. The gap between the bones may be narrow, with the bones joined by ligaments, or the gap may be wide and filled in by a broad sheet of connective tissue called an interosseous membrane.

In the forearm, the wide gap between the shaft portions of the radius and ulna bones are strongly united by an interosseous membrane (see Figure 1b). Similarly, in the leg, the shafts of the tibia and fibula are also united by an interosseous membrane. In addition, at the distal tibiofibular joint, the articulating surfaces of the bones lack cartilage and the narrow gap between the bones is anchored by fibrous connective tissue and ligaments on both the anterior and posterior aspects of the joint. Together, the interosseous membrane and these ligaments form the tibiofibular syndesmosis.

The syndesmoses found in the forearm and leg serve to unite parallel bones and prevent their separation. However, a syndesmosis does not prevent all movement between the bones, and thus this type of fibrous joint is functionally classified as an amphiarthrosis. In the leg, the syndesmosis between the tibia and fibula strongly unites the bones, allows for little movement, and firmly locks the talus bone in place between the tibia and fibula at the ankle joint. This provides strength and stability to the leg and ankle, which are important during weight bearing. In the forearm, the interosseous membrane is flexible enough to allow for rotation of the radius bone during forearm movements. Thus in contrast to the stability provided by the tibiofibular syndesmosis, the flexibility of the antebrachial interosseous membrane allows for the much greater mobility of the forearm.

The interosseous membranes of the leg and forearm also provide areas for muscle attachment. Damage to a syndesmotic joint, which usually results from a fracture of the bone with an accompanying tear of the interosseous membrane, will produce pain, loss of stability of the bones, and may damage the muscles attached to the interosseous membrane. If the fracture site is not properly immobilized with a cast or splint, contractile activity by these muscles can cause improper alignment of the broken bones during healing.

Gomphosis

A gomphosis (“fastened with bolts”) is the specialized fibrous joint that anchors the root of a tooth into its bony socket within the maxillary bone (upper jaw) or mandible bone (lower jaw) of the skull. A gomphosis is also known as a peg-and-socket joint. Spanning between the bony walls of the socket and the root of the tooth are numerous short bands of dense connective tissue, each of which is called a periodontal ligament (see Figure 1c). Due to the immobility of a gomphosis, this type of joint is functionally classified as a synarthrosis.

Glossary

fontanelles
expanded areas of fibrous connective tissue that separate the braincase bones of the skull prior to birth and during the first year after birth
gomphosis
type of fibrous joint in which the root of a tooth is anchored into its bony jaw socket by strong periodontal ligaments
interosseous membrane
wide sheet of fibrous connective tissue that fills the gap between two parallel bones, forming a syndesmosis; found between the radius and ulna of the forearm and between the tibia and fibula of the leg
ligament
strong band of dense connective tissue spanning between bones
periodontal ligament
band of dense connective tissue that anchors the root of a tooth into the bony jaw socket
suture
fibrous joint that connects the bones of the skull (except the mandible); an immobile joint (synarthrosis)
syndesmosis
type of fibrous joint in which two separated, parallel bones are connected by an interosseous membrane
synostosis
site at which adjacent bones or bony components have fused together
]]> 2117 2108 3 0 11.2 Naming Skeletal Muscles https://pressbooks.bccampus.ca/apdouglas2017/chapter/11-2-naming-skeletal-muscles/ Wed, 02 Aug 2017 21:58:41 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2174

Learning Objectives

By the end of this section, you will be able to:
  • Describe the criteria used to name skeletal muscles
  • Explain how understanding the muscle names helps describe shapes, location, and actions of various muscles

The Greeks and Romans conducted the first studies done on the human body in Western culture. The educated class of subsequent societies studied Latin and Greek, and therefore the early pioneers of anatomy continued to apply Latin and Greek terminology or roots when they named the skeletal muscles. The large number of muscles in the body and unfamiliar words can make learning the names of the muscles in the body seem daunting, but understanding the etymology can help. Etymology is the study of how the root of a particular word entered a language and how the use of the word evolved over time. Taking the time to learn the root of the words is crucial to understanding the vocabulary of anatomy and physiology. When you understand the names of muscles it will help you remember where the muscles are located and what they do (Figure 1, Figure 2, and Table 2). Pronunciation of words and terms will take a bit of time to master, but after you have some basic information; the correct names and pronunciations will become easier.

[caption id="" align="aligncenter" width="380"]The top panel shows the anterior view of the human body with the major muscles labeled. The bottom panel shows the posterior view of the human body with the major muscles labeled. Figure 1. Overview of the Muscular System. On the anterior and posterior views of the muscular system above, superficial muscles (those at the surface) are shown on the right side of the body while deep muscles (those underneath the superficial muscles) are shown on the left half of the body. For the legs, superficial muscles are shown in the anterior view while the posterior view shows both superficial and deep muscles.[/caption]
[caption id="" align="aligncenter" width="520"]This table shows two examples of muscle names and how to translate them based on their Latin roots. The first row uses abductor digiti minimi as an example. The word abductor comes from the Latin roots ab, which means away from, and duct, which means to move. Therefore an abductor is a muscle that moves away from something. The word digiti comes from the Latin root digititus, which means digit and refers to a finger or toe. The word minimi comes from the Latin root minimus, which means minimum, tiny, or little. Therefore, the abductor digiti minimi is a muscle that moves the little finger or toe away. The second row uses the adductor digiti minimi as an example. The word adductor comes from the Latin root ad, which means to or toward, and duct, which means to move. Therefore an adductor is a muscle that moves toward something. As with the abductor digiti minimi, digiti refers to a finger or toe and minimi refers to something that is little. Therefore the adductor digiti minimi is a muscle that moves the little finger or toe forward. Figure 2. Understanding a Muscle Name from the Latin[/caption]
Mnemonic Device for Latin Roots (Table 2)
Example Latin or Greek Translation Mnemonic Device
ad to; toward ADvance toward your goal
ab away from n/a
sub under SUBmarines move under water.
ductor something that moves A conDUCTOR makes a train move.
anti against If you are antisocial, you are against engaging in social activities.
epi on top of n/a
apo to the side of n/a
longissimus longest “Longissimus” is longer than the word “long.”
longus long long
brevis short brief
maximus large max
medius medium “Medius” and “medium” both begin with “med.”
minimus tiny; little mini
rectus straight To RECTify a situation is to straighten it out.
multi many If something is MULTIcolored, it has many colors.
uni one A UNIcorn has one horn.
bi/di two If a ring is DIcast, it is made of two metals.
tri three TRIple the amount of money is three times as much.
quad four QUADruplets are four children born at one birth.
externus outside EXternal
internus inside INternal

Anatomists name the skeletal muscles according to a number of criteria, each of which describes the muscle in some way. These include naming the muscle after its shape, its size compared to other muscles in the area, its location in the body or the location of its attachments to the skeleton, how many origins it has, or its action.

The skeletal muscle’s anatomical location or its relationship to a particular bone often determines its name. For example, the frontalis muscle is located on top of the frontal bone of the skull. Similarly, the shapes of some muscles are very distinctive and the names, such as orbicularis, reflect the shape. For the buttocks, the size of the muscles influences the names: gluteus maximus (largest), gluteus medius (medium), and the gluteus minimus (smallest). Names were given to indicate length—brevis (short), longus (long)—and to identify position relative to the midline: lateralis (to the outside away from the midline), and medialis (toward the midline). The direction of the muscle fibers and fascicles are used to describe muscles relative to the midline, such as the rectus (straight) abdominis, or the oblique (at an angle) muscles of the abdomen.

Some muscle names indicate the number of muscles in a group. One example of this is the quadriceps, a group of four muscles located on the anterior (front) thigh. Other muscle names can provide information as to how many origins a particular muscle has, such as the biceps brachii. The prefix bi indicates that the muscle has two origins and tri indicates three origins.

The location of a muscle’s attachment can also appear in its name. When the name of a muscle is based on the attachments, the origin is always named first. For instance, the sternocleidomastoid muscle of the neck has a dual origin on the sternum (sterno) and clavicle (cleido), and it inserts on the mastoid process of the temporal bone. The last feature by which to name a muscle is its action. When muscles are named for the movement they produce, one can find action words in their name. Some examples are flexor (decreases the angle at the joint), extensor (increases the angle at the joint), abductor (moves the bone away from the midline), or adductor (moves the bone toward the midline).

]]> 2174 2162 3 0 18.6 Blood Typing https://pressbooks.bccampus.ca/apdouglas2017/chapter/18-6-blood-typing/ Wed, 02 Aug 2017 21:59:16 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2221

Learning Objectives

By the end of this section, you will be able to:
  • Describe the two basic physiological consequences of transfusion of incompatible blood
  • Compare and contrast ABO and Rh blood groups
  • Identify which blood groups may be safely transfused into patients with different ABO types
  • Discuss the pathophysiology of hemolytic disease of the newborn

Blood transfusions in humans were risky procedures until the discovery of the major human blood groups by Karl Landsteiner, an Austrian biologist and physician, in 1900. Until that point, physicians did not understand that death sometimes followed blood transfusions, when the type of donor blood infused into the patient was incompatible with the patient’s own blood. Blood groups are determined by the presence or absence of specific marker molecules on the plasma membranes of erythrocytes. With their discovery, it became possible for the first time to match patient-donor blood types and prevent transfusion reactions and deaths.

Antigens, Antibodies, and Transfusion Reactions

Antigens are substances that the body does not recognize as belonging to the “self” and that therefore trigger a defensive response from the leukocytes of the immune system. (Seek more content for additional information on immunity.) Here, we will focus on the role of immunity in blood transfusion reactions. With RBCs in particular, you may see the antigens referred to as isoantigens or agglutinogens (surface antigens) and the antibodies referred to as isoantibodies or agglutinins. In this chapter, we will use the more common terms antigens and antibodies.

Antigens are generally large proteins, but may include other classes of organic molecules, including carbohydrates, lipids, and nucleic acids. Following an infusion of incompatible blood, erythrocytes with foreign antigens appear in the bloodstream and trigger an immune response. Proteins called antibodies (immunoglobulins), which are produced by certain B lymphocytes called plasma cells, attach to the antigens on the plasma membranes of the infused erythrocytes and cause them to adhere to one another.

  • Because the arms of the Y-shaped antibodies attach randomly to more than one nonself erythrocyte surface, they form clumps of erythrocytes. This process is called agglutination.
  • The clumps of erythrocytes block small blood vessels throughout the body, depriving tissues of oxygen and nutrients.
  • As the erythrocyte clumps are degraded, in a process called hemolysis, their hemoglobin is released into the bloodstream. This hemoglobin travels to the kidneys, which are responsible for filtration of the blood. However, the load of hemoglobin released can easily overwhelm the kidney’s capacity to clear it, and the patient can quickly develop kidney failure.

More than 50 antigens have been identified on erythrocyte membranes, but the most significant in terms of their potential harm to patients are classified in two groups: the ABO blood group and the Rh blood group.

The ABO Blood Group

Although the ABO blood group name consists of three letters, ABO blood typing designates the presence or absence of just two antigens, A and B. Both are glycoproteins. People whose erythrocytes have A antigens on their erythrocyte membrane surfaces are designated blood type A, and those whose erythrocytes have B antigens are blood type B. People can also have both A and B antigens on their erythrocytes, in which case they are blood type AB. People with neither A nor B antigens are designated blood type O. ABO blood types are genetically determined.

Normally the body must be exposed to a foreign antigen before an antibody can be produced. This is not the case for the ABO blood group. Individuals with type A blood—without any prior exposure to incompatible blood—have preformed antibodies to the B antigen circulating in their blood plasma. These antibodies, referred to as anti-B antibodies, will cause agglutination and hemolysis if they ever encounter erythrocytes with B antigens. Similarly, an individual with type B blood has pre-formed anti-A antibodies. Individuals with type AB blood, which has both antigens, do not have preformed antibodies to either of these. People with type O blood lack antigens A and B on their erythrocytes, but both anti-A and anti-B antibodies circulate in their blood plasma.

Rh Blood Groups

The Rh blood group is classified according to the presence or absence of a second erythrocyte antigen identified as Rh. (It was first discovered in a type of primate known as a rhesus macaque, which is often used in research, because its blood is similar to that of humans.) Although dozens of Rh antigens have been identified, only one, designated D, is clinically important. Those who have the Rh D antigen present on their erythrocytes—about 85 percent of Americans—are described as Rh positive (Rh+) and those who lack it are Rh negative (Rh−). Note that the Rh group is distinct from the ABO group, so any individual, no matter their ABO blood type, may have or lack this Rh antigen. When identifying a patient’s blood type, the Rh group is designated by adding the word positive or negative to the ABO type. For example, A positive (A+) means ABO group A blood with the Rh antigen present, and AB negative (AB−) means ABO group AB blood without the Rh antigen.

Table 2 summarizes the distribution of the ABO and Rh blood types within the United States.

Summary of ABO and Rh Blood Types within the United States (Table 2)
Blood Type African-Americans Asian-Americans Caucasian-Americans Latino/Latina-Americans
A+ 24 27 33 29
A− 2 0.5 7 2
B+ 18 25 9 9
B− 1 0.4 2 1
AB+ 4 7 3 2
AB− 0.3 0.1 1 0.2
O+ 47 39 37 53
O− 4 1 8 4

In contrast to the ABO group antibodies, which are preformed, antibodies to the Rh antigen are produced only in Rh− individuals after exposure to the antigen. This process, called sensitization, occurs following a transfusion with Rh-incompatible blood or, more commonly, with the birth of an Rh+ baby to an Rh− mother. Problems are rare in a first pregnancy, since the baby’s Rh+ cells rarely cross the placenta (the organ of gas and nutrient exchange between the baby and the mother). However, during or immediately after birth, the Rh− mother can be exposed to the baby’s Rh+ cells (Figure 1). Research has shown that this occurs in about 13−14 percent of such pregnancies. After exposure, the mother’s immune system begins to generate anti-Rh antibodies. If the mother should then conceive another Rh+ baby, the Rh antibodies she has produced can cross the placenta into the fetal bloodstream and destroy the fetal RBCs. This condition, known as hemolytic disease of the newborn (HDN) or erythroblastosis fetalis, may cause anemia in mild cases, but the agglutination and hemolysis can be so severe that without treatment the fetus may die in the womb or shortly after birth.

[caption id="" align="aligncenter" width="600"]This figure shows an umbilical artery and vein passing through the placenta on the top left. The top right panel shows the first exposure to Rh+ antibodies in the mother. The bottom right panel shows the response when the second exposure in the form of another fetus takes place. Textboxes detail the steps in each process. Figure 1. Erythroblastosis Fetalis. The first exposure of an Rh− mother to Rh+ erythrocytes during pregnancy induces sensitization. Anti-Rh antibodies begin to circulate in the mother’s bloodstream. A second exposure occurs with a subsequent pregnancy with an Rh+ fetus in the uterus. Maternal anti-Rh antibodies may cross the placenta and enter the fetal bloodstream, causing agglutination and hemolysis of fetal erythrocytes.[/caption]

A drug known as RhoGAM, short for Rh immune globulin, can temporarily prevent the development of Rh antibodies in the Rh− mother, thereby averting this potentially serious disease for the fetus. RhoGAM antibodies destroy any fetal Rh+ erythrocytes that may cross the placental barrier. RhoGAM is normally administered to Rh− mothers during weeks 26−28 of pregnancy and within 72 hours following birth. It has proven remarkably effective in decreasing the incidence of HDN. Earlier we noted that the incidence of HDN in an Rh+ subsequent pregnancy to an Rh− mother is about 13–14 percent without preventive treatment. Since the introduction of RhoGAM in 1968, the incidence has dropped to about 0.1 percent in the United States.

Determining ABO Blood Types

Clinicians are able to determine a patient’s blood type quickly and easily using commercially prepared antibodies. An unknown blood sample is allocated into separate wells. Into one well a small amount of anti-A antibody is added, and to another a small amount of anti-B antibody. If the antigen is present, the antibodies will cause visible agglutination of the cells (Figure 2). The blood should also be tested for Rh antibodies.

[caption id="" align="aligncenter" width="430"]This figure shows three different red blood cells with different blood types. Figure 2. Cross Matching Blood Types. This sample of a commercially produced “bedside” card enables quick typing of both a recipient’s and donor’s blood before transfusion. The card contains three reaction sites or wells. One is coated with an anti-A antibody, one with an anti-B antibody, and one with an anti-D antibody (tests for the presence of Rh factor D). Mixing a drop of blood and saline into each well enables the blood to interact with a preparation of type-specific antibodies, also called anti-seras. Agglutination of RBCs in a given site indicates a positive identification of the blood antigens, in this case A and Rh antigens for blood type A+. For the purpose of transfusion, the donor’s and recipient’s blood types must match.[/caption]

ABO Transfusion Protocols

To avoid transfusion reactions, it is best to transfuse only matching blood types; that is, a type B+ recipient should ideally receive blood only from a type B+ donor and so on. That said, in emergency situations, when acute hemorrhage threatens the patient’s life, there may not be time for cross matching to identify blood type. In these cases, blood from a universal donor—an individual with type O− blood—may be transfused. Recall that type O erythrocytes do not display A or B antigens. Thus, anti-A or anti-B antibodies that might be circulating in the patient’s blood plasma will not encounter any erythrocyte surface antigens on the donated blood and therefore will not be provoked into a response. One problem with this designation of universal donor is if the O− individual had prior exposure to Rh antigen, Rh antibodies may be present in the donated blood. Also, introducing type O blood into an individual with type A, B, or AB blood will nevertheless introduce antibodies against both A and B antigens, as these are always circulating in the type O blood plasma. This may cause problems for the recipient, but because the volume of blood transfused is much lower than the volume of the patient’s own blood, the adverse effects of the relatively few infused plasma antibodies are typically limited. Rh factor also plays a role. If Rh− individuals receiving blood have had prior exposure to Rh antigen, antibodies for this antigen may be present in the blood and trigger agglutination to some degree. Although it is always preferable to cross match a patient’s blood before transfusing, in a true life-threatening emergency situation, this is not always possible, and these procedures may be implemented.

A patient with blood type AB+ is known as the universal recipient. This patient can theoretically receive any type of blood, because the patient’s own blood—having both A and B antigens on the erythrocyte surface—does not produce anti-A or anti-B antibodies. In addition, an Rh+ patient can receive both Rh+ and Rh− blood. However, keep in mind that the donor’s blood will contain circulating antibodies, again with possible negative implications. Figure 3 summarizes the blood types and compatibilities.

At the scene of multiple-vehicle accidents, military engagements, and natural or human-caused disasters, many victims may suffer simultaneously from acute hemorrhage, yet type O blood may not be immediately available. In these circumstances, medics may at least try to replace some of the volume of blood that has been lost. This is done by intravenous administration of a saline solution that provides fluids and electrolytes in proportions equivalent to those of normal blood plasma. Research is ongoing to develop a safe and effective artificial blood that would carry out the oxygen-carrying function of blood without the RBCs, enabling transfusions in the field without concern for incompatibility. These blood substitutes normally contain hemoglobin- as well as perfluorocarbon-based oxygen carriers.

[caption id="" align="aligncenter" width="420"]This table shows the different blood types, the antibodies in plasma, the antigens in the red blood cell, and the blood compatible blood types in an emergency. Figure 3. ABO Blood Group. This chart summarizes the characteristics of the blood types in the ABO blood group. See the text for more on the concept of a universal donor or recipient.[/caption]

]]> 2221 2194 7 0 19.4 Cardiac Physiology https://pressbooks.bccampus.ca/apdouglas2017/chapter/19-4-cardiac-physiology/ Wed, 02 Aug 2017 21:59:31 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2260

Learning Objectives

By the end of this section, you will be able to:
  • Relate heart rate to cardiac output
  • Describe the effect of exercise on heart rate
  • Identify cardiovascular centers and cardiac reflexes that regulate heart function
  • Describe factors affecting heart rate
  • Distinguish between positive and negative factors that affect heart contractility
  • Summarize factors affecting stroke volume and cardiac output
  • Describe the cardiac response to variations in blood flow and pressure

The autorhythmicity inherent in cardiac cells keeps the heart beating at a regular pace; however, the heart is regulated by and responds to outside influences as well. Neural and endocrine controls are vital to the regulation of cardiac function. In addition, the heart is sensitive to several environmental factors, including electrolytes.

Resting Cardiac Output

Cardiac output (CO) is a measurement of the amount of blood pumped by each ventricle in one minute. To calculate this value, multiply stroke volume (SV), the amount of blood pumped by each ventricle, by heart rate (HR), in contractions per minute (or beats per minute, bpm). It can be represented mathematically by the following equation:

CO = HR × SV

SV is normally measured using an echocardiogram to record EDV and ESV, and calculating the difference: SV = EDV – ESV. SV can also be measured using a specialized catheter, but this is an invasive procedure and far more dangerous to the patient. A mean SV for a resting 70-kg (150-lb) individual would be approximately 70 mL. There are several important variables, including size of the heart, physical and mental condition of the individual, sex, contractility, duration of contraction, preload or EDV, and afterload or resistance. Normal range for SV would be 55–100 mL. An average resting HR would be approximately 75 bpm but could range from 60–100 in some individuals.

Using these numbers, the mean CO is 5.25 L/min, with a range of 4.0–8.0 L/min. Remember, however, that these numbers refer to CO from each ventricle separately, not the total for the heart. Factors influencing CO are summarized in Figure 1.

[caption id="" align="aligncenter" width="450"]This figure lists the different factors affecting the heart rate and stroke volume. It also shows how they both affect the cardiac output. Figure 1. Major Factors Influencing Cardiac Output. Cardiac output is influenced by heart rate and stroke volume, both of which are also variable.[/caption]

SVs are also used to calculate ejection fraction, which is the portion of the blood that is pumped or ejected from the heart with each contraction. To calculate ejection fraction, SV is divided by EDV. Despite the name, the ejection fraction is normally expressed as a percentage. Ejection fractions range from approximately 55–70 percent, with a mean of 58 percent.

Exercise and Maximum Cardiac Output

In healthy young individuals, HR may increase to 150 bpm during exercise. SV can also increase from 70 to approximately 130 mL due to increased strength of contraction. This would increase CO to approximately 19.5 L/min, 4–5 times the resting rate. Top cardiovascular athletes can achieve even higher levels. At their peak performance, they may increase resting CO by 7–8 times.

Since the heart is a muscle, exercising it increases its efficiency. The difference between maximum and resting CO is known as the cardiac reserve. It measures the residual capacity of the heart to pump blood.

Heart Rates

HRs vary considerably, not only with exercise and fitness levels, but also with age. Newborn resting HRs may be 120 bpm. HR gradually decreases until young adulthood and then gradually increases again with age.

Maximum HRs are normally in the range of 200–220 bpm, although there are some extreme cases in which they may reach higher levels. As one ages, the ability to generate maximum rates decreases. This may be estimated by taking the maximal value of 220 bpm and subtracting the individual’s age. So a 40-year-old individual would be expected to hit a maximum rate of approximately 180, and a 60-year-old person would achieve a HR of 160.

Disorders of the…

Heart: Abnormal Heart Rates For an adult, normal resting HR will be in the range of 60–100 bpm. Bradycardia is the condition in which resting rate drops below 60 bpm, and tachycardia is the condition in which the resting rate is above 100 bpm. Trained athletes typically have very low HRs. If the patient is not exhibiting other symptoms, such as weakness, fatigue, dizziness, fainting, chest discomfort, palpitations, or respiratory distress, bradycardia is not considered clinically significant. However, if any of these symptoms are present, they may indicate that the heart is not providing sufficient oxygenated blood to the tissues. The term relative bradycardia may be used with a patient who has a HR in the normal range but is still suffering from these symptoms. Most patients remain asymptomatic as long as the HR remains above 50 bpm.

Bradycardia may be caused by either inherent factors or causes external to the heart. While the condition may be inherited, typically it is acquired in older individuals. Inherent causes include abnormalities in either the SA or AV node. If the condition is serious, a pacemaker may be required. Other causes include ischemia to the heart muscle or diseases of the heart vessels or valves. External causes include metabolic disorders, pathologies of the endocrine system often involving the thyroid, electrolyte imbalances, neurological disorders including inappropriate autonomic responses, autoimmune pathologies, over-prescription of beta blocker drugs that reduce HR, recreational drug use, or even prolonged bed rest. Treatment relies upon establishing the underlying cause of the disorder and may necessitate supplemental oxygen.

Tachycardia is not normal in a resting patient but may be detected in pregnant women or individuals experiencing extreme stress. In the latter case, it would likely be triggered by stimulation from the limbic system or disorders of the autonomic nervous system. In some cases, tachycardia may involve only the atria. Some individuals may remain asymptomatic, but when present, symptoms may include dizziness, shortness of breath, lightheadedness, rapid pulse, heart palpations, chest pain, or fainting (syncope). While tachycardia is defined as a HR above 100 bpm, there is considerable variation among people. Further, the normal resting HRs of children are often above 100 bpm, but this is not considered to be tachycardia Many causes of tachycardia may be benign, but the condition may also be correlated with fever, anemia, hypoxia, hyperthyroidism, hypersecretion of catecholamines, some cardiomyopathies, some disorders of the valves, and acute exposure to radiation. Elevated rates in an exercising or resting patient are normal and expected. Resting rate should always be taken after recovery from exercise. Treatment depends upon the underlying cause but may include medications, implantable cardioverter defibrillators, ablation, or surgery.

Correlation Between Heart Rates and Cardiac Output

Initially, physiological conditions that cause HR to increase also trigger an increase in SV. During exercise, the rate of blood returning to the heart increases. However as the HR rises, there is less time spent in diastole and consequently less time for the ventricles to fill with blood. Even though there is less filling time, SV will initially remain high. However, as HR continues to increase, SV gradually decreases due to decreased filling time. CO will initially stabilize as the increasing HR compensates for the decreasing SV, but at very high rates, CO will eventually decrease as increasing rates are no longer able to compensate for the decreasing SV. Consider this phenomenon in a healthy young individual. Initially, as HR increases from resting to approximately 120 bpm, CO will rise. As HR increases from 120 to 160 bpm, CO remains stable, since the increase in rate is offset by decreasing ventricular filling time and, consequently, SV. As HR continues to rise above 160 bpm, CO actually decreases as SV falls faster than HR increases. So although aerobic exercises are critical to maintain the health of the heart, individuals are cautioned to monitor their HR to ensure they stay within the target heart rate range of between 120 and 160 bpm, so CO is maintained. The target HR is loosely defined as the range in which both the heart and lungs receive the maximum benefit from the aerobic workout and is dependent upon age.

Cardiovascular Centers

Nervous control over HR is centralized within the two paired cardiovascular centers of the medulla oblongata (Figure 2). The cardioaccelerator regions stimulate activity via sympathetic stimulation of the cardioaccelerator nerves, and the cardioinhibitory centers decrease heart activity via parasympathetic stimulation as one component of the vagus nerve, cranial nerve X. During rest, both centers provide slight stimulation to the heart, contributing to autonomic tone. This is a similar concept to tone in skeletal muscles. Normally, vagal stimulation predominates as, left unregulated, the SA node would initiate a sinus rhythm of approximately 100 bpm.

Both sympathetic and parasympathetic stimulations flow through a paired complex network of nerve fibers known as the cardiac plexus near the base of the heart. The cardioaccelerator center also sends additional fibers, forming the cardiac nerves via sympathetic ganglia (the cervical ganglia plus superior thoracic ganglia T1–T4) to both the SA and AV nodes, plus additional fibers to the atria and ventricles. The ventricles are more richly innervated by sympathetic fibers than parasympathetic fibers. Sympathetic stimulation causes the release of the neurotransmitter norepinephrine (NE) at the neuromuscular junction of the cardiac nerves. NE shortens the repolarization period, thus speeding the rate of depolarization and contraction, which results in an increase in HR. It opens chemical- or ligand-gated sodium and calcium ion channels, allowing an influx of positively charged ions.

NE binds to the beta-1 receptor. Some cardiac medications (for example, beta blockers) work by blocking these receptors, thereby slowing HR and are one possible treatment for hypertension. Overprescription of these drugs may lead to bradycardia and even stoppage of the heart.

[caption id="" align="aligncenter" width="280"]This figure shows the brain and the nerves connecting the brain to the heart. Figure 2. Autonomic Innervation of the Heart. Cardioaccelerator and cardioinhibitory areas are components of the paired cardiac centers located in the medulla oblongata of the brain. They innervate the heart via sympathetic cardiac nerves that increase cardiac activity and vagus (parasympathetic) nerves that slow cardiac activity.[/caption]

Parasympathetic stimulation originates from the cardioinhibitory region with impulses traveling via the vagus nerve (cranial nerve X). The vagus nerve sends branches to both the SA and AV nodes, and to portions of both the atria and ventricles. Parasympathetic stimulation releases the neurotransmitter acetylcholine (ACh) at the neuromuscular junction. ACh slows HR by opening chemical- or ligand-gated potassium ion channels to slow the rate of spontaneous depolarization, which extends repolarization and increases the time before the next spontaneous depolarization occurs. Without any nervous stimulation, the SA node would establish a sinus rhythm of approximately 100 bpm. Since resting rates are considerably less than this, it becomes evident that parasympathetic stimulation normally slows HR. This is similar to an individual driving a car with one foot on the brake pedal. To speed up, one need merely remove one’s foot from the break and let the engine increase speed. In the case of the heart, decreasing parasympathetic stimulation decreases the release of ACh, which allows HR to increase up to approximately 100 bpm. Any increases beyond this rate would require sympathetic stimulation. Figure 3 illustrates the effects of parasympathetic and sympathetic stimulation on the normal sinus rhythm.

[caption id="" align="aligncenter" width="350"]This figure shows three graphs. The top panel shows the normal or resting potential with time. The middle panel shows membrane potential with time in a parasympathetic stimulation where the heart rate is decreased. The bottom panel shows membrane potential with time in a sympathetic stimulation with increased heart rate. Figure 3. Effects of Parasympathetic and Sympathetic Stimulation on Normal Sinus Rhythm. The wave of depolarization in a normal sinus rhythm shows a stable resting HR. Following parasympathetic stimulation, HR slows. Following sympathetic stimulation, HR increases.[/caption]

Input to the Cardiovascular Center

The cardiovascular center receives input from a series of visceral receptors with impulses traveling through visceral sensory fibers within the vagus and sympathetic nerves via the cardiac plexus. Among these receptors are various proprioreceptors, baroreceptors, and chemoreceptors, plus stimuli from the limbic system. Collectively, these inputs normally enable the cardiovascular centers to regulate heart function precisely, a process known as cardiac reflexes. Increased physical activity results in increased rates of firing by various proprioreceptors located in muscles, joint capsules, and tendons. Any such increase in physical activity would logically warrant increased blood flow. The cardiac centers monitor these increased rates of firing, and suppress parasympathetic stimulation and increase sympathetic stimulation as needed in order to increase blood flow.

Similarly, baroreceptors are stretch receptors located in the aortic sinus, carotid bodies, the venae cavae, and other locations, including pulmonary vessels and the right side of the heart itself. Rates of firing from the baroreceptors represent blood pressure, level of physical activity, and the relative distribution of blood. The cardiac centers monitor baroreceptor firing to maintain cardiac homeostasis, a mechanism called the baroreceptor reflex. With increased pressure and stretch, the rate of baroreceptor firing increases, and the cardiac centers decrease sympathetic stimulation and increase parasympathetic stimulation. As pressure and stretch decrease, the rate of baroreceptor firing decreases, and the cardiac centers increase sympathetic stimulation and decrease parasympathetic stimulation.

There is a similar reflex, called the atrial reflex or Bainbridge reflex, associated with varying rates of blood flow to the atria. Increased venous return stretches the walls of the atria where specialized baroreceptors are located. However, as the atrial baroreceptors increase their rate of firing and as they stretch due to the increased blood pressure, the cardiac center responds by increasing sympathetic stimulation and inhibiting parasympathetic stimulation to increase HR. The opposite is also true.

Increased metabolic byproducts associated with increased activity, such as carbon dioxide, hydrogen ions, and lactic acid, plus falling oxygen levels, are detected by a suite of chemoreceptors innervated by the glossopharyngeal and vagus nerves. These chemoreceptors provide feedback to the cardiovascular centers about the need for increased or decreased blood flow, based on the relative levels of these substances.

The limbic system can also significantly impact HR related to emotional state. During periods of stress, it is not unusual to identify higher than normal HRs, often accompanied by a surge in the stress hormone cortisol. Individuals experiencing extreme anxiety may manifest panic attacks with symptoms that resemble those of heart attacks. These events are typically transient and treatable. Meditation techniques have been developed to ease anxiety and have been shown to lower HR effectively. Doing simple deep and slow breathing exercises with one’s eyes closed can also significantly reduce this anxiety and HR.

Disorders of the…

Heart: Broken Heart Syndrome

Extreme stress from such life events as the death of a loved one, an emotional break up, loss of income, or foreclosure of a home may lead to a condition commonly referred to as broken heart syndrome. This condition may also be called Takotsubo cardiomyopathy, transient apical ballooning syndrome, apical ballooning cardiomyopathy, stress-induced cardiomyopathy, Gebrochenes-Herz syndrome, and stress cardiomyopathy. The recognized effects on the heart include congestive heart failure due to a profound weakening of the myocardium not related to lack of oxygen. This may lead to acute heart failure, lethal arrhythmias, or even the rupture of a ventricle. The exact etiology is not known, but several factors have been suggested, including transient vasospasm, dysfunction of the cardiac capillaries, or thickening of the myocardium—particularly in the left ventricle—that may lead to the critical circulation of blood to this region. While many patients survive the initial acute event with treatment to restore normal function, there is a strong correlation with death. Careful statistical analysis by the Cass Business School, a prestigious institution located in London, published in 2008, revealed that within one year of the death of a loved one, women are more than twice as likely to die and males are six times as likely to die as would otherwise be expected.

Other Factors Influencing Heart Rate

Using a combination of autorhythmicity and innervation, the cardiovascular center is able to provide relatively precise control over HR. However, there are a number of other factors that have an impact on HR as well, including epinephrine, NE, and thyroid hormones; levels of various ions including calcium, potassium, and sodium; body temperature; hypoxia; and pH balance (Table 1 and Table 2). After reading this section, the importance of maintaining homeostasis should become even more apparent.

Major Factors Increasing Heart Rate and Force of Contraction (Table 1)
Factor Effect
Cardioaccelerator nerves Release of norepinephrine
Proprioreceptors Increased rates of firing during exercise
Chemoreceptors Decreased levels of O2; increased levels of H+, CO2, and lactic acid
Baroreceptors Decreased rates of firing, indicating falling blood volume/pressure
Limbic system Anticipation of physical exercise or strong emotions
Catecholamines Increased epinephrine and norepinephrine
Thyroid hormones Increased T3 and T4
Calcium Increased Ca2+
Potassium Decreased K+
Sodium Decreased Na+
Body temperature Increased body temperature
Nicotine and caffeine Stimulants, increasing heart rate
Factors Decreasing Heart Rate and Force of Contraction (Table 2)
Factor Effect
Cardioinhibitor nerves (vagus) Release of acetylcholine
Proprioreceptors Decreased rates of firing following exercise
Chemoreceptors Increased levels of O2; decreased levels of H+ and CO2
Baroreceptors Increased rates of firing, indicating higher blood volume/pressure
Limbic system Anticipation of relaxation
Catecholamines Decreased epinephrine and norepinephrine
Thyroid hormones Decreased T3 and T4
Calcium Decreased Ca2+
Potassium Increased K+
Sodium Increased Na+
Body temperature Decrease in body temperature

Epinephrine and Norepinephrine

The catecholamines, epinephrine and NE, secreted by the adrenal medulla form one component of the extended fight-or-flight mechanism. The other component is sympathetic stimulation. Epinephrine and NE have similar effects: binding to the beta-1 receptors, and opening sodium and calcium ion chemical- or ligand-gated channels. The rate of depolarization is increased by this additional influx of positively charged ions, so the threshold is reached more quickly and the period of repolarization is shortened. However, massive releases of these hormones coupled with sympathetic stimulation may actually lead to arrhythmias. There is no parasympathetic stimulation to the adrenal medulla.

Thyroid Hormones

In general, increased levels of thyroid hormone, or thyroxin, increase cardiac rate and contractility. The impact of thyroid hormone is typically of a much longer duration than that of the catecholamines. The physiologically active form of thyroid hormone, T3 or triiodothyronine, has been shown to directly enter cardiomyocytes and alter activity at the level of the genome. It also impacts the beta adrenergic response similar to epinephrine and NE described above. Excessive levels of thyroxin may trigger tachycardia.

Calcium

Calcium ion levels have great impacts upon both HR and contractility; as the levels of calcium ions increase, so do HR and contractility. High levels of calcium ions (hypercalcemia) may be implicated in a short QT interval and a widened T wave in the ECG. The QT interval represents the time from the start of depolarization to repolarization of the ventricles, and includes the period of ventricular systole. Extremely high levels of calcium may induce cardiac arrest. Drugs known as calcium channel blockers slow HR by binding to these channels and blocking or slowing the inward movement of calcium ions.

Caffeine and Nicotine

Caffeine and nicotine are not found naturally within the body. Both of these nonregulated drugs have an excitatory effect on membranes of neurons in general and have a stimulatory effect on the cardiac centers specifically, causing an increase in HR. Caffeine works by increasing the rates of depolarization at the SA node, whereas nicotine stimulates the activity of the sympathetic neurons that deliver impulses to the heart.

Although it is the world’s most widely consumed psychoactive drug, caffeine is legal and not regulated. While precise quantities have not been established, “normal” consumption is not considered harmful to most people, although it may cause disruptions to sleep and acts as a diuretic. Its consumption by pregnant women is cautioned against, although no evidence of negative effects has been confirmed. Tolerance and even physical and mental addiction to the drug result in individuals who routinely consume the substance.

Nicotine, too, is a stimulant and produces addiction. While legal and nonregulated, concerns about nicotine’s safety and documented links to respiratory and cardiac disease have resulted in warning labels on cigarette packages.

Factors Decreasing Heart Rate

HR can be slowed when a person experiences altered sodium and potassium levels, hypoxia, acidosis, alkalosis, and hypothermia (see Table 1). The relationship between electrolytes and HR is complex, but maintaining electrolyte balance is critical to the normal wave of depolarization. Of the two ions, potassium has the greater clinical significance. Initially, both hyponatremia (low sodium levels) and hypernatremia (high sodium levels) may lead to tachycardia. Severely high hypernatremia may lead to fibrillation, which may cause CO to cease. Severe hyponatremia leads to both bradycardia and other arrhythmias. Hypokalemia (low potassium levels) also leads to arrhythmias, whereas hyperkalemia (high potassium levels) causes the heart to become weak and flaccid, and ultimately to fail.

Heart muscle relies exclusively on aerobic metabolism for energy. Hypoxia (an insufficient supply of oxygen) leads to decreasing HRs, since metabolic reactions fueling heart contraction are restricted.

Acidosis is a condition in which excess hydrogen ions are present, and the patient’s blood expresses a low pH value. Alkalosis is a condition in which there are too few hydrogen ions, and the patient’s blood has an elevated pH. Normal blood pH falls in the range of 7.35–7.45, so a number lower than this range represents acidosis and a higher number represents alkalosis. Recall that enzymes are the regulators or catalysts of virtually all biochemical reactions; they are sensitive to pH and will change shape slightly with values outside their normal range. These variations in pH and accompanying slight physical changes to the active site on the enzyme decrease the rate of formation of the enzyme-substrate complex, subsequently decreasing the rate of many enzymatic reactions, which can have complex effects on HR. Severe changes in pH will lead to denaturation of the enzyme.

The last variable is body temperature. Elevated body temperature is called hyperthermia, and suppressed body temperature is called hypothermia. Slight hyperthermia results in increasing HR and strength of contraction. Hypothermia slows the rate and strength of heart contractions. This distinct slowing of the heart is one component of the larger diving reflex that diverts blood to essential organs while submerged. If sufficiently chilled, the heart will stop beating, a technique that may be employed during open heart surgery. In this case, the patient’s blood is normally diverted to an artificial heart-lung machine to maintain the body’s blood supply and gas exchange until the surgery is complete, and sinus rhythm can be restored. Excessive hyperthermia and hypothermia will both result in death, as enzymes drive the body systems to cease normal function, beginning with the central nervous system.

Stroke Volume

Many of the same factors that regulate HR also impact cardiac function by altering SV. While a number of variables are involved, SV is ultimately dependent upon the difference between EDV and ESV. The three primary factors to consider are preload, or the stretch on the ventricles prior to contraction; the contractility, or the force or strength of the contraction itself; and afterload, the force the ventricles must generate to pump blood against the resistance in the vessels. These factors are summarized in Table 1 and Table 2.

Preload

Preload is another way of expressing EDV. Therefore, the greater the EDV is, the greater the preload is. One of the primary factors to consider is filling time, or the duration of ventricular diastole during which filling occurs. The more rapidly the heart contracts, the shorter the filling time becomes, and the lower the EDV and preload are. This effect can be partially overcome by increasing the second variable, contractility, and raising SV, but over time, the heart is unable to compensate for decreased filling time, and preload also decreases.

With increasing ventricular filling, both EDV or preload increase, and the cardiac muscle itself is stretched to a greater degree. At rest, there is little stretch of the ventricular muscle, and the sarcomeres remain short. With increased ventricular filling, the ventricular muscle is increasingly stretched and the sarcomere length increases. As the sarcomeres reach their optimal lengths, they will contract more powerfully, because more of the myosin heads can bind to the actin on the thin filaments, forming cross bridges and increasing the strength of contraction and SV. If this process were to continue and the sarcomeres stretched beyond their optimal lengths, the force of contraction would decrease. However, due to the physical constraints of the location of the heart, this excessive stretch is not a concern.

The relationship between ventricular stretch and contraction has been stated in the well-known Frank-Starling mechanism or simply Starling’s Law of the Heart. This principle states that, within physiological limits, the force of heart contraction is directly proportional to the initial length of the muscle fiber. This means that the greater the stretch of the ventricular muscle (within limits), the more powerful the contraction is, which in turn increases SV. Therefore, by increasing preload, you increase the second variable, contractility.

Otto Frank (1865–1944) was a German physiologist; among his many published works are detailed studies of this important heart relationship. Ernest Starling (1866–1927) was an important English physiologist who also studied the heart. Although they worked largely independently, their combined efforts and similar conclusions have been recognized in the name “Frank-Starling mechanism.”

Any sympathetic stimulation to the venous system will increase venous return to the heart, which contributes to ventricular filling, and EDV and preload. While much of the ventricular filling occurs while both atria and ventricles are in diastole, the contraction of the atria, the atrial kick, plays a crucial role by providing the last 20–30 percent of ventricular filling.

Contractility

It is virtually impossible to consider preload or ESV without including an early mention of the concept of contractility. Indeed, the two parameters are intimately linked. Contractility refers to the force of the contraction of the heart muscle, which controls SV, and is the primary parameter for impacting ESV. The more forceful the contraction is, the greater the SV and smaller the ESV are. Less forceful contractions result in smaller SVs and larger ESVs. Factors that increase contractility are described as positive inotropic factors, and those that decrease contractility are described as negative inotropic factors (ino- = “fiber;” -tropic = “turning toward”).

Not surprisingly, sympathetic stimulation is a positive inotrope, whereas parasympathetic stimulation is a negative inotrope. Sympathetic stimulation triggers the release of NE at the neuromuscular junction from the cardiac nerves and also stimulates the adrenal cortex to secrete epinephrine and NE. In addition to their stimulatory effects on HR, they also bind to both alpha and beta receptors on the cardiac muscle cell membrane to increase metabolic rate and the force of contraction. This combination of actions has the net effect of increasing SV and leaving a smaller residual ESV in the ventricles. In comparison, parasympathetic stimulation releases ACh at the neuromuscular junction from the vagus nerve. The membrane hyperpolarizes and inhibits contraction to decrease the strength of contraction and SV, and to raise ESV. Since parasympathetic fibers are more widespread in the atria than in the ventricles, the primary site of action is in the upper chambers. Parasympathetic stimulation in the atria decreases the atrial kick and reduces EDV, which decreases ventricular stretch and preload, thereby further limiting the force of ventricular contraction. Stronger parasympathetic stimulation also directly decreases the force of contraction of the ventricles.

Several synthetic drugs, including dopamine and isoproterenol, have been developed that mimic the effects of epinephrine and NE by stimulating the influx of calcium ions from the extracellular fluid. Higher concentrations of intracellular calcium ions increase the strength of contraction. Excess calcium (hypercalcemia) also acts as a positive inotropic agent. The drug digitalis lowers HR and increases the strength of the contraction, acting as a positive inotropic agent by blocking the sequestering of calcium ions into the sarcoplasmic reticulum. This leads to higher intracellular calcium levels and greater strength of contraction. In addition to the catecholamines from the adrenal medulla, other hormones also demonstrate positive inotropic effects. These include thyroid hormones and glucagon from the pancreas.

Negative inotropic agents include hypoxia, acidosis, hyperkalemia, and a variety of synthetic drugs. These include numerous beta blockers and calcium channel blockers. Early beta blocker drugs include propranolol and pronethalol, and are credited with revolutionizing treatment of cardiac patients experiencing angina pectoris. There is also a large class of dihydropyridine, phenylalkylamine, and benzothiazepine calcium channel blockers that may be administered decreasing the strength of contraction and SV.

Afterload

Afterload refers to the tension that the ventricles must develop to pump blood effectively against the resistance in the vascular system. Any condition that increases resistance requires a greater afterload to force open the semilunar valves and pump the blood. Damage to the valves, such as stenosis, which makes them harder to open will also increase afterload. Any decrease in resistance decreases the afterload. Figure 4 summarizes the major factors influencing SV, Figure 5 summarizes the major factors influencing CO, and Table 3 and Table 4 summarize cardiac responses to increased and decreased blood flow and pressure in order to restore homeostasis.

[caption id="" align="aligncenter" width="550"]This table describes major factors influencing stroke volume. Preload may be raised due to fast filling time or increased venous return. These factors increase end diastolic volume and increase stroke volume. Preload may be lowered due to decreased thyroid hormones, decreased calcium ions, high or low potassium ions, high or low sodium, low body temperature, hypoxia, abnormal pH balance, or drugs (for example, calcium channel blockers). These factors decrease end diastolic volume and decrease stroke volume. Contractility may be raised due to sympathetic stimulation, epinephrine and norepinephrine, high intracellular calcium ions, high blood calcium level, thyroid hormones, or glucagon. These factors decrease end systolic volume and increase stroke volume. Contractility may be lowered due to parasympathetic stimulation, acetylcholine, hypoxia, or hyperkalemia. These factors increase end systolic volume and decrease stroke volume. Afterload may be raised due to increased vascular resistance or semilunar valve damage. These factors increase end systolic volume and decrease stroke volume. Afterload may be lowered due to decreased vascular resistance. This factor decreases end systolic volume and increases stroke volume. Figure 4. Major Factors Influencing Stroke Volume. Multiple factors impact preload, afterload, and contractility, and are the major considerations influencing SV.[/caption]
[caption id="" align="aligncenter" width="550"]This flowchart lists all the important factors that affect cardiac output. Figure 5. Summary of Major Factors Influencing Cardiac Output. The primary factors influencing HR include autonomic innervation plus endocrine control. Not shown are environmental factors, such as electrolytes, metabolic products, and temperature. The primary factors controlling SV include preload, contractility, and afterload. Other factors such as electrolytes may be classified as either positive or negative inotropic agents.[/caption]
Cardiac Response to Decreasing Blood Flow and Pressure Due to Decreasing Cardiac Output (Table 3)
Baroreceptors (aorta, carotid arteries, venae cavae, and atria) Chemoreceptors (both central nervous system and in proximity to baroreceptors)
Sensitive to Decreasing stretch Decreasing O2 and increasing CO2, H+, and lactic acid
Target Parasympathetic stimulation suppressed Sympathetic stimulation increased
Response of heart Increasing heart rate and increasing stroke volume Increasing heart rate and increasing stroke volume
Overall effect Increasing blood flow and pressure due to increasing cardiac output; hemostasis restored Increasing blood flow and pressure due to increasing cardiac output; hemostasis restored
Cardiac Response to Increasing Blood Flow and Pressure Due to Increasing Cardiac Output (Table 4)
Baroreceptors (aorta, carotid arteries, venae cavae, and atria) Chemoreceptors (both central nervous system and in proximity to baroreceptors)
Sensitive to Increasing stretch Increasing O2 and decreasing CO2, H+, and lactic acid
Target Parasympathetic stimulation increased Sympathetic stimulation suppressed
Response of heart Decreasing heart rate and decreasing stroke volume Decreasing heart rate and decreasing stroke volume
Overall effect Decreasing blood flow and pressure due to decreasing cardiac output; hemostasis restored Decreasing blood flow and pressure due to decreasing cardiac output; hemostasis restored

]]> 2260 2222 5 0 26.1 Body Fluids and Fluid Compartments https://pressbooks.bccampus.ca/apdouglas2017/chapter/26-1-body-fluids-and-fluid-compartments/ Wed, 02 Aug 2017 22:00:41 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2404

Learning Objectives

By the end of this section, you will be able to:
  • Explain the importance of water in the body
  • Contrast the composition of the intracellular fluid with that of the extracellular fluid
  • Explain the importance of protein channels in the movement of solutes
  • Identify the causes and symptoms of edema

The chemical reactions of life take place in aqueous solutions. The dissolved substances in a solution are called solutes. In the human body, solutes vary in different parts of the body, but may include proteins—including those that transport lipids, carbohydrates, and, very importantly, electrolytes. Often in medicine, a mineral dissociated from a salt that carries an electrical charge (an ion) is called and electrolyte. For instance, sodium ions (Na+) and chloride ions (Cl-) are often referred to as electrolytes.

In the body, water moves through semi-permeable membranes of cells and from one compartment of the body to another by a process called osmosis. Osmosis is basically the diffusion of water from regions of higher concentration to regions of lower concentration, along an osmotic gradient across a semi-permeable membrane. As a result, water will move into and out of cells and tissues, depending on the relative concentrations of the water and solutes found there. An appropriate balance of solutes inside and outside of cells must be maintained to ensure normal function.

Body Water Content

Human beings are mostly water, ranging from about 75 percent of body mass in infants to about 50–60 percent in adult men and women, to as low as 45 percent in old age. The percent of body water changes with development, because the proportions of the body given over to each organ and to muscles, fat, bone, and other tissues change from infancy to adulthood (Figure 1). Your brain and kidneys have the highest proportions of water, which composes 80–85 percent of their masses. In contrast, teeth have the lowest proportion of water, at 8–10 percent.

[caption id="" align="aligncenter" width="450"]This illustration shows a silhouette of a human body with various organs highlighted. The percent of water contained in each organ is given. The brain typically contains 80% to 85% water, teeth contain 8% to 10% water, a single lung contains 75% to 80% water, the heart contains 75% to 80% water, the bones contain 20% to 25% water, the liver contains 70% to 75% water, the kidneys contain 80% to 85% water, the skin contains 70% to 75% water and the muscles also contain 70% to 75% water. Figure 1. Water Content of the Body’s Organs and Tissues. Water content varies in different body organs and tissues, from as little as 8 percent in the teeth to as much as 85 percent in the brain.[/caption]

Fluid Compartments

Body fluids can be discussed in terms of their specific fluid compartment, a location that is largely separate from another compartment by some form of a physical barrier. The intracellular fluid (ICF) compartment is the system that includes all fluid enclosed in cells by their plasma membranes. Extracellular fluid (ECF) surrounds all cells in the body. Extracellular fluid has two primary constituents: the fluid component of the blood (called plasma) and the interstitial fluid (IF) that surrounds all cells not in the blood (Figure 2).

[caption id="" align="aligncenter" width="380"]This diagram shows a small blood vessel surrounded by several body cells. The fluid between the body cells is the interstitial fluid (IF), which is a type of extracellular fluid (ECF). The fluid in the blood vessel is also an example of extracellular fluid. The fluid in the cytoplasm of each body cell is intracellular fluid, or ICF. Figure 2. Fluid Compartments in the Human Body. The intracellular fluid (ICF) is the fluid within cells. The interstitial fluid (IF) is part of the extracellular fluid (ECF) between the cells. Blood plasma is the second part of the ECF. Materials travel between cells and the plasma in capillaries through the IF.[/caption]

Intracellular Fluid

The ICF lies within cells and is the principal component of the cytosol/cytoplasm. The ICF makes up about 60 percent of the total water in the human body, and in an average-size adult male, the ICF accounts for about 25 liters (seven gallons) of fluid (Figure 3). This fluid volume tends to be very stable, because the amount of water in living cells is closely regulated. If the amount of water inside a cell falls to a value that is too low, the cytosol becomes too concentrated with solutes to carry on normal cellular activities; if too much water enters a cell, the cell may burst and be destroyed.

[caption id="" align="aligncenter" width="380"]This pie chart shows that about 55% of water in the human body is intracellular fluid. About 30% of the water in the human body is interstitial fluid. Most of the remaining 15% of water is plasma, along with a small percentage labeled “other fluid”. Figure 3. A Pie Graph Showing the Proportion of Total Body Fluid in Each of the Body’s Fluid Compartments. Most of the water in the body is intracellular fluid. The second largest volume is the interstitial fluid, which surrounds cells that are not blood cells.[/caption]

Extracellular Fluid

The ECF accounts for the other one-third of the body’s water content. Approximately 20 percent of the ECF is found in plasma. Plasma travels through the body in blood vessels and transports a range of materials, including blood cells, proteins (including clotting factors and antibodies), electrolytes, nutrients, gases, and wastes. Gases, nutrients, and waste materials travel between capillaries and cells through the IF. Cells are separated from the IF by a selectively permeable cell membrane that helps regulate the passage of materials between the IF and the interior of the cell.

The body has other water-based ECF. These include the cerebrospinal fluid that bathes the brain and spinal cord, lymph, the synovial fluid in joints, the pleural fluid in the pleural cavities, the pericardial fluid in the cardiac sac, the peritoneal fluid in the peritoneal cavity, and the aqueous humor of the eye. Because these fluids are outside of cells, these fluids are also considered components of the ECF compartment.

Composition of Body Fluids

The compositions of the two components of the ECF—plasma and IF—are more similar to each other than either is to the ICF (Figure 4). Blood plasma has high concentrations of sodium, chloride, bicarbonate, and protein. The IF has high concentrations of sodium, chloride, and bicarbonate, but a relatively lower concentration of protein. In contrast, the ICF has elevated amounts of potassium, phosphate, magnesium, and protein. Overall, the ICF contains high concentrations of potassium and phosphate (HPO42−HPO42−), whereas both plasma and the ECF contain high concentrations of sodium and chloride.

[caption id="" align="aligncenter" width="380"]This bar graph shows the concentration of several ions and proteins in intracellular fluid, interstitial fluid and plasma. The ions and proteins are categories on the X axis . The Y axis shows concentration, in milliequivalents per liter, ranging from zero to 160. Three different colored bars are shown above each compound on the X axis. One bar represents intracellular fluid (ICF), a second bar represents interstitial fluid (IF, which is part of ECF) and the third bar represents plasma (ECF). Intracellular fluid contains high concentrations of K plus and HPO four two minus. It has lower concentrations of MG two plus and protein, and negligible amounts of the other compounds. Interstitial fluid contains high concentrations of NA plus and CL minus, along with a smaller amount of HCO 3 minus, and negligible amounts of the other compounds. Plasma contains large concentrations of NA plus and CL minus, with smaller concentrations of HCO 3 minus and protein, and negligible amounts of the other compounds. Figure 4. The Concentrations of Different Elements in Key Bodily Fluids. The graph shows the composition of the ICF, IF, and plasma. The compositions of plasma and IF are similar to one another but are quite different from the composition of the ICF.[/caption]

Most body fluids are neutral in charge. Thus, cations, or positively charged ions, and anions, or negatively charged ions, are balanced in fluids. As seen in the previous graph, sodium (Na+) ions and chloride (Cl-) ions are concentrated in the ECF of the body, whereas potassium (K+) ions are concentrated inside cells. Although sodium and potassium can “leak” through “pores” into and out of cells, respectively, the high levels of potassium and low levels of sodium in the ICF are maintained by sodium-potassium pumps in the cell membranes. These pumps use the energy supplied by ATP to pump sodium out of the cell and potassium into the cell (Figure 5).

[caption id="" align="aligncenter" width="520"]This diagram shows a sodium potassium pump embedded in the cell membrane. In the first step, the pump is opened to the cytosol and closed to the extracellular fluid. First, three sodium ions move into the pump from the cytosol. An ATP molecule binds to the cytosol side of the pump, causing the pump to change shape and open to the extracellular fluid. The pump is now closed to the cytosol. The sodium ions are then released into the extracellular fluid, after which two potassium ions enter the pump. Also at this point, the used ADP detaches from the cytosol side of the pump, leaving a single phosphate attached. The pump then changes shape again so that it closes to the extracellular fluid and again opens to the cytosol. This releases the two potassium ions into the cytosol. The single phosphate also detaches from the pump at this point so that the cycle can start anew. Two bars along the right hand side of the figure indicate that sodium normally diffuses into the cell down its concentration gradient while potassium usually diffuses out of the cell down its concentration gradient. Therefore, the sodium potassium pump is working against these natural concentration gradients. Figure 5. The Sodium-Potassium Pump. The sodium-potassium pump is powered by ATP to transfer sodium out of the cytoplasm and into the ECF. The pump also transfers potassium out of the ECF and into the cytoplasm. (credit: modification of work by Mariana Ruiz Villarreal)[/caption]

Fluid Movement between Compartments

Hydrostatic pressure, the force exerted by a fluid against a wall, causes movement of fluid between compartments. The hydrostatic pressure of blood is the pressure exerted by blood against the walls of the blood vessels by the pumping action of the heart. In capillaries, hydrostatic pressure (also known as capillary blood pressure) is higher than the opposing “colloid osmotic pressure” in blood—a “constant” pressure primarily produced by circulating albumin—at the arteriolar end of the capillary (Figure 6). This pressure forces plasma and nutrients out of the capillaries and into surrounding tissues. Fluid and the cellular wastes in the tissues enter the capillaries at the venule end, where the hydrostatic pressure is less than the osmotic pressure in the vessel. Filtration pressure squeezes fluid from the plasma in the blood to the IF surrounding the tissue cells. The surplus fluid in the interstitial space that is not returned directly back to the capillaries is drained from tissues by the lymphatic system, and then re-enters the vascular system at the subclavian veins.
[caption id="" align="aligncenter" width="480"]Alt text to come. Figure 6. Capillary Exchange. Net filtration occurs near the arterial end of the capillary since capillary hydrostatic pressure (CHP) is greater than blood colloidal osmotic pressure (BCOP). There is no net movement of fluid near the midpoint of the capillary since CHP = BCOP. Net reabsorption occurs near the venous end of the capillary since BCOP is greater than CHP.[/caption]

Hydrostatic pressure is especially important in governing the movement of water in the nephrons of the kidneys to ensure proper filtering of the blood to form urine. As hydrostatic pressure in the kidneys increases, the amount of water leaving the capillaries also increases, and more urine filtrate is formed. If hydrostatic pressure in the kidneys drops too low, as can happen in dehydration, the functions of the kidneys will be impaired, and less nitrogenous wastes will be removed from the bloodstream. Extreme dehydration can result in kidney failure.

Fluid also moves between compartments along an osmotic gradient. Recall that an osmotic gradient is produced by the difference in concentration of all solutes on either side of a semi-permeable membrane. The magnitude of the osmotic gradient is proportional to the difference in the concentration of solutes on one side of the cell membrane to that on the other side. Water will move by osmosis from the side where its concentration is high (and the concentration of solute is low) to the side of the membrane where its concentration is low (and the concentration of solute is high). In the body, water moves by osmosis from plasma to the IF (and the reverse) and from the IF to the ICF (and the reverse). In the body, water moves constantly into and out of fluid compartments as conditions change in different parts of the body.

For example, if you are sweating, you will lose water through your skin. Sweating depletes your tissues of water and increases the solute concentration in those tissues. As this happens, water diffuses from your blood into sweat glands and surrounding skin tissues that have become dehydrated because of the osmotic gradient. Additionally, as water leaves the blood, it is replaced by the water in other tissues throughout your body that are not dehydrated. If this continues, dehydration spreads throughout the body. When a dehydrated person drinks water and rehydrates, the water is redistributed by the same gradient, but in the opposite direction, replenishing water in all of the tissues.

Solute Movement between Compartments

The movement of some solutes between compartments is active, which consumes energy and is an active transport process, whereas the movement of other solutes is passive, which does not require energy. Active transport allows cells to move a specific substance against its concentration gradient through a membrane protein, requiring energy in the form of ATP. For example, the sodium-potassium pump employs active transport to pump sodium out of cells and potassium into cells, with both substances moving against their concentration gradients.

Passive transport of a molecule or ion depends on its ability to pass through the membrane, as well as the existence of a concentration gradient that allows the molecules to diffuse from an area of higher concentration to an area of lower concentration. Some molecules, like gases, lipids, and water itself (which also utilizes water channels in the membrane called aquaporins), slip fairly easily through the cell membrane; others, including polar molecules like glucose, amino acids, and ions do not. Some of these molecules enter and leave cells using facilitated transport, whereby the molecules move down a concentration gradient through specific protein channels in the membrane. This process does not require energy. For example, glucose is transferred into cells by glucose transporters that use facilitated transport (Figure 7).

[caption id="" align="aligncenter" width="480"]This diagram shows a carrier protein embedded in the plasma membrane between the cytoplasm and the extracellular fluid. There are several glucose molecules in the extracellular fluid. In the first step, the carrier protein is open to the extracellular fluid and closed to the cytosol. One of the glucose molecules travels from the extracellular fluid into the carrier protein. The protein then changes shape, closing at both ends. This pushes the glucose down into the carrier protein, closer to the cytosol end. The protein then opens on the cytosol side and closes on the extracellular fluid side, allowing the glucose to enter the cytosol. Figure 7. Facilitated Diffusion. Glucose molecules use facilitated diffusion to move down a concentration gradient through the carrier protein channels in the membrane. (credit: modification of work by Mariana Ruiz Villarreal)[/caption]
Disorders of the…

Fluid Balance: Edema Edema is the accumulation of excess water in the tissues. It is most common in the soft tissues of the extremities. The physiological causes of edema include water leakage from blood capillaries. Edema is almost always caused by an underlying medical condition, by the use of certain therapeutic drugs, by pregnancy, by localized injury, or by an allergic reaction. In the limbs, the symptoms of edema include swelling of the subcutaneous tissues, an increase in the normal size of the limb, and stretched, tight skin. One quick way to check for subcutaneous edema localized in a limb is to press a finger into the suspected area. Edema is likely if the depression persists for several seconds after the finger is removed (which is called “pitting”).

Pulmonary edema is excess fluid in the air sacs of the lungs, a common symptom of heart and/or kidney failure. People with pulmonary edema likely will experience difficulty breathing, and they may experience chest pain. Pulmonary edema can be life threatening, because it compromises gas exchange in the lungs, and anyone having symptoms should immediately seek medical care.

In pulmonary edema resulting from heart failure, excessive leakage of water occurs because fluids get “backed up” in the pulmonary capillaries of the lungs, when the left ventricle of the heart is unable to pump sufficient blood into the systemic circulation. Because the left side of the heart is unable to pump out its normal volume of blood, the blood in the pulmonary circulation gets “backed up,” starting with the left atrium, then into the pulmonary veins, and then into pulmonary capillaries. The resulting increased hydrostatic pressure within pulmonary capillaries, as blood is still coming in from the pulmonary arteries, causes fluid to be pushed out of them and into lung tissues.

Other causes of edema include damage to blood vessels and/or lymphatic vessels, or a decrease in osmotic pressure in chronic and severe liver disease, where the liver is unable to manufacture plasma proteins (Figure 8). A decrease in the normal levels of plasma proteins results in a decrease of colloid osmotic pressure (which counterbalances the hydrostatic pressure) in the capillaries. This process causes loss of water from the blood to the surrounding tissues, resulting in edema.

[caption id="" align="aligncenter" width="350"]This photo shows the dorsal surfaces of a person’s right and left hands. The left hand is normal, with the several blood vessels visible under the skin. However, the top of the right hand is swollen and no blood vessels are visible. Figure 8. Edema. An allergic reaction can cause capillaries in the hand to leak excess fluid that accumulates in the tissues. (credit: Jane Whitney)[/caption]

Mild, transient edema of the feet and legs may be caused by sitting or standing in the same position for long periods of time, as in the work of a toll collector or a supermarket cashier. This is because deep veins in the lower limbs rely on skeletal muscle contractions to push on the veins and thus “pump” blood back to the heart. Otherwise, the venous blood pools in the lower limbs and can leak into surrounding tissues.

Medications that can result in edema include vasodilators, calcium channel blockers used to treat hypertension, non-steroidal anti-inflammatory drugs, estrogen therapies, and some diabetes medications. Underlying medical conditions that can contribute to edema include congestive heart failure, kidney damage and kidney disease, disorders that affect the veins of the legs, and cirrhosis and other liver disorders.

Therapy for edema usually focuses on elimination of the cause. Activities that can reduce the effects of the condition include appropriate exercises to keep the blood and lymph flowing through the affected areas. Other therapies include elevation of the affected part to assist drainage, massage and compression of the areas to move the fluid out of the tissues, and decreased salt intake to decrease sodium and water retention.

]]> 2404 2395 1 0 1.1 Overview of Anatomy and Physiology https://pressbooks.bccampus.ca/apdouglas2017/chapter/1-1-overview-of-anatomy-and-physiology-2/ Wed, 02 Aug 2017 22:00:53 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2422

Learning Objectives

  • Compare and contrast anatomy and physiology, including their specializations and methods of study
  • Discuss the fundamental relationship between anatomy and physiology

Human anatomy is the scientific study of the body’s structures. Some of these structures are very small and can only be observed and analyzed with the assistance of a microscope. Other larger structures can readily be seen, manipulated, measured, and weighed. The word “anatomy” comes from a Greek root that means “to cut apart.” Human anatomy was first studied by observing the exterior of the body and observing the wounds of soldiers and other injuries. Later, physicians were allowed to dissect bodies of the dead to augment their knowledge. When a body is dissected, its structures are cut apart in order to observe their physical attributes and their relationships to one another. Dissection is still used in medical schools, anatomy courses, and in pathology labs. In order to observe structures in living people, however, a number of imaging techniques have been developed. These techniques allow clinicians to visualize structures inside the living body such as a cancerous tumor or a fractured bone.

Like most scientific disciplines, anatomy has areas of specialization. Gross anatomy is the study of the larger structures of the body, those visible without the aid of magnification (Figure 1a). Macro- means “large,” thus, gross anatomy is also referred to as macroscopic anatomy. In contrast, micro- means “small,” and microscopic anatomy is the study of structures that can be observed only with the use of a microscope or other magnification devices (Figure 1b). Microscopic anatomy includes cytology, the study of cells and histology, the study of tissues. As the technology of microscopes has advanced, anatomists have been able to observe smaller and smaller structures of the body, from slices of large structures like the heart, to the three-dimensional structures of large molecules in the body.

[caption id="" align="aligncenter" width="480"]Photo A shows an entire human brain which has a lumpy and deeply striated appearance. Photo B is a micrograph of neural tissue. It contains two roughly diamond-shaped cells with dark nuclei. The cells are embedded in a light colored tissue containing smaller cells and fiber strands. Figure 1. Gross and Microscopic Anatomy. (a) Gross anatomy considers large structures such as the brain. (b) Microscopic anatomy can deal with the same structures, though at a different scale. This is a micrograph of nerve cells from the brain. LM × 1600. (credit a: “WriterHound”/Wikimedia Commons; credit b: Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Anatomists take two general approaches to the study of the body’s structures: regional and systemic. Regional anatomy is the study of the interrelationships of all of the structures in a specific body region, such as the abdomen. Studying regional anatomy helps us appreciate the interrelationships of body structures, such as how muscles, nerves, blood vessels, and other structures work together to serve a particular body region. In contrast, systemic anatomy is the study of the structures that make up a discrete body system—that is, a group of structures that work together to perform a unique body function. For example, a systemic anatomical study of the muscular system would consider all of the skeletal muscles of the body.

Whereas anatomy is about structure, physiology is about function. Human physiology is the scientific study of the chemistry and physics of the structures of the body and the ways in which they work together to support the functions of life. Much of the study of physiology centers on the body’s tendency toward homeostasis. Homeostasis is the state of steady internal conditions maintained by living things. The study of physiology certainly includes observation, both with the naked eye and with microscopes, as well as manipulations and measurements. However, current advances in physiology usually depend on carefully designed laboratory experiments that reveal the functions of the many structures and chemical compounds that make up the human body.

Like anatomists, physiologists typically specialize in a particular branch of physiology. For example, neurophysiology is the study of the brain, spinal cord, and nerves and how these work together to perform functions as complex and diverse as vision, movement, and thinking. Physiologists may work from the organ level (exploring, for example, what different parts of the brain do) to the molecular level (such as exploring how an electrochemical signal travels along nerves).

Form is closely related to function in all living things. For example, the thin flap of your eyelid can snap down to clear away dust particles and almost instantaneously slide back up to allow you to see again. At the microscopic level, the arrangement and function of the nerves and muscles that serve the eyelid allow for its quick action and retreat. At a smaller level of analysis, the function of these nerves and muscles likewise relies on the interactions of specific molecules and ions. Even the three-dimensional structure of certain molecules is essential to their function.

Your study of anatomy and physiology will make more sense if you continually relate the form of the structures you are studying to their function. In fact, it can be somewhat frustrating to attempt to study anatomy without an understanding of the physiology that a body structure supports. Imagine, for example, trying to appreciate the unique arrangement of the bones of the human hand if you had no conception of the function of the hand. Fortunately, your understanding of how the human hand manipulates tools—from pens to cell phones—helps you appreciate the unique alignment of the thumb in opposition to the four fingers, making your hand a structure that allows you to pinch and grasp objects and type text messages.

]]> 2422 2420 1 0 1.4 Requirements for Human Life https://pressbooks.bccampus.ca/apdouglas2017/chapter/1-4-requirements-for-human-life/ Wed, 02 Aug 2017 22:01:06 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2425

Learning Objectives

By the end of this section, you will be able to:
  • Discuss the role of oxygen and nutrients in maintaining human survival
  • Explain why extreme heat and extreme cold threaten human survival
  • Explain how the pressure exerted by gases and fluids influences human survival

Humans have been adapting to life on Earth for at least the past 200,000 years. Earth and its atmosphere have provided us with air to breathe, water to drink, and food to eat, but these are not the only requirements for survival. Although you may rarely think about it, you also cannot live outside of a certain range of temperature and pressure that the surface of our planet and its atmosphere provides. The next sections explore these four requirements of life.

Oxygen

Atmospheric air is only about 20 percent oxygen, but that oxygen is a key component of the chemical reactions that keep the body alive, including the reactions that produce ATP. Brain cells are especially sensitive to lack of oxygen because of their requirement for a high-and-steady production of ATP. Brain damage is likely within five minutes without oxygen, and death is likely within ten minutes.

Nutrients

A nutrient is a substance in foods and beverages that is essential to human survival. The three basic classes of nutrients are water, the energy-yielding and body-building nutrients, and the micronutrients (vitamins and minerals).

The most critical nutrient is water. Depending on the environmental temperature and our state of health, we may be able to survive for only a few days without water. The body’s functional chemicals are dissolved and transported in water, and the chemical reactions of life take place in water. Moreover, water is the largest component of cells, blood, and the fluid between cells, and water makes up about 70 percent of an adult’s body mass. Water also helps regulate our internal temperature and cushions, protects, and lubricates joints and many other body structures.

The energy-yielding nutrients are primarily carbohydrates and lipids, while proteins mainly supply the amino acids that are the building blocks of the body itself. You ingest these in plant and animal foods and beverages, and the digestive system breaks them down into molecules small enough to be absorbed. The breakdown products of carbohydrates and lipids can then be used in the metabolic processes that convert them to ATP. Although you might feel as if you are starving after missing a single meal, you can survive without consuming the energy-yielding nutrients for at least several weeks.

Water and the energy-yielding nutrients are also referred to as macronutrients because the body needs them in large amounts. In contrast, micronutrients are vitamins and minerals. These elements and compounds participate in many essential chemical reactions and processes, such as nerve impulses, and some, such as calcium, also contribute to the body’s structure. Your body can store some of the micronutrients in its tissues, and draw on those reserves if you fail to consume them in your diet for a few days or weeks. Some others micronutrients, such as vitamin C and most of the B vitamins, are water-soluble and cannot be stored, so you need to consume them every day or two.

Narrow Range of Temperature

You have probably seen news stories about athletes who died of heat stroke, or hikers who died of exposure to cold. Such deaths occur because the chemical reactions upon which the body depends can only take place within a narrow range of body temperature, from just below to just above 37°C (98.6°F). When body temperature rises well above or drops well below normal, certain proteins (enzymes) that facilitate chemical reactions lose their normal structure and their ability to function and the chemical reactions of metabolism cannot proceed.

That said, the body can respond effectively to short-term exposure to heat (Figure 1) or cold. One of the body’s responses to heat is, of course, sweating. As sweat evaporates from skin, it removes some thermal energy from the body, cooling it. Adequate water (from the extracellular fluid in the body) is necessary to produce sweat, so adequate fluid intake is essential to balance that loss during the sweat response. Not surprisingly, the sweat response is much less effective in a humid environment because the air is already saturated with water. Thus, the sweat on the skin’s surface is not able to evaporate, and internal body temperature can get dangerously high.
[caption id="" align="aligncenter" width="420"]This photo shows two white-clad men riding camels through a sparse desert. Two canvas tents are visible in the background. Figure 1. Extreme Heat. Humans adapt to some degree to repeated exposure to high temperatures. (credit: McKay Savage/flickr)[/caption]

The body can also respond effectively to short-term exposure to cold. One response to cold is shivering, which is random muscle movement that generates heat. Another response is increased breakdown of stored energy to generate heat. When that energy reserve is depleted, however, and the core temperature begins to drop significantly, red blood cells will lose their ability to give up oxygen, denying the brain of this critical component of ATP production. This lack of oxygen can cause confusion, lethargy, and eventually loss of consciousness and death. The body responds to cold by reducing blood circulation to the extremities, the hands and feet, in order to prevent blood from cooling there and so that the body’s core can stay warm. Even when core body temperature remains stable, however, tissues exposed to severe cold, especially the fingers and toes, can develop frostbite when blood flow to the extremities has been much reduced. This form of tissue damage can be permanent and lead to gangrene, requiring amputation of the affected region.

Everyday Connection

Controlled Hypothermia As you have learned, the body continuously engages in coordinated physiological processes to maintain a stable temperature. In some cases, however, overriding this system can be useful, or even life-saving. Hypothermia is the clinical term for an abnormally low body temperature (hypo- = “below” or “under”). Controlled hypothermia is clinically induced hypothermia performed in order to reduce the metabolic rate of an organ or of a person’s entire body.

Controlled hypothermia often is used, for example, during open-heart surgery because it decreases the metabolic needs of the brain, heart, and other organs, reducing the risk of damage to them. When controlled hypothermia is used clinically, the patient is given medication to prevent shivering. The body is then cooled to 25–32°C (79–89°F). The heart is stopped and an external heart-lung pump maintains circulation to the patient’s body. The heart is cooled further and is maintained at a temperature below 15°C (60°F) for the duration of the surgery. This very cold temperature helps the heart muscle to tolerate its lack of blood supply during the surgery.

Some emergency department physicians use controlled hypothermia to reduce damage to the heart in patients who have suffered a cardiac arrest. In the emergency department, the physician induces coma and lowers the patient’s body temperature to approximately 91 degrees. This condition, which is maintained for 24 hours, slows the patient’s metabolic rate. Because the patient’s organs require less blood to function, the heart’s workload is reduced.

Narrow Range of Atmospheric Pressure

Pressure is a force exerted by a substance that is in contact with another substance. Atmospheric pressure is pressure exerted by the mixture of gases (primarily nitrogen and oxygen) in the Earth’s atmosphere. Although you may not perceive it, atmospheric pressure is constantly pressing down on your body. This pressure keeps gases within your body, such as the gaseous nitrogen in body fluids, dissolved. If you were suddenly ejected from a space ship above Earth’s atmosphere, you would go from a situation of normal pressure to one of very low pressure. The pressure of the nitrogen gas in your blood would be much higher than the pressure of nitrogen in the space surrounding your body. As a result, the nitrogen gas in your blood would expand, forming bubbles that could block blood vessels and even cause cells to break apart.

Atmospheric pressure does more than just keep blood gases dissolved. Your ability to breathe—that is, to take in oxygen and release carbon dioxide—also depends upon a precise atmospheric pressure. Altitude sickness occurs in part because the atmosphere at high altitudes exerts less pressure, reducing the exchange of these gases, and causing shortness of breath, confusion, headache, lethargy, and nausea. Mountain climbers carry oxygen to reduce the effects of both low oxygen levels and low barometric pressure at higher altitudes (Figure 2).

[caption id="" align="aligncenter" width="380"]This photo shows Mount Everest as seen from a distance. It is a large, pyramid-shaped, craggy peak with many smaller snow-covered peaks in the foreground. The peak of Mount Everest is partially occluded by clouds. Figure 2. Harsh Conditions. Climbers on Mount Everest must accommodate extreme cold, low oxygen levels, and low barometric pressure in an environment hostile to human life. (credit: Melanie Ko/flickr)[/caption]
Homeostatic Imbalances

Decompression Sickness Decompression sickness (DCS) is a condition in which gases dissolved in the blood or in other body tissues are no longer dissolved following a reduction in pressure on the body. This condition affects underwater divers who surface from a deep dive too quickly, and it can affect pilots flying at high altitudes in planes with unpressurized cabins. Divers often call this condition “the bends,” a reference to joint pain that is a symptom of DCS.

In all cases, DCS is brought about by a reduction in barometric pressure. At high altitude, barometric pressure is much less than on Earth’s surface because pressure is produced by the weight of the column of air above the body pressing down on the body. The very great pressures on divers in deep water are likewise from the weight of a column of water pressing down on the body. For divers, DCS occurs at normal barometric pressure (at sea level), but it is brought on by the relatively rapid decrease of pressure as divers rise from the high pressure conditions of deep water to the now low, by comparison, pressure at sea level. Not surprisingly, diving in deep mountain lakes, where barometric pressure at the surface of the lake is less than that at sea level is more likely to result in DCS than diving in water at sea level.

In DCS, gases dissolved in the blood (primarily nitrogen) come rapidly out of solution, forming bubbles in the blood and in other body tissues. This occurs because when pressure of a gas over a liquid is decreased, the amount of gas that can remain dissolved in the liquid also is decreased. It is air pressure that keeps your normal blood gases dissolved in the blood. When pressure is reduced, less gas remains dissolved. You have seen this in effect when you open a carbonated drink. Removing the seal of the bottle reduces the pressure of the gas over the liquid. This in turn causes bubbles as dissolved gases (in this case, carbon dioxide) come out of solution in the liquid.

The most common symptoms of DCS are pain in the joints, with headache and disturbances of vision occurring in 10 percent to 15 percent of cases. Left untreated, very severe DCS can result in death. Immediate treatment is with pure oxygen. The affected person is then moved into a hyperbaric chamber. A hyperbaric chamber is a reinforced, closed chamber that is pressurized to greater than atmospheric pressure. It treats DCS by repressurizing the body so that pressure can then be removed much more gradually. Because the hyperbaric chamber introduces oxygen to the body at high pressure, it increases the concentration of oxygen in the blood. This has the effect of replacing some of the nitrogen in the blood with oxygen, which is easier to tolerate out of solution.

The dynamic pressure of body fluids is also important to human survival. For example, blood pressure, which is the pressure exerted by blood as it flows within blood vessels, must be great enough to enable blood to reach all body tissues, and yet low enough to ensure that the delicate blood vessels can withstand the friction and force of the pulsating flow of pressurized blood.

]]> 2425 2420 2 0 2.1 Elements and Atoms: the Building Blocks of Matter https://pressbooks.bccampus.ca/apdouglas2017/chapter/2-1-elements-and-atoms-the-building-blocks-of-matter/ Wed, 02 Aug 2017 22:01:35 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2437

Learning Objectives

By the end of this section, you will be able to:
  • Discuss the relationships between matter, mass, elements, compounds, atoms, and subatomic particles
  • Distinguish between atomic number and mass number
  • Identify the key distinction between isotopes of the same element
  • Explain how electrons occupy electron shells and their contribution to an atom’s relative stability

The substance of the universe—from a grain of sand to a star—is called matter. Scientists define matter as anything that occupies space and has mass. An object’s mass and its weight are related concepts, but not quite the same. An object’s mass is the amount of matter contained in the object, and the object’s mass is the same whether that object is on Earth or in the zero-gravity environment of outer space. An object’s weight, on the other hand, is its mass as affected by the pull of gravity. Where gravity strongly pulls on an object’s mass its weight is greater than it is where gravity is less strong. An object of a certain mass weighs less on the moon, for example, than it does on Earth because the gravity of the moon is less than that of Earth. In other words, weight is variable, and is influenced by gravity. A piece of cheese that weighs a pound on Earth weighs only a few ounces on the moon.

Elements and Compounds

All matter in the natural world is composed of one or more of the 92 fundamental substances called elements. An element is a pure substance that is distinguished from all other matter by the fact that it cannot be created or broken down by ordinary chemical means. While your body can assemble many of the chemical compounds needed for life from their constituent elements, it cannot make elements. They must come from the environment. A familiar example of an element that you must take in is calcium (Ca++). Calcium is essential to the human body; it is absorbed and used for a number of processes, including strengthening bones. When you consume dairy products your digestive system breaks down the food into components small enough to cross into the bloodstream. Among these is calcium, which, because it is an element, cannot be broken down further. The elemental calcium in cheese, therefore, is the same as the calcium that forms your bones. Some other elements you might be familiar with are oxygen, sodium, and iron. The elements in the human body are shown in Figure 1, beginning with the most abundant: oxygen (O), carbon (C), hydrogen (H), and nitrogen (N). Each element’s name can be replaced by a one- or two-letter symbol; you will become familiar with some of these during this course. All the elements in your body are derived from the foods you eat and the air you breathe.

[caption id="" align="aligncenter" width="520"]This figure shows a human body with the percentage of the main elements in the body, in the left panel. In the right panel, a table lists the elements and the percentages in the body. Figure 1. Elements of the Human Body. The main elements that compose the human body are shown from most abundant to least abundant.[/caption]

In nature, elements rarely occur alone. Instead, they combine to form compounds. A compound is a substance composed of two or more elements joined by chemical bonds. For example, the compound glucose is an important body fuel. It is always composed of the same three elements: carbon, hydrogen, and oxygen. Moreover, the elements that make up any given compound always occur in the same relative amounts. In glucose, there are always six carbon and six oxygen units for every twelve hydrogen units. But what, exactly, are these “units” of elements?

Atoms and Subatomic Particles

An atom is the smallest quantity of an element that retains the unique properties of that element. In other words, an atom of hydrogen is a unit of hydrogen—the smallest amount of hydrogen that can exist. As you might guess, atoms are almost unfathomably small. The period at the end of this sentence is millions of atoms wide.

Atomic Structure and Energy

Atoms are made up of even smaller subatomic particles, three types of which are important: the proton, neutron, and electron. The number of positively-charged protons and non-charged (“neutral”) neutrons, gives mass to the atom, and the number of each in the nucleus of the atom determine the element. The number of negatively-charged electrons that “spin” around the nucleus at close to the speed of light equals the number of protons. An electron has about 1/2000th the mass of a proton or neutron.

Figure 2 shows two models that can help you imagine the structure of an atom—in this case, helium (He). In the planetary model, helium’s two electrons are shown circling the nucleus in a fixed orbit depicted as a ring. Although this model is helpful in visualizing atomic structure, in reality, electrons do not travel in fixed orbits, but whiz around the nucleus erratically in a so-called electron cloud.

[caption id="" align="aligncenter" width="285"]The top panel of this figure shows two electrons orbiting around the nucleus of a Helium atom. The bottom panel of this figure shows a cloud of electrons surrounding the nucleus of a Helium atom. Figure 2. Two Models of Atomic Structure. (a) In the planetary model, the electrons of helium are shown in fixed orbits, depicted as rings, at a precise distance from the nucleus, somewhat like planets orbiting the sun. (b) In the electron cloud model, the electrons of carbon are shown in the variety of locations they would have at different distances from the nucleus over time.[/caption]

An atom’s protons and electrons carry electrical charges. Protons, with their positive charge, are designated p+. Electrons, which have a negative charge, are designated e–. An atom’s neutrons have no charge: they are electrically neutral. Just as a magnet sticks to a steel refrigerator because their opposite charges attract, the positively charged protons attract the negatively charged electrons. This mutual attraction gives the atom some structural stability. The attraction by the positively charged nucleus helps keep electrons from straying far. The number of protons and electrons within a neutral atom are equal, thus, the atom’s overall charge is balanced.

Atomic Number and Mass Number

An atom of carbon is unique to carbon, but a proton of carbon is not. One proton is the same as another, whether it is found in an atom of carbon, sodium (Na), or iron (Fe). The same is true for neutrons and electrons. So, what gives an element its distinctive properties—what makes carbon so different from sodium or iron? The answer is the unique quantity of protons each contains. Carbon by definition is an element whose atoms contain six protons. No other element has exactly six protons in its atoms. Moreover, all atoms of carbon, whether found in your liver or in a lump of coal, contain six protons. Thus, the atomic number, which is the number of protons in the nucleus of the atom, identifies the element. Because an atom usually has the same number of electrons as protons, the atomic number identifies the usual number of electrons as well.

In their most common form, many elements also contain the same number of neutrons as protons. The most common form of carbon, for example, has six neutrons as well as six protons, for a total of 12 subatomic particles in its nucleus. An element’s mass number is the sum of the number of protons and neutrons in its nucleus. So the most common form of carbon’s mass number is 12. (Electrons have so little mass that they do not appreciably contribute to the mass of an atom.) Carbon is a relatively light element. Uranium (U), in contrast, has a mass number of 238 and is referred to as a heavy metal. Its atomic number is 92 (it has 92 protons) but it contains 146 neutrons; it has the most mass of all the naturally occurring elements.

The periodic table of the elements, shown in Figure 3, is a chart identifying the 92 elements found in nature, as well as several larger, unstable elements discovered experimentally. The elements are arranged in order of their atomic number, with hydrogen and helium at the top of the table, and the more massive elements below. The periodic table is a useful device because for each element, it identifies the chemical symbol, the atomic number, and the mass number, while organizing elements according to their propensity to react with other elements. The number of protons and electrons in an element are equal. The number of protons and neutrons may be equal for some elements, but are not equal for all.

[caption id="" align="aligncenter" width="650"]This figure shows the periodic table. Figure 3. The Periodic Table of the Elements. (credit: R.A. Dragoset, A. Musgrove, C.W. Clark, W.C. Martin)[/caption]
[caption id="" align="aligncenter" width="120"]QR Code representing a URL Visit this website to view the periodic table.[/caption] Visit this website to view the periodic table. In the periodic table of the elements, elements in a single column have the same number of electrons that can participate in a chemical reaction. These electrons are known as “valence electrons.” For example, the elements in the first column all have a single valence electron, an electron that can be “donated” in a chemical reaction with another atom. What is the meaning of a mass number shown in parentheses?

Isotopes

Although each element has a unique number of protons, it can exist as different isotopes. An isotope is one of the different forms of an element, distinguished from one another by different numbers of neutrons. The standard isotope of carbon is 12C, commonly called carbon twelve. 12C has six protons and six neutrons, for a mass number of twelve. All of the isotopes of carbon have the same number of protons; therefore, 13C has seven neutrons, and 14C has eight neutrons. The different isotopes of an element can also be indicated with the mass number hyphenated (for example, C-12 instead of 12C). Hydrogen has three common isotopes, shown in Figure 4.

[caption id="" align="aligncenter" width="380"]This figure shows the three isotopes of hydrogen: hydrogen, deuterium, and tritium. Figure 4. Isotopes of Hydrogen. Protium, designated 1H, has one proton and no neutrons. It is by far the most abundant isotope of hydrogen in nature. Deuterium, designated 2H, has one proton and one neutron. Tritium, designated 3H, has two neutrons.[/caption]

An isotope that contains more than the usual number of neutrons is referred to as a heavy isotope. An example is 14C. Heavy isotopes tend to be unstable, and unstable isotopes are radioactive. A radioactive isotope is an isotope whose nucleus readily decays, giving off subatomic particles and electromagnetic energy. Different radioactive isotopes (also called radioisotopes) differ in their half-life, the time it takes for half of any size sample of an isotope to decay. For example, the half-life of tritium—a radioisotope of hydrogen—is about 12 years, indicating it takes 12 years for half of the tritium nuclei in a sample to decay. Excessive exposure to radioactive isotopes can damage human cells and even cause cancer and birth defects, but when exposure is controlled, some radioactive isotopes can be useful in medicine. For more information, see the Career Connections.

Career Connection
Interventional Radiologist The controlled use of radioisotopes has advanced medical diagnosis and treatment of disease. Interventional radiologists are physicians who treat disease by using minimally invasive techniques involving radiation. Many conditions that could once only be treated with a lengthy and traumatic operation can now be treated non-surgically, reducing the cost, pain, length of hospital stay, and recovery time for patients. For example, in the past, the only options for a patient with one or more tumors in the liver were surgery and chemotherapy (the administration of drugs to treat cancer). Some liver tumors, however, are difficult to access surgically, and others could require the surgeon to remove too much of the liver. Moreover, chemotherapy is highly toxic to the liver, and certain tumors do not respond well to it anyway. In some such cases, an interventional radiologist can treat the tumors by disrupting their blood supply, which they need if they are to continue to grow. In this procedure, called radioembolization, the radiologist accesses the liver with a fine needle, threaded through one of the patient’s blood vessels. The radiologist then inserts tiny radioactive “seeds” into the blood vessels that supply the tumors. In the days and weeks following the procedure, the radiation emitted from the seeds destroys the vessels and directly kills the tumor cells in the vicinity of the treatment.

Radioisotopes emit subatomic particles that can be detected and tracked by imaging technologies. One of the most advanced uses of radioisotopes in medicine is the positron emission tomography (PET) scanner, which detects the activity in the body of a very small injection of radioactive glucose, the simple sugar that cells use for energy. The PET camera reveals to the medical team which of the patient’s tissues are taking up the most glucose. Thus, the most metabolically active tissues show up as bright “hot spots” on the images (Figure 5). PET can reveal some cancerous masses because cancer cells consume glucose at a high rate to fuel their rapid reproduction.

[caption id="" align="aligncenter" width="480"]This figure shows multiple images from a PET scan. Figure 5. PET Scan. PET highlights areas in the body where there is relatively high glucose use, which is characteristic of cancerous tissue. This PET scan shows sites of the spread of a large primary tumor to other sites.[/caption]

The Behavior of Electrons

In the human body, atoms do not exist as independent entities. Rather, they are constantly reacting with other atoms to form and to break down more complex substances. To fully understand anatomy and physiology you must grasp how atoms participate in such reactions. The key is understanding the behavior of electrons.

Although electrons do not follow rigid orbits a set distance away from the atom’s nucleus, they do tend to stay within certain regions of space called electron shells. An electron shell is a layer of electrons that encircle the nucleus at a distinct energy level.

The atoms of the elements found in the human body have from one to five electron shells, and all electron shells hold eight electrons except the first shell, which can only hold two. This configuration of electron shells is the same for all atoms. The precise number of shells depends on the number of electrons in the atom. Hydrogen and helium have just one and two electrons, respectively. If you take a look at the periodic table of the elements, you will notice that hydrogen and helium are placed alone on either sides of the top row; they are the only elements that have just one electron shell (Figure 6). A second shell is necessary to hold the electrons in all elements larger than hydrogen and helium.

Lithium (Li), whose atomic number is 3, has three electrons. Two of these fill the first electron shell, and the third spills over into a second shell. The second electron shell can accommodate as many as eight electrons. Carbon, with its six electrons, entirely fills its first shell, and half-fills its second. With ten electrons, neon (Ne) entirely fills its two electron shells. Again, a look at the periodic table reveals that all of the elements in the second row, from lithium to neon, have just two electron shells. Atoms with more than ten electrons require more than two shells. These elements occupy the third and subsequent rows of the periodic table.
[caption id="" align="aligncenter" width="420"]This four panel figure shows four different atoms with the electrons in orbit around the nucleus. Figure 6. Electron Shells. Electrons orbit the atomic nucleus at distinct levels of energy called electron shells. (a) With one electron, hydrogen only half-fills its electron shell. Helium also has a single shell, but its two electrons completely fill it. (b) The electrons of carbon completely fill its first electron shell, but only half-fills its second. (c) Neon, an element that does not occur in the body, has 10 electrons, filling both of its electron shells.[/caption]

The factor that most strongly governs the tendency of an atom to participate in chemical reactions is the number of electrons in its valence shell. A valence shell is an atom’s outermost electron shell. If the valence shell is full, the atom is stable; meaning its electrons are unlikely to be pulled away from the nucleus by the electrical charge of other atoms. If the valence shell is not full, the atom is reactive; meaning it will tend to react with other atoms in ways that make the valence shell full. Consider hydrogen, with its one electron only half-filling its valence shell. This single electron is likely to be drawn into relationships with the atoms of other elements, so that hydrogen’s single valence shell can be stabilized.

All atoms (except hydrogen and helium with their single electron shells) are most stable when there are exactly eight electrons in their valence shell. This principle is referred to as the octet rule, and it states that an atom will give up, gain, or share electrons with another atom so that it ends up with eight electrons in its own valence shell. For example, oxygen, with six electrons in its valence shell, is likely to react with other atoms in a way that results in the addition of two electrons to oxygen’s valence shell, bringing the number to eight. When two hydrogen atoms each share their single electron with oxygen, covalent bonds are formed, resulting in a molecule of water, H2O.

In nature, atoms of one element tend to join with atoms of other elements in characteristic ways. For example, carbon commonly fills its valence shell by linking up with four atoms of hydrogen. In so doing, the two elements form the simplest of organic molecules, methane, which also is one of the most abundant and stable carbon-containing compounds on Earth. As stated above, another example is water; oxygen needs two electrons to fill its valence shell. It commonly interacts with two atoms of hydrogen, forming H2O. Incidentally, the name “hydrogen” reflects its contribution to water (hydro- = “water”; -gen = “maker”). Thus, hydrogen is the “water maker.”

]]> 2437 2429 1 0 13.4 The Peripheral Nervous System https://pressbooks.bccampus.ca/apdouglas2017/chapter/13-4-the-peripheral-nervous-system/ Wed, 02 Aug 2017 22:02:16 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2534

Learning Objectives

By the end of this section, you will be able to:
  • Describe the structures found in the PNS
  • Distinguish between somatic and autonomic structures, including the special peripheral structures of the enteric nervous system
  • Name the twelve cranial nerves and explain the functions associated with each
  • Describe the sensory and motor components of spinal nerves and the plexuses that they pass through
The PNS is not as contained as the CNS because it is defined as everything that is not the CNS. Some peripheral structures are incorporated into the other organs of the body. In describing the anatomy of the PNS, it is necessary to describe the common structures, the nerves and the ganglia, as they are found in various parts of the body. Many of the neural structures that are incorporated into other organs are features of the digestive system; these structures are known as the enteric nervous system and are a special subset of the PNS.

Ganglia

A ganglion is a group of neuron cell bodies in the periphery. Ganglia can be categorized, for the most part, as either sensory ganglia or autonomic ganglia, referring to their primary functions. The most common type of sensory ganglion is a dorsal (posterior) root ganglion. These ganglia are the cell bodies of neurons with axons that are sensory endings in the periphery, such as in the skin, and that extend into the CNS through the dorsal nerve root. The ganglion is an enlargement of the nerve root. Under microscopic inspection, it can be seen to include the cell bodies of the neurons, as well as bundles of fibers that are the posterior nerve root (Figure 1). The cells of the dorsal root ganglion are unipolar cells, classifying them by shape. Also, the small round nuclei of satellite cells can be seen surrounding—as if they were orbiting—the neuron cell bodies.

[caption id="" align="aligncenter" width="580"]This micrograph shows the structure of the dorsal root ganglion. The cell bodies of the neurons and the axon bundles are also labeled. Figure 1. Dorsal Root Ganglion. The cell bodies of sensory neurons, which are unipolar neurons by shape, are seen in this photomicrograph. Also, the fibrous region is composed of the axons of these neurons that are passing through the ganglion to be part of the dorsal nerve root (tissue source: canine). LM × 40. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]
[caption id="" align="aligncenter" width="580"]This micrograph shows a magnified view of the dorsal root ganglion, showing the satellite cells and the cell bodies of sensory neurons. Figure 2. Spinal Cord and Root Ganglion. The slide includes both a cross-section of the lumbar spinal cord and a section of the dorsal root ganglion (see also Figure 1) (tissue source: canine). LM × 1600. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Another type of sensory ganglion is a cranial nerve ganglion. This is analogous to the dorsal root ganglion, except that it is associated with a cranial nerve instead of a spinal nerve. The roots of cranial nerves are within the cranium, whereas the ganglia are outside the skull. For example, the trigeminal ganglion is superficial to the temporal bone whereas its associated nerve is attached to the mid-pons region of the brain stem. The neurons of cranial nerve ganglia are also unipolar in shape with associated satellite cells.

The other major category of ganglia are those of the autonomic nervous system, which is divided into the sympathetic and parasympathetic nervous systems. The sympathetic chain ganglia constitute a row of ganglia along the vertebral column that receive central input from the lateral horn of the thoracic and upper lumbar spinal cord. Superior to the chain ganglia are three paravertebral ganglia in the cervical region. Three other autonomic ganglia that are related to the sympathetic chain are the prevertebral ganglia, which are located outside of the chain but have similar functions. They are referred to as prevertebral because they are anterior to the vertebral column. The neurons of these autonomic ganglia are multipolar in shape, with dendrites radiating out around the cell body where synapses from the spinal cord neurons are made. The neurons of the chain, paravertebral, and prevertebral ganglia then project to organs in the head and neck, thoracic, abdominal, and pelvic cavities to regulate the sympathetic aspect of homeostatic mechanisms.

Another group of autonomic ganglia are the terminal ganglia that receive input from cranial nerves or sacral spinal nerves and are responsible for regulating the parasympathetic aspect of homeostatic mechanisms. These two sets of ganglia, sympathetic and parasympathetic, often project to the same organs—one input from the chain ganglia and one input from a terminal ganglion—to regulate the overall function of an organ. For example, the heart receives two inputs such as these; one increases heart rate, and the other decreases it. The terminal ganglia that receive input from cranial nerves are found in the head and neck, as well as the thoracic and upper abdominal cavities, whereas the terminal ganglia that receive sacral input are in the lower abdominal and pelvic cavities.

Terminal ganglia below the head and neck are often incorporated into the wall of the target organ as a plexus. A plexus, in a general sense, is a network of fibers or vessels. This can apply to nervous tissue (as in this instance) or structures containing blood vessels (such as a choroid plexus). For example, the enteric plexus is the extensive network of axons and neurons in the wall of the small and large intestines. The enteric plexus is actually part of the enteric nervous system, along with the gastric plexuses and the esophageal plexus. Though the enteric nervous system receives input originating from central neurons of the autonomic nervous system, it does not require CNS input to function. In fact, it operates independently to regulate the digestive system.

Nerves

Bundles of axons in the PNS are referred to as nerves. These structures in the periphery are different than the central counterpart, called a tract. Nerves are composed of more than just nervous tissue. They have connective tissues invested in their structure, as well as blood vessels supplying the tissues with nourishment. The outer surface of a nerve is a surrounding layer of fibrous connective tissue called the epineurium. Within the nerve, axons are further bundled into fascicles, which are each surrounded by their own layer of fibrous connective tissue called perineurium. Finally, individual axons are surrounded by loose connective tissue called the endoneurium (Figure 3). These three layers are similar to the connective tissue sheaths for muscles. Nerves are associated with the region of the CNS to which they are connected, either as cranial nerves connected to the brain or spinal nerves connected to the spinal cord.
[caption id="" align="aligncenter" width="480"]This figure shows the structure of a nerve. The top panel shows the cross section of a spinal nerve and the major parts are labeled. The bottom panel shows a micrograph of the cross-section of a spinal nerve. Figure 3. Nerve Structure. The structure of a nerve is organized by the layers of connective tissue on the outside, around each fascicle, and surrounding the individual nerve fibers (tissue source: simian). LM × 40. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]
[caption id="" align="aligncenter" width="580"]This micrograph shows a magnified view of the nerve. The perineurium and the endoneurium are labeled. Figure 4. Close-Up of Nerve Trunk. Zoom in on this slide of a nerve trunk to examine the endoneurium, perineurium, and epineurium in greater detail (tissue source: simian). LM × 1600. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Cranial Nerves

The nerves attached to the brain are the cranial nerves, which are primarily responsible for the sensory and motor functions of the head and neck (one of these nerves targets organs in the thoracic and abdominal cavities as part of the parasympathetic nervous system). There are twelve cranial nerves, which are designated CNI through CNXII for “Cranial Nerve,” using Roman numerals for 1 through 12. They can be classified as sensory nerves, motor nerves, or a combination of both, meaning that the axons in these nerves originate out of sensory ganglia external to the cranium or motor nuclei within the brain stem. Sensory axons enter the brain to synapse in a nucleus. Motor axons connect to skeletal muscles of the head or neck. Three of the nerves are solely composed of sensory fibers; five are strictly motor; and the remaining four are mixed nerves.

Learning the cranial nerves is a tradition in anatomy courses, and students have always used mnemonic devices to remember the nerve names. A traditional mnemonic is the rhyming couplet, “On Old Olympus’ Towering Tops/A Finn And German Viewed Some Hops,” in which the initial letter of each word corresponds to the initial letter in the name of each nerve. The names of the nerves have changed over the years to reflect current usage and more accurate naming. An exercise to help learn this sort of information is to generate a mnemonic using words that have personal significance. The names of the cranial nerves are listed in Table 3 along with a brief description of their function, their source (sensory ganglion or motor nucleus), and their target (sensory nucleus or skeletal muscle). They are listed here with a brief explanation of each nerve (Figure 5).

The olfactory nerve and optic nerve are responsible for the sense of smell and vision, respectively. The oculomotor nerve is responsible for eye movements by controlling four of the extraocular muscles. It is also responsible for lifting the upper eyelid when the eyes point up, and for pupillary constriction. The trochlear nerve and the abducens nerve are both responsible for eye movement, but do so by controlling different extraocular muscles. The trigeminal nerve is responsible for cutaneous sensations of the face and controlling the muscles of mastication. The facial nerve is responsible for the muscles involved in facial expressions, as well as part of the sense of taste and the production of saliva. The vestibulocochlear nerve is responsible for the senses of hearing and balance. The glossopharyngeal nerve is responsible for controlling muscles in the oral cavity and upper throat, as well as part of the sense of taste and the production of saliva. The vagus nerve is responsible for contributing to homeostatic control of the organs of the thoracic and upper abdominal cavities. The spinal accessory nerve is responsible for controlling the muscles of the neck, along with cervical spinal nerves. The hypoglossal nerve is responsible for controlling the muscles of the lower throat and tongue.
[caption id="" align="aligncenter" width="480"]This diagrams shows the brain and the main nerves in the brain are labeled. Figure 5. The Cranial Nerves. The anatomical arrangement of the roots of the cranial nerves observed from an inferior view of the brain.[/caption]
Three of the cranial nerves also contain autonomic fibers, and a fourth is almost purely a component of the autonomic system. The oculomotor, facial, and glossopharyngeal nerves contain fibers that contact autonomic ganglia. The oculomotor fibers initiate pupillary constriction, whereas the facial and glossopharyngeal fibers both initiate salivation. The vagus nerve primarily targets autonomic ganglia in the thoracic and upper abdominal cavities.

Another important aspect of the cranial nerves that lends itself to a mnemonic is the functional role each nerve plays. The nerves fall into one of three basic groups. They are sensory, motor, or both (see Table 3). The sentence, “Some Say Marry Money But My Brother Says Brains Beauty Matter More,” corresponds to the basic function of each nerve. The first, second, and eighth nerves are purely sensory: the olfactory (CNI), optic (CNII), and vestibulocochlear (CNVIII) nerves. The three eye-movement nerves are all motor: the oculomotor (CNIII), trochlear (CNIV), and abducens (CNVI). The spinal accessory (CNXI) and hypoglossal (CNXII) nerves are also strictly motor. The remainder of the nerves contain both sensory and motor fibers. They are the trigeminal (CNV), facial (CNVII), glossopharyngeal (CNIX), and vagus (CNX) nerves. The nerves that convey both are often related to each other. The trigeminal and facial nerves both concern the face; one concerns the sensations and the other concerns the muscle movements. The facial and glossopharyngeal nerves are both responsible for conveying gustatory, or taste, sensations as well as controlling salivary glands. The vagus nerve is involved in visceral responses to taste, namely the gag reflex. This is not an exhaustive list of what these combination nerves do, but there is a thread of relation between them.

Cranial Nerves (Table 3)
Mnemonic # Name Function (S/M/B) Central connection (nuclei) Peripheral connection (ganglion or muscle)
On I Olfactory Smell (S) Olfactory bulb Olfactory epithelium
Old II Optic Vision (S) Hypothalamus/thalamus/midbrain Retina (retinal ganglion cells)
Olympus’ III Oculomotor Eye movements (M) Oculomotor nucleus Extraocular muscles (other 4), levator palpebrae superioris, ciliary ganglion (autonomic)
Towering IV Trochlear Eye movements (M) Trochlear nucleus Superior oblique muscle
Tops V Trigeminal Sensory/motor – face (B) Trigeminal nuclei in the midbrain, pons, and medulla Trigeminal
A VI Abducens Eye movements (M) Abducens nucleus Lateral rectus muscle
Finn VII Facial Motor – face, Taste (B) Facial nucleus, solitary nucleus, superior salivatory nucleus Facial muscles, Geniculate ganglion, Pterygopalatine ganglion (autonomic)
And VIII Auditory (Vestibulocochlear) Hearing/balance (S) Cochlear nucleus, Vestibular nucleus/cerebellum Spiral ganglion (hearing), Vestibular ganglion (balance)
German IX Glossopharyngeal Motor – throat Taste (B) Solitary nucleus, inferior salivatory nucleus, nucleus ambiguus Pharyngeal muscles, Geniculate ganglion, Otic ganglion (autonomic)
Viewed X Vagus Motor/sensory – viscera (autonomic) (B) Medulla Terminal ganglia serving thoracic and upper abdominal organs (heart and small intestines)
Some XI Spinal Accessory Motor – head and neck (M) Spinal accessory nucleus Neck muscles
Hops XII Hypoglossal Motor – lower throat (M) Hypoglossal nucleus Muscles of the larynx and lower pharynx

Spinal Nerves

The nerves connected to the spinal cord are the spinal nerves. The arrangement of these nerves is much more regular than that of the cranial nerves. All of the spinal nerves are combined sensory and motor axons that separate into two nerve roots. The sensory axons enter the spinal cord as the dorsal nerve root. The motor fibers, both somatic and autonomic, emerge as the ventral nerve root. The dorsal root ganglion for each nerve is an enlargement of the spinal nerve.

There are 31 spinal nerves, named for the level of the spinal cord at which each one emerges. There are eight pairs of cervical nerves designated C1 to C8, twelve thoracic nerves designated T1 to T12, five pairs of lumbar nerves designated L1 to L5, five pairs of sacral nerves designated S1 to S5, and one pair of coccygeal nerves. The nerves are numbered from the superior to inferior positions, and each emerges from the vertebral column through the intervertebral foramen at its level. The first nerve, C1, emerges between the first cervical vertebra and the occipital bone. The second nerve, C2, emerges between the first and second cervical vertebrae. The same occurs for C3 to C7, but C8 emerges between the seventh cervical vertebra and the first thoracic vertebra. For the thoracic and lumbar nerves, each one emerges between the vertebra that has the same designation and the next vertebra in the column. The sacral nerves emerge from the sacral foramina along the length of that unique vertebra.

Spinal nerves extend outward from the vertebral column to enervate the periphery. The nerves in the periphery are not straight continuations of the spinal nerves, but rather the reorganization of the axons in those nerves to follow different courses. Axons from different spinal nerves will come together into a systemic nerve. This occurs at four places along the length of the vertebral column, each identified as a nerve plexus, whereas the other spinal nerves directly correspond to nerves at their respective levels. In this instance, the word plexus is used to describe networks of nerve fibers with no associated cell bodies.

Of the four nerve plexuses, two are found at the cervical level, one at the lumbar level, and one at the sacral level (Figure 6). The cervical plexus is composed of axons from spinal nerves C1 through C5 and branches into nerves in the posterior neck and head, as well as the phrenic nerve, which connects to the diaphragm at the base of the thoracic cavity. The other plexus from the cervical level is the brachial plexus. Spinal nerves C4 through T1 reorganize through this plexus to give rise to the nerves of the arms, as the name brachial suggests. A large nerve from this plexus is the radial nerve from which the axillary nerve branches to go to the armpit region. The radial nerve continues through the arm and is paralleled by the ulnar nerve and the median nerve. The lumbar plexus arises from all the lumbar spinal nerves and gives rise to nerves enervating the pelvic region and the anterior leg. The femoral nerve is one of the major nerves from this plexus, which gives rise to the saphenous nerve as a branch that extends through the anterior lower leg. The sacral plexus comes from the lower lumbar nerves L4 and L5 and the sacral nerves S1 to S4. The most significant systemic nerve to come from this plexus is the sciatic nerve, which is a combination of the tibial nerve and the fibular nerve. The sciatic nerve extends across the hip joint and is most commonly associated with the condition sciatica, which is the result of compression or irritation of the nerve or any of the spinal nerves giving rise to it.

These plexuses are described as arising from spinal nerves and giving rise to certain systemic nerves, but they contain fibers that serve sensory functions or fibers that serve motor functions. This means that some fibers extend from cutaneous or other peripheral sensory surfaces and send action potentials into the CNS. Those are axons of sensory neurons in the dorsal root ganglia that enter the spinal cord through the dorsal nerve root. Other fibers are the axons of motor neurons of the anterior horn of the spinal cord, which emerge in the ventral nerve root and send action potentials to cause skeletal muscles to contract in their target regions. For example, the radial nerve contains fibers of cutaneous sensation in the arm, as well as motor fibers that move muscles in the arm.

Spinal nerves of the thoracic region, T2 through T11, are not part of the plexuses but rather emerge and give rise to the intercostal nerves found between the ribs, which articulate with the vertebrae surrounding the spinal nerve.

[caption id="" align="aligncenter" width="440"]This figure shows a torso of a human body. The spinal cord is shown in the body and the main nerves along the spinal cord are labeled. Figure 6. Nerve Plexuses of the Body. There are four main nerve plexuses in the human body. The cervical plexus supplies nerves to the posterior head and neck, as well as to the diaphragm. The brachial plexus supplies nerves to the arm. The lumbar plexus supplies nerves to the anterior leg. The sacral plexus supplies nerves to the posterior leg.[/caption]
]]> 2534 2512 3 0 14.2 Central Processing https://pressbooks.bccampus.ca/apdouglas2017/chapter/14-2-central-processing/ Wed, 02 Aug 2017 22:02:35 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2563

Learning Objectives

By the end of this section, you will be able to:
  • Describe the pathways that sensory systems follow into the central nervous system
  • Differentiate between the two major ascending pathways in the spinal cord
  • Describe the pathway of somatosensory input from the face and compare it to the ascending pathways in the spinal cord
  • Explain topographical representations of sensory information in at least two systems
  • Describe two pathways of visual processing and the functions associated with each

Sensory Pathways

Specific regions of the CNS coordinate different somatic processes using sensory inputs and motor outputs of peripheral nerves. A simple case is a reflex caused by a synapse between a dorsal sensory neuron axon and a motor neuron in the ventral horn. More complex arrangements are possible to integrate peripheral sensory information with higher processes. The important regions of the CNS that play a role in somatic processes can be separated into the spinal cord brain stem, diencephalon, cerebral cortex, and subcortical structures.

Spinal Cord and Brain Stem

A sensory pathway that carries peripheral sensations to the brain is referred to as an ascending pathway, or ascending tract. The various sensory modalities each follow specific pathways through the CNS. Tactile and other somatosensory stimuli activate receptors in the skin, muscles, tendons, and joints throughout the entire body. However, the somatosensory pathways are divided into two separate systems on the basis of the location of the receptor neurons. Somatosensory stimuli from below the neck pass along the sensory pathways of the spinal cord, whereas somatosensory stimuli from the head and neck travel through the cranial nerves—specifically, the trigeminal system.

The dorsal column system (sometimes referred to as the dorsal column–medial lemniscus) and the spinothalamic tract are two major pathways that bring sensory information to the brain (Figure 1). The sensory pathways in each of these systems are composed of three successive neurons.

The dorsal column system begins with the axon of a dorsal root ganglion neuron entering the dorsal root and joining the dorsal column white matter in the spinal cord. As axons of this pathway enter the dorsal column, they take on a positional arrangement so that axons from lower levels of the body position themselves medially, whereas axons from upper levels of the body position themselves laterally. The dorsal column is separated into two component tracts, the fasciculus gracilis that contains axons from the legs and lower body, and the fasciculus cuneatus that contains axons from the upper body and arms.

The axons in the dorsal column terminate in the nuclei of the medulla, where each synapses with the second neuron in their respective pathway. The nucleus gracilis is the target of fibers in the fasciculus gracilis, whereas the nucleus cuneatus is the target of fibers in the fasciculus cuneatus. The second neuron in the system projects from one of the two nuclei and then decussates, or crosses the midline of the medulla. These axons then continue to ascend the brain stem as a bundle called the medial lemniscus. These axons terminate in the thalamus, where each synapses with the third neuron in their respective pathway. The third neuron in the system projects its axons to the postcentral gyrus of the cerebral cortex, where somatosensory stimuli are initially processed and the conscious perception of the stimulus occurs.

The spinothalamic tract also begins with neurons in a dorsal root ganglion. These neurons extend their axons to the dorsal horn, where they synapse with the second neuron in their respective pathway. The name “spinothalamic” comes from this second neuron, which has its cell body in the spinal cord gray matter and connects to the thalamus. Axons from these second neurons then decussate within the spinal cord and ascend to the brain and enter the thalamus, where each synapses with the third neuron in its respective pathway. The neurons in the thalamus then project their axons to the spinothalamic tract, which synapses in the postcentral gyrus of the cerebral cortex.

These two systems are similar in that they both begin with dorsal root ganglion cells, as with most general sensory information. The dorsal column system is primarily responsible for touch sensations and proprioception, whereas the spinothalamic tract pathway is primarily responsible for pain and temperature sensations. Another similarity is that the second neurons in both of these pathways are contralateral, because they project across the midline to the other side of the brain or spinal cord. In the dorsal column system, this decussation takes place in the brain stem; in the spinothalamic pathway, it takes place in the spinal cord at the same spinal cord level at which the information entered. The third neurons in the two pathways are essentially the same. In both, the second neuron synapses in the thalamus, and the thalamic neuron projects to the somatosensory cortex.

[caption id="" align="aligncenter" width="530"]The left panel shows the dorsal column system and its connection to the brain. The right column shows the spinothalamic tract and its connection to the brain. Figure 1. Ascending Sensory Pathways of the Spinal Cord. The dorsal column system and spinothalamic tract are the major ascending pathways that connect the periphery with the brain.[/caption]

The trigeminal pathway carries somatosensory information from the face, head, mouth, and nasal cavity. As with the previously discussed nerve tracts, the sensory pathways of the trigeminal pathway each involve three successive neurons. First, axons from the trigeminal ganglion enter the brain stem at the level of the pons. These axons project to one of three locations. The spinal trigeminal nucleus of the medulla receives information similar to that carried by spinothalamic tract, such as pain and temperature sensations. Other axons go to either the chief sensory nucleus in the pons or the mesencephalic nuclei in the midbrain. These nuclei receive information like that carried by the dorsal column system, such as touch, pressure, vibration, and proprioception. Axons from the second neuron decussate and ascend to the thalamus along the trigeminothalamic tract. In the thalamus, each axon synapses with the third neuron in its respective pathway. Axons from the third neuron then project from the thalamus to the primary somatosensory cortex of the cerebrum.

The sensory pathway for gustation travels along the facial and glossopharyngeal cranial nerves, which synapse with neurons of the solitary nucleus in the brain stem. Axons from the solitary nucleus then project to the ventral posterior nucleus of the thalamus. Finally, axons from the ventral posterior nucleus project to the gustatory cortex of the cerebral cortex, where taste is processed and consciously perceived.

The sensory pathway for audition travels along the vestibulocochlear nerve, which synapses with neurons in the cochlear nuclei of the superior medulla. Within the brain stem, input from either ear is combined to extract location information from the auditory stimuli. Whereas the initial auditory stimuli received at the cochlea strictly represent the frequency—or pitch—of the stimuli, the locations of sounds can be determined by comparing information arriving at both ears.

Sound localization is a feature of central processing in the auditory nuclei of the brain stem. Sound localization is achieved by the brain calculating the interaural time difference and the interaural intensity difference. A sound originating from a specific location will arrive at each ear at different times, unless the sound is directly in front of the listener. If the sound source is slightly to the left of the listener, the sound will arrive at the left ear microseconds before it arrives at the right ear (Figure 2). This time difference is an example of an interaural time difference. Also, the sound will be slightly louder in the left ear than in the right ear because some of the sound waves reaching the opposite ear are blocked by the head. This is an example of an interaural intensity difference.

[caption id="" align="aligncenter" width="290"]The top panel shows a person hearing a sound source that arrives in both his ears at the same time with the same intensity. The bottom panel shows a sound source that is not centered and arrives at different times with different intensities in each ear. Figure 2. Auditory Brain Stem Mechanisms of Sound Localization. Localizing sound in the horizontal plane is achieved by processing in the medullary nuclei of the auditory system. Connections between neurons on either side are able to compare very slight differences in sound stimuli that arrive at either ear and represent interaural time and intensity differences.[/caption]

Auditory processing continues on to a nucleus in the midbrain called the inferior colliculus. Axons from the inferior colliculus project to two locations, the thalamus and the superior colliculus. The medial geniculate nucleus of the thalamus receives the auditory information and then projects that information to the auditory cortex in the temporal lobe of the cerebral cortex. The superior colliculus receives input from the visual and somatosensory systems, as well as the ears, to initiate stimulation of the muscles that turn the head and neck toward the auditory stimulus.

Balance is coordinated through the vestibular system, the nerves of which are composed of axons from the vestibular ganglion that carries information from the utricle, saccule, and semicircular canals. The system contributes to controlling head and neck movements in response to vestibular signals. An important function of the vestibular system is coordinating eye and head movements to maintain visual attention. Most of the axons terminate in the vestibular nuclei of the medulla. Some axons project from the vestibular ganglion directly to the cerebellum, with no intervening synapse in the vestibular nuclei. The cerebellum is primarily responsible for initiating movements on the basis of equilibrium information.

Neurons in the vestibular nuclei project their axons to targets in the brain stem. One target is the reticular formation, which influences respiratory and cardiovascular functions in relation to body movements. A second target of the axons of neurons in the vestibular nuclei is the spinal cord, which initiates the spinal reflexes involved with posture and balance. To assist the visual system, fibers of the vestibular nuclei project to the oculomotor, trochlear, and abducens nuclei to influence signals sent along the cranial nerves. These connections constitute the pathway of the vestibulo-ocular reflex (VOR), which compensates for head and body movement by stabilizing images on the retina (Figure 3). Finally, the vestibular nuclei project to the thalamus to join the proprioceptive pathway of the dorsal column system, allowing conscious perception of equilibrium.

[caption id="" align="aligncenter" width="480"]This image shows how the excitation of eye muscles on one side, the inhibition of these muscles on the other side, and the compensating eye movements work together in vestibular ocular reflex. Figure 3. Vestibulo-ocular Reflex. Connections between the vestibular system and the cranial nerves controlling eye movement keep the eyes centered on a visual stimulus, even though the head is moving. During head movement, the eye muscles move the eyes in the opposite direction as the head movement, keeping the visual stimulus centered in the field of view.[/caption]

The connections of the optic nerve are more complicated than those of other cranial nerves. Instead of the connections being between each eye and the brain, visual information is segregated between the left and right sides of the visual field. In addition, some of the information from one side of the visual field projects to the opposite side of the brain. Within each eye, the axons projecting from the medial side of the retina decussate at the optic chiasm. For example, the axons from the medial retina of the left eye cross over to the right side of the brain at the optic chiasm. However, within each eye, the axons projecting from the lateral side of the retina do not decussate. For example, the axons from the lateral retina of the right eye project back to the right side of the brain. Therefore the left field of view of each eye is processed on the right side of the brain, whereas the right field of view of each eye is processed on the left side of the brain (Figure 4).

[caption id="" align="aligncenter" width="395"]This image shows the right and left visual fields in the brain. It describes how the optical fields map to different sides of the brain. Figure 4. Segregation of Visual Field Information at the Optic Chiasm. Contralateral visual field information from the lateral retina projects to the ipsilateral brain, whereas ipsilateral visual field information has to decussate at the optic chiasm to reach the opposite side of the brain.[/caption]

A unique clinical presentation that relates to this anatomic arrangement is the loss of lateral peripheral vision, known as bilateral hemianopia. This is different from “tunnel vision” because the superior and inferior peripheral fields are not lost. Visual field deficits can be disturbing for a patient, but in this case, the cause is not within the visual system itself. A growth of the pituitary gland presses against the optic chiasm and interferes with signal transmission. However, the axons projecting to the same side of the brain are unaffected. Therefore, the patient loses the outermost areas of their field of vision and cannot see objects to their right and left.

Extending from the optic chiasm, the axons of the visual system are referred to as the optic tract instead of the optic nerve. The optic tract has three major targets, two in the diencephalon and one in the midbrain. The connection between the eyes and diencephalon is demonstrated during development, in which the neural tissue of the retina differentiates from that of the diencephalon by the growth of the secondary vesicles. The connections of the retina into the CNS are a holdover from this developmental association. The majority of the connections of the optic tract are to the thalamus—specifically, the lateral geniculate nucleus. Axons from this nucleus then project to the visual cortex of the cerebrum, located in the occipital lobe. Another target of the optic tract is the superior colliculus.

In addition, a very small number of RGC axons project from the optic chiasm to the suprachiasmatic nucleus of the hypothalamus. These RGCs are photosensitive, in that they respond to the presence or absence of light. Unlike the photoreceptors, however, these photosensitive RGCs cannot be used to perceive images. By simply responding to the absence or presence of light, these RGCs can send information about day length. The perceived proportion of sunlight to darkness establishes the circadian rhythm of our bodies, allowing certain physiological events to occur at approximately the same time every day.

Diencephalon

The diencephalon is beneath the cerebrum and includes the thalamus and hypothalamus. In the somatic nervous system, the thalamus is an important relay for communication between the cerebrum and the rest of the nervous system. The hypothalamus has both somatic and autonomic functions. In addition, the hypothalamus communicates with the limbic system, which controls emotions and memory functions.

Sensory input to the thalamus comes from most of the special senses and ascending somatosensory tracts. Each sensory system is relayed through a particular nucleus in the thalamus. The thalamus is a required transfer point for most sensory tracts that reach the cerebral cortex, where conscious sensory perception begins. The one exception to this rule is the olfactory system. The olfactory tract axons from the olfactory bulb project directly to the cerebral cortex, along with the limbic system and hypothalamus.

The thalamus is a collection of several nuclei that can be categorized into three anatomical groups. White matter running through the thalamus defines the three major regions of the thalamus, which are an anterior nucleus, a medial nucleus, and a lateral group of nuclei. The anterior nucleus serves as a relay between the hypothalamus and the emotion and memory-producing limbic system. The medial nuclei serve as a relay for information from the limbic system and basal ganglia to the cerebral cortex. This allows memory creation during learning, but also determines alertness. The special and somatic senses connect to the lateral nuclei, where their information is relayed to the appropriate sensory cortex of the cerebrum.

Cortical Processing

As described earlier, many of the sensory axons are positioned in the same way as their corresponding receptor cells in the body. This allows identification of the position of a stimulus on the basis of which receptor cells are sending information. The cerebral cortex also maintains this sensory topography in the particular areas of the cortex that correspond to the position of the receptor cells. The somatosensory cortex provides an example in which, in essence, the locations of the somatosensory receptors in the body are mapped onto the somatosensory cortex. This mapping is often depicted using a sensory homunculus (Figure 5).

The term homunculus comes from the Latin word for “little man” and refers to a map of the human body that is laid across a portion of the cerebral cortex. In the somatosensory cortex, the external genitals, feet, and lower legs are represented on the medial face of the gyrus within the longitudinal fissure. As the gyrus curves out of the fissure and along the surface of the parietal lobe, the body map continues through the thighs, hips, trunk, shoulders, arms, and hands. The head and face are just lateral to the fingers as the gyrus approaches the lateral sulcus. The representation of the body in this topographical map is medial to lateral from the lower to upper body. It is a continuation of the topographical arrangement seen in the dorsal column system, where axons from the lower body are carried in the fasciculus gracilis, whereas axons from the upper body are carried in the fasciculus cuneatus. As the dorsal column system continues into the medial lemniscus, these relationships are maintained. Also, the head and neck axons running from the trigeminal nuclei to the thalamus run adjacent to the upper body fibers. The connections through the thalamus maintain topography such that the anatomic information is preserved. Note that this correspondence does not result in a perfectly miniature scale version of the body, but rather exaggerates the more sensitive areas of the body, such as the fingers and lower face. Less sensitive areas of the body, such as the shoulders and back, are mapped to smaller areas on the cortex.

[caption id="" align="aligncenter" width="380"]This image shows the areas of the brain that control and respond to the different senses. Figure 5. The Sensory Homunculus. A cartoon representation of the sensory homunculus arranged adjacent to the cortical region in which the processing takes place.[/caption]

Likewise, the topographic relationship between the retina and the visual cortex is maintained throughout the visual pathway. The visual field is projected onto the two retinae, as described above, with sorting at the optic chiasm. The right peripheral visual field falls on the medial portion of the right retina and the lateral portion of the left retina. The right medial retina then projects across the midline through the optic chiasm. This results in the right visual field being processed in the left visual cortex. Likewise, the left visual field is processed in the right visual cortex (see Figure 4). Though the chiasm is helping to sort right and left visual information, superior and inferior visual information is maintained topographically in the visual pathway. Light from the superior visual field falls on the inferior retina, and light from the inferior visual field falls on the superior retina. This topography is maintained such that the superior region of the visual cortex processes the inferior visual field and vice versa. Therefore, the visual field information is inverted and reversed as it enters the visual cortex—up is down, and left is right. However, the cortex processes the visual information such that the final conscious perception of the visual field is correct. The topographic relationship is evident in that information from the foveal region of the retina is processed in the center of the primary visual cortex. Information from the peripheral regions of the retina are correspondingly processed toward the edges of the visual cortex. Similar to the exaggerations in the sensory homunculus of the somatosensory cortex, the foveal-processing area of the visual cortex is disproportionately larger than the areas processing peripheral vision.

In an experiment performed in the 1960s, subjects wore prism glasses so that the visual field was inverted before reaching the eye. On the first day of the experiment, subjects would duck when walking up to a table, thinking it was suspended from the ceiling. However, after a few days of acclimation, the subjects behaved as if everything were represented correctly. Therefore, the visual cortex is somewhat flexible in adapting to the information it receives from our eyes (Figure 6).

[caption id="" align="aligncenter" width="380"]This image shows the mapping of the right and left visual fields on the brain. It also explains how the brain merges images from both visual fields. Figure 6. Topographic Mapping of the Retina onto the Visual Cortex. The visual field projects onto the retina through the lenses and falls on the retinae as an inverted, reversed image. The topography of this image is maintained as the visual information travels through the visual pathway to the cortex.[/caption]

The cortex has been described as having specific regions that are responsible for processing specific information; there is the visual cortex, somatosensory cortex, gustatory cortex, etc. However, our experience of these senses is not divided. Instead, we experience what can be referred to as a seamless percept. Our perceptions of the various sensory modalities—though distinct in their content—are integrated by the brain so that we experience the world as a continuous whole.

In the cerebral cortex, sensory processing begins at the primary sensory cortex, then proceeds to an association area, and finally, into a multimodal integration area. For example, the visual pathway projects from the retinae through the thalamus to the primary visual cortex in the occipital lobe. This area is primarily in the medial wall within the longitudinal fissure. Here, visual stimuli begin to be recognized as basic shapes. Edges of objects are recognized and built into more complex shapes. Also, inputs from both eyes are compared to extract depth information. Because of the overlapping field of view between the two eyes, the brain can begin to estimate the distance of stimuli based on binocular depth cues.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Watch this video to learn more about how the brain perceives 3-D motion.[/caption]

Depth Perception, 3-D Movies, and Optical Illusions The visual field is projected onto the retinal surface, where photoreceptors transduce light energy into neural signals for the brain to interpret. The retina is a two-dimensional surface, so it does not encode three-dimensional information. However, we can perceive depth. How is that accomplished?

Two ways in which we can extract depth information from the two-dimensional retinal signal are based on monocular cues and binocular cues, respectively. Monocular depth cues are those that are the result of information within the two-dimensional visual field. One object that overlaps another object has to be in front. Relative size differences are also a cue. For example, if a basketball appears larger than the basket, then the basket must be further away. On the basis of experience, we can estimate how far away the basket is. Binocular depth cues compare information represented in the two retinae because they do not see the visual field exactly the same.

The centers of the two eyes are separated by a small distance, which is approximately 6 to 6.5 cm in most people. Because of this offset, visual stimuli do not fall on exactly the same spot on both retinae unless we are fixated directly on them and they fall on the fovea of each retina. All other objects in the visual field, either closer or farther away than the fixated object, will fall on different spots on the retina. When vision is fixed on an object in space, closer objects will fall on the lateral retina of each eye, and more distant objects will fall on the medial retina of either eye (Figure 7). This is easily observed by holding a finger up in front of your face as you look at a more distant object. You will see two images of your finger that represent the two disparate images that are falling on either retina.

These depth cues, both monocular and binocular, can be exploited to make the brain think there are three dimensions in two-dimensional information. This is the basis of 3-D movies. The projected image on the screen is two dimensional, but it has disparate information embedded in it. The 3-D glasses that are available at the theater filter the information so that only one eye sees one version of what is on the screen, and the other eye sees the other version. If you take the glasses off, the image on the screen will have varying amounts of blur because both eyes are seeing both layers of information, and the third dimension will not be evident. Some optical illusions can take advantage of depth cues as well, though those are more often using monocular cues to fool the brain into seeing different parts of the scene as being at different depths.

[caption id="" align="aligncenter" width="380"]This image shows how the left and right eye view objects that are closer and farther away. Figure 7. Retinal Disparity. Because of the interocular distance, which results in objects of different distances falling on different spots of the two retinae, the brain can extract depth perception from the two-dimensional information of the visual field.[/caption]

There are two main regions that surround the primary cortex that are usually referred to as areas V2 and V3 (the primary visual cortex is area V1). These surrounding areas are the visual association cortex. The visual association regions develop more complex visual perceptions by adding color and motion information. The information processed in these areas is then sent to regions of the temporal and parietal lobes. Visual processing has two separate streams of processing: one into the temporal lobe and one into the parietal lobe. These are the ventral and dorsal streams, respectively (Figure 8). The ventral stream identifies visual stimuli and their significance. Because the ventral stream uses temporal lobe structures, it begins to interact with the non-visual cortex and may be important in visual stimuli becoming part of memories. The dorsal stream locates objects in space and helps in guiding movements of the body in response to visual inputs. The dorsal stream enters the parietal lobe, where it interacts with somatosensory cortical areas that are important for our perception of the body and its movements. The dorsal stream can then influence frontal lobe activity where motor functions originate.

[caption id="" align="aligncenter" width="475"]This image shows the side of the human brain and maps different regions to different visual functions. Figure 8. Ventral and Dorsal Visual Streams. From the primary visual cortex in the occipital lobe, visual processing continues in two streams—one into the temporal lobe and one into the parietal lobe.[/caption]
]]> 2563 2535 2 0 15.2 Autonomic Reflexes and Homeostasis https://pressbooks.bccampus.ca/apdouglas2017/chapter/15-2-autonomic-reflexes-and-homeostasis/ Wed, 02 Aug 2017 22:03:05 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2580

Learning Objectives

By the end of this section, you will be able to:
  • Compare the structure of somatic and autonomic reflex arcs
  • Explain the differences in sympathetic and parasympathetic reflexes
  • Differentiate between short and long reflexes
  • Determine the effect of the autonomic nervous system on the regulation of the various organ systems on the basis of the signaling molecules involved
  • Describe the effects of drugs that affect autonomic function

The autonomic nervous system regulates organ systems through circuits that resemble the reflexes described in the somatic nervous system. The main difference between the somatic and autonomic systems is in what target tissues are effectors. Somatic responses are solely based on skeletal muscle contraction. The autonomic system, however, targets cardiac and smooth muscle, as well as glandular tissue. Whereas the basic circuit is a reflex arc, there are differences in the structure of those reflexes for the somatic and autonomic systems.

The Structure of Reflexes

One difference between a somatic reflex, such as the withdrawal reflex, and a visceral reflex, which is an autonomic reflex, is in the efferent branch. The output of a somatic reflex is the lower motor neuron in the ventral horn of the spinal cord that projects directly to a skeletal muscle to cause its contraction. The output of a visceral reflex is a two-step pathway starting with the preganglionic fiber emerging from a lateral horn neuron in the spinal cord, or a cranial nucleus neuron in the brain stem, to a ganglion—followed by the postganglionic fiber projecting to a target effector. The other part of a reflex, the afferent branch, is often the same between the two systems. Sensory neurons receiving input from the periphery—with cell bodies in the sensory ganglia, either of a cranial nerve or a dorsal root ganglion adjacent to the spinal cord—project into the CNS to initiate the reflex (Figure 1). The Latin root “effere” means “to carry.” Adding the prefix “ef-” suggests the meaning “to carry away,” whereas adding the prefix “af-” suggests “to carry toward or inward.”

[caption id="" align="aligncenter" width="500"]The top panel in this figure shows the autonomic efferent pathway. The spinal cord is shown on the left, and a myelinated axon is shown going from the spinal cord to the central neuron. An unmyelinated axon is shown going from the central neuron to the target effector. The bottom panel shows the somatic efferent pathway. The spinal cord is shown on the left, and a myelinated axon is shown going from the spinal cord to the target effector director. In both cases, magnified images show the synapses. Figure 1. Comparison of Somatic and Visceral Reflexes. The afferent inputs to somatic and visceral reflexes are essentially the same, whereas the efferent branches are different. Somatic reflexes, for instance, involve a direct connection from the ventral horn of the spinal cord to the skeletal muscle. Visceral reflexes involve a projection from the central neuron to a ganglion, followed by a second projection from the ganglion to the target effector.[/caption]

Afferent Branch

The afferent branch of a reflex arc does differ between somatic and visceral reflexes in some instances. Many of the inputs to visceral reflexes are from special or somatic senses, but particular senses are associated with the viscera that are not part of the conscious perception of the environment through the somatic nervous system. For example, there is a specific type of mechanoreceptor, called a baroreceptor, in the walls of the aorta and carotid sinuses that senses the stretch of those organs when blood volume or pressure increases. You do not have a conscious perception of having high blood pressure, but that is an important afferent branch of the cardiovascular and, particularly, vasomotor reflexes. The sensory neuron is essentially the same as any other general sensory neuron. The baroreceptor apparatus is part of the ending of a unipolar neuron that has a cell body in a sensory ganglion. The baroreceptors from the carotid arteries have axons in the glossopharyngeal nerve, and those from the aorta have axons in the vagus nerve.

Though visceral senses are not primarily a part of conscious perception, those sensations sometimes make it to conscious awareness. If a visceral sense is strong enough, it will be perceived. The sensory homunculus—the representation of the body in the primary somatosensory cortex—only has a small region allotted for the perception of internal stimuli. If you swallow a large bolus of food, for instance, you will probably feel the lump of that food as it pushes through your esophagus, or even if your stomach is distended after a large meal. If you inhale especially cold air, you can feel it as it enters your larynx and trachea. These sensations are not the same as feeling high blood pressure or blood sugar levels.

When particularly strong visceral sensations rise to the level of conscious perception, the sensations are often felt in unexpected places. For example, strong visceral sensations of the heart will be felt as pain in the left shoulder and left arm. This irregular pattern of projection of conscious perception of visceral sensations is called referred pain. Depending on the organ system affected, the referred pain will project to different areas of the body (Figure 2). The location of referred pain is not random, but a definitive explanation of the mechanism has not been established. The most broadly accepted theory for this phenomenon is that the visceral sensory fibers enter into the same level of the spinal cord as the somatosensory fibers of the referred pain location. By this explanation, the visceral sensory fibers from the mediastinal region, where the heart is located, would enter the spinal cord at the same level as the spinal nerves from the shoulder and arm, so the brain misinterprets the sensations from the mediastinal region as being from the axillary and brachial regions. Projections from the medial and inferior divisions of the cervical ganglia do enter the spinal cord at the middle to lower cervical levels, which is where the somatosensory fibers enter.

[caption id="" align="aligncenter" width="520"]The figure shows the different organs in the human body. The left panel shows the front view, and the right panel shows the back view. Figure 2. Referred Pain Chart. Conscious perception of visceral sensations map to specific regions of the body, as shown in this chart. Some sensations are felt locally, whereas others are perceived as affecting areas that are quite distant from the involved organ.[/caption]

Efferent Branch

The efferent branch of the visceral reflex arc begins with the projection from the central neuron along the preganglionic fiber. This fiber then makes a synapse on the ganglionic neuron that projects to the target effector.

The effector organs that are the targets of the autonomic system range from the iris and ciliary body of the eye to the urinary bladder and reproductive organs. The thoracolumbar output, through the various sympathetic ganglia, reaches all of these organs. The cranial component of the parasympathetic system projects from the eye to part of the intestines. The sacral component picks up with the majority of the large intestine and the pelvic organs of the urinary and reproductive systems.

Short and Long Reflexes

Somatic reflexes involve sensory neurons that connect sensory receptors to the CNS and motor neurons that project back out to the skeletal muscles. Visceral reflexes that involve the thoracolumbar or craniosacral systems share similar connections. However, there are reflexes that do not need to involve any CNS components. A long reflex has afferent branches that enter the spinal cord or brain and involve the efferent branches, as previously explained. A short reflex is completely peripheral and only involves the local integration of sensory input with motor output (Figure 3).

[caption id="" align="aligncenter" width="520"]The top panel in this figure shows a long reflex, where the spinal cord is connected to the sensory receptor cell and the peripheral ganglion. The bottom panel shows a short reflex, where the sensory receptor cell is directly connected to the peripheral ganglion. Figure 3. Short and Long Reflexes. Sensory input can stimulate either a short or a long reflex. A sensory neuron can project to the CNS or to an autonomic ganglion. The short reflex involves the direct stimulation of a postganglionic fiber by the sensory neuron, whereas the long reflex involves integration in the spinal cord or brain.[/caption]

The difference between short and long reflexes is in the involvement of the CNS. Somatic reflexes always involve the CNS, even in a monosynaptic reflex in which the sensory neuron directly activates the motor neuron. That synapse is in the spinal cord or brain stem, so it has to involve the CNS. However, in the autonomic system there is the possibility that the CNS is not involved. Because the efferent branch of a visceral reflex involves two neurons—the central neuron and the ganglionic neuron—a “short circuit” can be possible. If a sensory neuron projects directly to the ganglionic neuron and causes it to activate the effector target, then the CNS is not involved.

A division of the nervous system that is related to the autonomic nervous system is the enteric nervous system. The word enteric refers to the digestive organs, so this represents the nervous tissue that is part of the digestive system. There are a few myenteric plexuses in which the nervous tissue in the wall of the digestive tract organs can directly influence digestive function. If stretch receptors in the stomach are activated by the filling and distension of the stomach, a short reflex will directly activate the smooth muscle fibers of the stomach wall to increase motility to digest the excessive food in the stomach. No CNS involvement is needed because the stretch receptor is directly activating a neuron in the wall of the stomach that causes the smooth muscle to contract. That neuron, connected to the smooth muscle, is a postganglionic parasympathetic neuron that can be controlled by a fiber found in the vagus nerve.

[caption id="" align="aligncenter" width="120"]QR Code representing a URL Read this article to learn about a teenager who experiences a series of spells that suggest a stroke.[/caption]

Balance in Competing Autonomic Reflex Arcs

The autonomic nervous system is important for homeostasis because its two divisions compete at the target effector. The balance of homeostasis is attributable to the competing inputs from the sympathetic and parasympathetic divisions (dual innervation). At the level of the target effector, the signal of which system is sending the message is strictly chemical. A signaling molecule binds to a receptor that causes changes in the target cell, which in turn causes the tissue or organ to respond to the changing conditions of the body.

Competing Neurotransmitters

The postganglionic fibers of the sympathetic and parasympathetic divisions both release neurotransmitters that bind to receptors on their targets. Postganglionic sympathetic fibers release norepinephrine, with a minor exception, whereas postganglionic parasympathetic fibers release ACh. For any given target, the difference in which division of the autonomic nervous system is exerting control is just in what chemical binds to its receptors. The target cells will have adrenergic and muscarinic receptors. If norepinephrine is released, it will bind to the adrenergic receptors present on the target cell, and if ACh is released, it will bind to the muscarinic receptors on the target cell.

In the sympathetic system, there are exceptions to this pattern of dual innervation. The postganglionic sympathetic fibers that contact the blood vessels within skeletal muscle and that contact sweat glands do not release norepinephrine, they release ACh. This does not create any problem because there is no parasympathetic input to the sweat glands. Sweat glands have muscarinic receptors and produce and secrete sweat in response to the presence of ACh.

At most of the other targets of the autonomic system, the effector response is based on which neurotransmitter is released and what receptor is present. For example, regions of the heart that establish heart rate are contacted by postganglionic fibers from both systems. If norepinephrine is released onto those cells, it binds to an adrenergic receptor that causes the cells to depolarize faster, and the heart rate increases. If ACh is released onto those cells, it binds to a muscarinic receptor that causes the cells to hyperpolarize so that they cannot reach threshold as easily, and the heart rate slows. Without this parasympathetic input, the heart would work at a rate of approximately 100 beats per minute (bpm). The sympathetic system speeds that up, as it would during exercise, to 120–140 bpm, for example. The parasympathetic system slows it down to the resting heart rate of 60–80 bpm.

Another example is in the control of pupillary size (Figure 4). The afferent branch responds to light hitting the retina. Photoreceptors are activated, and the signal is transferred to the retinal ganglion cells that send an action potential along the optic nerve into the diencephalon. If light levels are low, the sympathetic system sends a signal out through the upper thoracic spinal cord to the superior cervical ganglion of the sympathetic chain. The postganglionic fiber then projects to the iris, where it releases norepinephrine onto the radial fibers of the iris (a smooth muscle). When those fibers contract, the pupil dilates—increasing the amount of light hitting the retina. If light levels are too high, the parasympathetic system sends a signal out from the Eddinger–Westphal nucleus through the oculomotor nerve. This fiber synapses in the ciliary ganglion in the posterior orbit. The postganglionic fiber then projects to the iris, where it releases ACh onto the circular fibers of the iris—another smooth muscle. When those fibers contract, the pupil constricts to limit the amount of light hitting the retina.

[caption id="" align="aligncenter" width="580"]This diagram shows how the size of pupils is regulated. The top level of this image shows an eye and the axons going to the brain via the optic nerve. The second level shows the effects of dim light, which results in pupils dilating. The third level shows the effect of bright light, which results in pupils constricting. Figure 4. Autonomic Control of Pupillary Size. Activation of the pupillary reflex comes from the amount of light activating the retinal ganglion cells, as sent along the optic nerve. The output of the sympathetic system projects through the superior cervical ganglion, whereas the parasympathetic system originates out of the midbrain and projects through the oculomotor nerve to the ciliary ganglion, which then projects to the iris. The postganglionic fibers of either division release neurotransmitters onto the smooth muscles of the iris to cause changes in the pupillary size. Norepinephrine results in dilation and ACh results in constriction.[/caption]

In this example, the autonomic system is controlling how much light hits the retina. It is a homeostatic reflex mechanism that keeps the activation of photoreceptors within certain limits. In the context of avoiding a threat like the lioness on the savannah, the sympathetic response for fight or flight will increase pupillary diameter so that more light hits the retina and more visual information is available for running away. Likewise, the parasympathetic response of rest reduces the amount of light reaching the retina, allowing the photoreceptors to cycle through bleaching and be regenerated for further visual perception; this is what the homeostatic process is attempting to maintain.

Autonomic Tone

Organ systems are balanced between the input from the sympathetic and parasympathetic divisions. When something upsets that balance, the homeostatic mechanisms strive to return it to its regular state. For each organ system, there may be more of a sympathetic or parasympathetic tendency to the resting state, which is known as the autonomic tone of the system. For example, the heart rate was described above. Because the resting heart rate is the result of the parasympathetic system slowing the heart down from its intrinsic rate of 100 bpm, the heart can be said to be in parasympathetic tone.

In a similar fashion, another aspect of the cardiovascular system is primarily under sympathetic control. Blood pressure is partially determined by the contraction of smooth muscle in the walls of blood vessels. These tissues have adrenergic receptors that respond to the release of norepinephrine from postganglionic sympathetic fibers by constricting and increasing blood pressure. The hormones released from the adrenal medulla—epinephrine and norepinephrine—will also bind to these receptors. Those hormones travel through the bloodstream where they can easily interact with the receptors in the vessel walls. The parasympathetic system has no significant input to the systemic blood vessels, so the sympathetic system determines their tone.

There are a limited number of blood vessels that respond to sympathetic input in a different fashion. Blood vessels in skeletal muscle, particularly those in the lower limbs, are more likely to dilate. It does not have an overall effect on blood pressure to alter the tone of the vessels, but rather allows for blood flow to increase for those skeletal muscles that will be active in the fight-or-flight response. The blood vessels that have a parasympathetic projection are limited to those in the erectile tissue of the reproductive organs. Acetylcholine released by these postganglionic parasympathetic fibers cause the vessels to dilate, leading to the engorgement of the erectile tissue.

Homeostatic Imbalances

Orthostatic Hypotension Have you ever stood up quickly and felt dizzy for a moment? This is because, for one reason or another, blood is not getting to your brain so it is briefly deprived of oxygen. When you change position from sitting or lying down to standing, your cardiovascular system has to adjust for a new challenge, keeping blood pumping up into the head while gravity is pulling more and more blood down into the legs.

The reason for this is a sympathetic reflex that maintains the output of the heart in response to postural change. When a person stands up, proprioceptors indicate that the body is changing position. A signal goes to the CNS, which then sends a signal to the upper thoracic spinal cord neurons of the sympathetic division. The sympathetic system then causes the heart to beat faster and the blood vessels to constrict. Both changes will make it possible for the cardiovascular system to maintain the rate of blood delivery to the brain. Blood is being pumped superiorly through the internal branch of the carotid arteries into the brain, against the force of gravity. Gravity is not increasing while standing, but blood is more likely to flow down into the legs as they are extended for standing. This sympathetic reflex keeps the brain well oxygenated so that cognitive and other neural processes are not interrupted.

Sometimes this does not work properly. If the sympathetic system cannot increase cardiac output, then blood pressure into the brain will decrease, and a brief neurological loss can be felt. This can be brief, as a slight “wooziness” when standing up too quickly, or a loss of balance and neurological impairment for a period of time. The name for this is orthostatic hypotension, which means that blood pressure goes below the homeostatic set point when standing. It can be the result of standing up faster than the reflex can occur, which may be referred to as a benign “head rush,” or it may be the result of an underlying cause.

There are two basic reasons that orthostatic hypotension can occur. First, blood volume is too low and the sympathetic reflex is not effective. This hypovolemia may be the result of dehydration or medications that affect fluid balance, such as diuretics or vasodilators. Both of these medications are meant to lower blood pressure, which may be necessary in the case of systemic hypertension, and regulation of the medications may alleviate the problem. Sometimes increasing fluid intake or water retention through salt intake can improve the situation.

The second underlying cause of orthostatic hypotension is autonomic failure. There are several disorders that result in compromised sympathetic functions. The disorders range from diabetes to multiple system atrophy (a loss of control over many systems in the body), and addressing the underlying condition can improve the hypotension. For example, with diabetes, peripheral nerve damage can occur, which would affect the postganglionic sympathetic fibers. Getting blood glucose levels under control can improve neurological deficits associated with diabetes.

]]> 2580 2569 2 0 15.3 Central Control https://pressbooks.bccampus.ca/apdouglas2017/chapter/15-3-central-control/ Wed, 02 Aug 2017 22:03:20 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2584

Learning Objectives

By the end of this section, you will be able to:
  • Describe the role of higher centers of the brain in autonomic regulation
  • Explain the connection of the hypothalamus to homeostasis
  • Describe the regions of the CNS that link the autonomic system with emotion
  • Describe the pathways important to descending control of the autonomic system

The pupillary light reflex (Figure 1) begins when light hits the retina and causes a signal to travel along the optic nerve. This is visual sensation, because the afferent branch of this reflex is simply sharing the special sense pathway. Bright light hitting the retina leads to the parasympathetic response, through the oculomotor nerve, followed by the postganglionic fiber from the ciliary ganglion, which stimulates the circular fibers of the iris to contract and constrict the pupil. When light hits the retina in one eye, both pupils contract. When that light is removed, both pupils dilate again back to the resting position. When the stimulus is unilateral (presented to only one eye), the response is bilateral (both eyes). The same is not true for somatic reflexes. If you touch a hot radiator, you only pull that arm back, not both. Central control of autonomic reflexes is different than for somatic reflexes. The hypothalamus, along with other CNS locations, controls the autonomic system.

[caption id="" align="aligncenter" width="530"]This diagram shows the connections between the different nerves and pathways in the eyes. A hand is shown shining a light on the right eye, and arrows and text callouts indicate the different pathways that are activated. Figure 1. Pupillary Reflex Pathways. The pupil is under competing autonomic control in response to light levels hitting the retina. The sympathetic system will dilate the pupil when the retina is not receiving enough light, and the parasympathetic system will constrict the pupil when too much light hits the retina.[/caption]

Forebrain Structures

Autonomic control is based on the visceral reflexes, composed of the afferent and efferent branches. These homeostatic mechanisms are based on the balance between the two divisions of the autonomic system, which results in tone for various organs that is based on the predominant input from the sympathetic or parasympathetic systems. Coordinating that balance requires integration that begins with forebrain structures like the hypothalamus and continues into the brain stem and spinal cord.

The Hypothalamus

The hypothalamus is the control center for many homeostatic mechanisms. It regulates both autonomic function and endocrine function. The roles it plays in the pupillary reflexes demonstrates the importance of this control center. The optic nerve projects primarily to the thalamus, which is the necessary relay to the occipital cortex for conscious visual perception. Another projection of the optic nerve, however, goes to the hypothalamus.

The hypothalamus then uses this visual system input to drive the pupillary reflexes. If the retina is activated by high levels of light, the hypothalamus stimulates the parasympathetic response. If the optic nerve message shows that low levels of light are falling on the retina, the hypothalamus activates the sympathetic response. Output from the hypothalamus follows two main tracts, the dorsal longitudinal fasciculus and the medial forebrain bundle (Figure 2). Along these two tracts, the hypothalamus can influence the Eddinger–Westphal nucleus of the oculomotor complex or the lateral horns of the thoracic spinal cord.

[caption id="" align="aligncenter" width="480"]This figure shows the human brain on the left panel, and a magnified image shows the location of the medial forebrain bundle and the dorsal longitudinal fasciculus in the brain. Figure 2. Fiber Tracts of the Central Autonomic System. The hypothalamus is the source of most of the central control of autonomic function. It receives input from cerebral structures and projects to brain stem and spinal cord structures to regulate the balance of sympathetic and parasympathetic input to the organ systems of the body. The main pathways for this are the medial forebrain bundle and the dorsal longitudinal fasciculus.[/caption]
These two tracts connect the hypothalamus with the major parasympathetic nuclei in the brain stem and the preganglionic (central) neurons of the thoracolumbar spinal cord. The hypothalamus also receives input from other areas of the forebrain through the medial forebrain bundle. The olfactory cortex, the septal nuclei of the basal forebrain, and the amygdala project into the hypothalamus through the medial forebrain bundle. These forebrain structures inform the hypothalamus about the state of the nervous system and can influence the regulatory processes of homeostasis. A good example of this is found in the amygdala, which is found beneath the cerebral cortex of the temporal lobe and plays a role in our ability to remember and feel emotions.

The Amygdala

The amygdala is a group of nuclei in the medial region of the temporal lobe that is part of the limbic lobe (Figure 3). The limbic lobe includes structures that are involved in emotional responses, as well as structures that contribute to memory function. The limbic lobe has strong connections with the hypothalamus and influences the state of its activity on the basis of emotional state. For example, when you are anxious or scared, the amygdala will send signals to the hypothalamus along the medial forebrain bundle that will stimulate the sympathetic fight-or-flight response. The hypothalamus will also stimulate the release of stress hormones through its control of the endocrine system in response to amygdala input.

[caption id="" align="aligncenter" width="510"]This figure shows the location of the limbic lobe and its major parts in the human brain. Figure 3. The Limbic Lobe. Structures arranged around the edge of the cerebrum constitute the limbic lobe, which includes the amygdala, hippocampus, and cingulate gyrus, and connects to the hypothalamus.[/caption]

The Medulla

The medulla contains nuclei referred to as the cardiovascular center, which controls the smooth and cardiac muscle of the cardiovascular system through autonomic connections. When the homeostasis of the cardiovascular system shifts, such as when blood pressure changes, the coordination of the autonomic system can be accomplished within this region. Furthermore, when descending inputs from the hypothalamus stimulate this area, the sympathetic system can increase activity in the cardiovascular system, such as in response to anxiety or stress. The preganglionic sympathetic fibers that are responsible for increasing heart rate are referred to as the cardiac accelerator nerves, whereas the preganglionic sympathetic fibers responsible for constricting blood vessels compose the vasomotor nerves.

Several brain stem nuclei are important for the visceral control of major organ systems. One brain stem nucleus involved in cardiovascular function is the solitary nucleus. It receives sensory input about blood pressure and cardiac function from the glossopharyngeal and vagus nerves, and its output will activate sympathetic stimulation of the heart or blood vessels through the upper thoracic lateral horn. Another brain stem nucleus important for visceral control is the dorsal motor nucleus of the vagus nerve, which is the motor nucleus for the parasympathetic functions ascribed to the vagus nerve, including decreasing the heart rate, relaxing bronchial tubes in the lungs, and activating digestive function through the enteric nervous system. The nucleus ambiguus, which is named for its ambiguous histology, also contributes to the parasympathetic output of the vagus nerve and targets muscles in the pharynx and larynx for swallowing and speech, as well as contributing to the parasympathetic tone of the heart along with the dorsal motor nucleus of the vagus.

]]> 2584 2569 3 0 20.6 Development of Blood Vessels and Fetal Circulation https://pressbooks.bccampus.ca/apdouglas2017/chapter/20-6-development-of-blood-vessels-and-fetal-circulation/ Wed, 02 Aug 2017 22:03:37 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2617

Learning Objectives

By the end of this section, you will be able to:
  • Describe the development of blood vessels
  • Describe the fetal circulation

In a developing embryo,the heart has developed enough by day 21 post-fertilization to begin beating. Circulation patterns are clearly established by the fourth week of embryonic life. It is critical to the survival of the developing human that the circulatory system forms early to supply the growing tissue with nutrients and gases, and to remove waste products. Blood cells and vessel production in structures outside the embryo proper called the yolk sac, chorion, and connecting stalk begin about 15 to 16 days following fertilization. Development of these circulatory elements within the embryo itself begins approximately 2 days later. You will learn more about the formation and function of these early structures when you study the chapter on development. During those first few weeks, blood vessels begin to form from the embryonic mesoderm. The precursor cells are known as hemangioblasts. These in turn differentiate into angioblasts, which give rise to the blood vessels and pluripotent stem cells, which differentiate into the formed elements of blood. (Seek additional content for more detail on fetal development and circulation.) Together, these cells form masses known as blood islands scattered throughout the embryonic disc. Spaces appear on the blood islands that develop into vessel lumens. The endothelial lining of the vessels arise from the angioblasts within these islands. Surrounding mesenchymal cells give rise to the smooth muscle and connective tissue layers of the vessels. While the vessels are developing, the pluripotent stem cells begin to form the blood.

Vascular tubes also develop on the blood islands, and they eventually connect to one another as well as to the developing, tubular heart. Thus, the developmental pattern, rather than beginning from the formation of one central vessel and spreading outward, occurs in many regions simultaneously with vessels later joining together. This angiogenesis—the creation of new blood vessels from existing ones—continues as needed throughout life as we grow and develop.

Blood vessel development often follows the same pattern as nerve development and travels to the same target tissues and organs. This occurs because the many factors directing growth of nerves also stimulate blood vessels to follow a similar pattern. Whether a given vessel develops into an artery or a vein is dependent upon local concentrations of signaling proteins.

As the embryo grows within the mother’s uterus, its requirements for nutrients and gas exchange also grow. The placenta—a circulatory organ unique to pregnancy—develops jointly from the embryo and uterine wall structures to fill this need. Emerging from the placenta is the umbilical vein, which carries oxygen-rich blood from the mother to the fetal inferior vena cava via the ductus venosus to the heart that pumps it into fetal circulation. Two umbilical arteries carry oxygen-depleted fetal blood, including wastes and carbon dioxide, to the placenta. Remnants of the umbilical arteries remain in the adult. (Seek additional content for more information on the role of the placenta in fetal circulation.)

There are three major shunts—alternate paths for blood flow—found in the circulatory system of the fetus. Two of these shunts divert blood from the pulmonary to the systemic circuit, whereas the third connects the umbilical vein to the inferior vena cava. The first two shunts are critical during fetal life, when the lungs are compressed, filled with amniotic fluid, and nonfunctional, and gas exchange is provided by the placenta. These shunts close shortly after birth, however, when the newborn begins to breathe. The third shunt persists a bit longer but becomes nonfunctional once the umbilical cord is severed. The three shunts are as follows (Figure 1):
  • The foramen ovale is an opening in the interatrial septum that allows blood to flow from the right atrium to the left atrium. A valve associated with this opening prevents backflow of blood during the fetal period. As the newborn begins to breathe and blood pressure in the atria increases, this shunt closes. The fossa ovalis remains in the interatrial septum after birth, marking the location of the former foramen ovale.
  • The ductus arteriosus is a short, muscular vessel that connects the pulmonary trunk to the aorta. Most of the blood pumped from the right ventricle into the pulmonary trunk is thereby diverted into the aorta. Only enough blood reaches the fetal lungs to maintain the developing lung tissue. When the newborn takes the first breath, pressure within the lungs drops dramatically, and both the lungs and the pulmonary vessels expand. As the amount of oxygen increases, the smooth muscles in the wall of the ductus arteriosus constrict, sealing off the passage. Eventually, the muscular and endothelial components of the ductus arteriosus degenerate, leaving only the connective tissue component of the ligamentum arteriosum.
  • The ductus venosus is a temporary blood vessel that branches from the umbilical vein, allowing much of the freshly oxygenated blood from the placenta—the organ of gas exchange between the mother and fetus—to bypass the fetal liver and go directly to the fetal heart. The ductus venosus closes slowly during the first weeks of infancy and degenerates to become the ligamentum venosum.
[caption id="" align="aligncenter" width="495"]This figure shows the blood vessels in a fetus. Figure 1. Fetal Shunts. The foramen ovale in the interatrial septum allows blood to flow from the right atrium to the left atrium. The ductus arteriosus is a temporary vessel, connecting the aorta to the pulmonary trunk. The ductus venosus links the umbilical vein to the inferior vena cava largely through the liver.[/caption]
]]> 2617 2615 1 0 22.4 Gas Exchange https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-4-gas-exchange-2/ Wed, 02 Aug 2017 22:03:54 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2622

Learning Objectives

By the end of this section, you will be able to:
  • Compare the composition of atmospheric air and alveolar air
  • Describe the mechanisms that drive gas exchange
  • Discuss the importance of sufficient ventilation and perfusion, and how the body adapts when they are insufficient
  • Discuss the process of external respiration
  • Describe the process of internal respiration
The purpose of the respiratory system is to perform gas exchange. Pulmonary ventilation provides air to the alveoli for this gas exchange process. At the respiratory membrane, where the alveolar and capillary walls meet, gases move across the membranes, with oxygen entering the bloodstream and carbon dioxide exiting. It is through this mechanism that blood is oxygenated and carbon dioxide, the waste product of cellular respiration, is removed from the body.

Gas Exchange

In order to understand the mechanisms of gas exchange in the lung, it is important to understand the underlying principles of gases and their behavior. In addition to Boyle’s law, several other gas laws help to describe the behavior of gases.

Gas Laws and Air Composition

Gas molecules exert force on the surfaces with which they are in contact; this force is called pressure. In natural systems, gases are normally present as a mixture of different types of molecules. For example, the atmosphere consists of oxygen, nitrogen, carbon dioxide, and other gaseous molecules, and this gaseous mixture exerts a certain pressure referred to as atmospheric pressure (Table 2). Partial pressure (Px) is the pressure of a single type of gas in a mixture of gases. For example, in the atmosphere, oxygen exerts a partial pressure, and nitrogen exerts another partial pressure, independent of the partial pressure of oxygen (Figure 1). Total pressure is the sum of all the partial pressures of a gaseous mixture. Dalton’s law describes the behavior of nonreactive gases in a gaseous mixture and states that a specific gas type in a mixture exerts its own pressure; thus, the total pressure exerted by a mixture of gases is the sum of the partial pressures of the gases in the mixture.
Partial Pressures of Atmospheric Gases (Table 2)
Gas Percent of total composition Partial pressure
(mm Hg)
Nitrogen (N2) 78.6 597.4
Oxygen (O2) 20.9 158.8
Water (H2O) 0.04 3.0
Carbon dioxide (CO2) 0.004 0.3
Others 0.0006 0.5
Total composition/total atmospheric pressure 100% 760.0
[caption id="" align="aligncenter" width="420"]The left panel of this figure shows a canister of oxygen. The middle panel shows a canister of nitrogen. The right panel shows a canister containing a mixture of oxygen and nitrogen. A pressure gauge on each container shows the pressure exerted by the gas in that container. Figure 1. Partial and Total Pressures of a Gas. Partial pressure is the force exerted by a gas. The sum of the partial pressures of all the gases in a mixture equals the total pressure.[/caption]
Partial pressure is extremely important in predicting the movement of gases. Recall that gases tend to equalize their pressure in two regions that are connected. A gas will move from an area where its partial pressure is higher to an area where its partial pressure is lower. In addition, the greater the partial pressure difference between the two areas, the more rapid is the movement of gases.

Solubility of Gases in Liquids

Henry’s law describes the behavior of gases when they come into contact with a liquid, such as blood. Henry’s law states that the concentration of gas in a liquid is directly proportional to the solubility and partial pressure of that gas. The greater the partial pressure of the gas, the greater the number of gas molecules that will dissolve in the liquid. The concentration of the gas in a liquid is also dependent on the solubility of the gas in the liquid. For example, although nitrogen is present in the atmosphere, very little nitrogen dissolves into the blood, because the solubility of nitrogen in blood is very low. The exception to this occurs in scuba divers; the composition of the compressed air that divers breathe causes nitrogen to have a higher partial pressure than normal, causing it to dissolve in the blood in greater amounts than normal. Too much nitrogen in the bloodstream results in a serious condition that can be fatal if not corrected. Gas molecules establish an equilibrium between those molecules dissolved in liquid and those in air. The composition of air in the atmosphere and in the alveoli differs. In both cases, the relative concentration of gases is nitrogen > oxygen > water vapor > carbon dioxide. The amount of water vapor present in alveolar air is greater than that in atmospheric air (Table 3). Recall that the respiratory system works to humidify incoming air, thereby causing the air present in the alveoli to have a greater amount of water vapor than atmospheric air. In addition, alveolar air contains a greater amount of carbon dioxide and less oxygen than atmospheric air. This is no surprise, as gas exchange removes oxygen from and adds carbon dioxide to alveolar air. Both deep and forced breathing cause the alveolar air composition to be changed more rapidly than during quiet breathing. As a result, the partial pressures of oxygen and carbon dioxide change, affecting the diffusion process that moves these materials across the membrane. This will cause oxygen to enter and carbon dioxide to leave the blood more quickly.
Composition and Partial Pressures of Alveolar Air (Table 3)
Gas Percent of total composition Partial pressure
(mm Hg)
Nitrogen (N2) 74.9 569
Oxygen (O2) 13.7 104
Water (H2O) 6.2 40
Carbon dioxide (CO2) 5.2 47
Total composition/total alveolar pressure 100% 760.0

Ventilation and Perfusion

Two important aspects of gas exchange in the lung are ventilation and perfusion. Ventilation is the movement of air into and out of the lungs, and perfusion is the flow of blood in the pulmonary capillaries. For gas exchange to be efficient, the volumes involved in ventilation and perfusion should be compatible. However, factors such as regional gravity effects on blood, blocked alveolar ducts, or disease can cause ventilation and perfusion to be imbalanced. The partial pressure of oxygen in alveolar air is about 104 mm Hg, whereas the partial pressure of the oxygenated pulmonary venous blood is about 100 mm Hg. When ventilation is sufficient, oxygen enters the alveoli at a high rate, and the partial pressure of oxygen in the alveoli remains high. In contrast, when ventilation is insufficient, the partial pressure of oxygen in the alveoli drops. Without the large difference in partial pressure between the alveoli and the blood, oxygen does not diffuse efficiently across the respiratory membrane. The body has mechanisms that counteract this problem. In cases when ventilation is not sufficient for an alveolus, the body redirects blood flow to alveoli that are receiving sufficient ventilation. This is achieved by constricting the pulmonary arterioles that serves the dysfunctional alveolus, which redirects blood to other alveoli that have sufficient ventilation. At the same time, the pulmonary arterioles that serve alveoli receiving sufficient ventilation vasodilate, which brings in greater blood flow. Factors such as carbon dioxide, oxygen, and pH levels can all serve as stimuli for adjusting blood flow in the capillary networks associated with the alveoli. Ventilation is regulated by the diameter of the airways, whereas perfusion is regulated by the diameter of the blood vessels. The diameter of the bronchioles is sensitive to the partial pressure of carbon dioxide in the alveoli. A greater partial pressure of carbon dioxide in the alveoli causes the bronchioles to increase their diameter as will a decreased level of oxygen in the blood supply, allowing carbon dioxide to be exhaled from the body at a greater rate. As mentioned above, a greater partial pressure of oxygen in the alveoli causes the pulmonary arterioles to dilate, increasing blood flow.

Gas Exchange

Gas exchange occurs at two sites in the body: in the lungs, where oxygen is picked up and carbon dioxide is released at the respiratory membrane, and at the tissues, where oxygen is released and carbon dioxide is picked up. External respiration is the exchange of gases with the external environment, and occurs in the alveoli of the lungs. Internal respiration is the exchange of gases with the internal environment, and occurs in the tissues. The actual exchange of gases occurs due to simple diffusion. Energy is not required to move oxygen or carbon dioxide across membranes. Instead, these gases follow pressure gradients that allow them to diffuse. The anatomy of the lung maximizes the diffusion of gases: The respiratory membrane is highly permeable to gases; the respiratory and blood capillary membranes are very thin; and there is a large surface area throughout the lungs.

External Respiration

The pulmonary artery carries deoxygenated blood into the lungs from the heart, where it branches and eventually becomes the capillary network composed of pulmonary capillaries. These pulmonary capillaries create the respiratory membrane with the alveoli (Figure 2). As the blood is pumped through this capillary network, gas exchange occurs. Although a small amount of the oxygen is able to dissolve directly into plasma from the alveoli, most of the oxygen is picked up by erythrocytes (red blood cells) and binds to a protein called hemoglobin, a process described later in this chapter. Oxygenated hemoglobin is red, causing the overall appearance of bright red oxygenated blood, which returns to the heart through the pulmonary veins. Carbon dioxide is released in the opposite direction of oxygen, from the blood to the alveoli. Some of the carbon dioxide is returned on hemoglobin, but can also be dissolved in plasma or is present as a converted form, also explained in greater detail later in this chapter. External respiration occurs as a function of partial pressure differences in oxygen and carbon dioxide between the alveoli and the blood in the pulmonary capillaries.
[caption id="" align="aligncenter" width="425"]This figure shows the pathway in which external respiration takes place. The exchange of oxygen and carbon dioxide between the alveolus and blood plasma is detailed. Figure 3. External Respiration. In external respiration, oxygen diffuses across the respiratory membrane from the alveolus to the capillary, whereas carbon dioxide diffuses out of the capillary into the alveolus.[/caption]
Although the solubility of oxygen in blood is not high, there is a drastic difference in the partial pressure of oxygen in the alveoli versus in the blood of the pulmonary capillaries. This difference is about 64 mm Hg: The partial pressure of oxygen in the alveoli is about 104 mm Hg, whereas its partial pressure in the blood of the capillary is about 40 mm Hg. This large difference in partial pressure creates a very strong pressure gradient that causes oxygen to rapidly cross the respiratory membrane from the alveoli into the blood. The partial pressure of carbon dioxide is also different between the alveolar air and the blood of the capillary. However, the partial pressure difference is less than that of oxygen, about 5 mm Hg. The partial pressure of carbon dioxide in the blood of the capillary is about 45 mm Hg, whereas its partial pressure in the alveoli is about 40 mm Hg. However, the solubility of carbon dioxide is much greater than that of oxygen—by a factor of about 20—in both blood and alveolar fluids. As a result, the relative concentrations of oxygen and carbon dioxide that diffuse across the respiratory membrane are similar.

Internal Respiration

Internal respiration is gas exchange that occurs at the level of body tissues (Figure 3). Similar to external respiration, internal respiration also occurs as simple diffusion due to a partial pressure gradient. However, the partial pressure gradients are opposite of those present at the respiratory membrane. The partial pressure of oxygen in tissues is low, about 40 mm Hg, because oxygen is continuously used for cellular respiration. In contrast, the partial pressure of oxygen in the blood is about 100 mm Hg. This creates a pressure gradient that causes oxygen to dissociate from hemoglobin, diffuse out of the blood, cross the interstitial space, and enter the tissue. Hemoglobin that has little oxygen bound to it loses much of its brightness, so that blood returning to the heart is more burgundy in color. Considering that cellular respiration continuously produces carbon dioxide, the partial pressure of carbon dioxide is lower in the blood than it is in the tissue, causing carbon dioxide to diffuse out of the tissue, cross the interstitial fluid, and enter the blood. It is then carried back to the lungs either bound to hemoglobin, dissolved in plasma, or in a converted form. By the time blood returns to the heart, the partial pressure of oxygen has returned to about 40 mm Hg, and the partial pressure of carbon dioxide has returned to about 45 mm Hg. The blood is then pumped back to the lungs to be oxygenated once again during external respiration.
[caption id="" align="aligncenter" width="425"]This diagram details the pathway of internal respiration. The exchange of oxygen and carbon dioxide between a red blood cell and a tissue cell is shown. Figure 4. Internal Respiration. Oxygen diffuses out of the capillary and into cells, whereas carbon dioxide diffuses out of cells and into the capillary.[/caption]
]]> 2622 2618 1 0 22.6 Modifications in Respiratory Functions https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-6-modifications-in-respiratory-functions-2/ Wed, 02 Aug 2017 22:04:22 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2631

Learning Objectives

By the end of this section, you will be able to:
  • Define the terms hyperpnea and hyperventilation
  • Describe the effect of exercise on the respiratory system
  • Describe the effect of high altitude on the respiratory system
  • Discuss the process of acclimatization
At rest, the respiratory system performs its functions at a constant, rhythmic pace, as regulated by the respiratory centers of the brain. At this pace, ventilation provides sufficient oxygen to all the tissues of the body. However, there are times that the respiratory system must alter the pace of its functions in order to accommodate the oxygen demands of the body.

Hyperpnea

Hyperpnea is an increased depth and rate of ventilation to meet an increase in oxygen demand as might be seen in exercise or disease, particularly diseases that target the respiratory or digestive tracts. This does not significantly alter blood oxygen or carbon dioxide levels, but merely increases the depth and rate of ventilation to meet the demand of the cells. In contrast, hyperventilation is an increased ventilation rate that is independent of the cellular oxygen needs and leads to abnormally low blood carbon dioxide levels and high (alkaline) blood pH. Interestingly, exercise does not cause hyperpnea as one might think. Muscles that perform work during exercise do increase their demand for oxygen, stimulating an increase in ventilation. However, hyperpnea during exercise appears to occur before a drop in oxygen levels within the muscles can occur. Therefore, hyperpnea must be driven by other mechanisms, either instead of or in addition to a drop in oxygen levels. The exact mechanisms behind exercise hyperpnea are not well understood, and some hypotheses are somewhat controversial. However, in addition to low oxygen, high carbon dioxide, and low pH levels, there appears to be a complex interplay of factors related to the nervous system and the respiratory centers of the brain. First, a conscious decision to partake in exercise, or another form of physical exertion, results in a psychological stimulus that may trigger the respiratory centers of the brain to increase ventilation. In addition, the respiratory centers of the brain may be stimulated through the activation of motor neurons that innervate muscle groups that are involved in the physical activity. Finally, physical exertion stimulates proprioceptors, which are receptors located within the muscles, joints, and tendons, which sense movement and stretching; proprioceptors thus create a stimulus that may also trigger the respiratory centers of the brain. These neural factors are consistent with the sudden increase in ventilation that is observed immediately as exercise begins. Because the respiratory centers are stimulated by psychological, motor neuron, and proprioceptor inputs throughout exercise, the fact that there is also a sudden decrease in ventilation immediately after the exercise ends when these neural stimuli cease, further supports the idea that they are involved in triggering the changes of ventilation.

High Altitude Effects

An increase in altitude results in a decrease in atmospheric pressure. Although the proportion of oxygen relative to gases in the atmosphere remains at 21 percent, its partial pressure decreases (Table 4). As a result, it is more difficult for a body to achieve the same level of oxygen saturation at high altitude than at low altitude, due to lower atmospheric pressure. In fact, hemoglobin saturation is lower at high altitudes compared to hemoglobin saturation at sea level. For example, hemoglobin saturation is about 67 percent at 19,000 feet above sea level, whereas it reaches about 98 percent at sea level.
Partial Pressure of Oxygen at Different Altitudes (Table 4)
Example location Altitude (feet above sea level) Atmospheric pressure (mm Hg) Partial pressure of oxygen (mm Hg)
New York City, New York 0 760 159
Boulder, Colorado 5000 632 133
Aspen, Colorado 8000 565 118
Pike’s Peak, Colorado 14,000 447 94
Denali (Mt. McKinley), Alaska 20,000 350 73
Mt. Everest, Tibet 29,000 260 54
As you recall, partial pressure is extremely important in determining how much gas can cross the respiratory membrane and enter the blood of the pulmonary capillaries. A lower partial pressure of oxygen means that there is a smaller difference in partial pressures between the alveoli and the blood, so less oxygen crosses the respiratory membrane. As a result, fewer oxygen molecules are bound by hemoglobin. Despite this, the tissues of the body still receive a sufficient amount of oxygen during rest at high altitudes. This is due to two major mechanisms. First, the number of oxygen molecules that enter the tissue from the blood is nearly equal between sea level and high altitudes. At sea level, hemoglobin saturation is higher, but only a quarter of the oxygen molecules are actually released into the tissue. At high altitudes, a greater proportion of molecules of oxygen are released into the tissues. Secondly, at high altitudes, a greater amount of BPG is produced by erythrocytes, which enhances the dissociation of oxygen from hemoglobin. Physical exertion, such as skiing or hiking, can lead to altitude sickness due to the low amount of oxygen reserves in the blood at high altitudes. At sea level, there is a large amount of oxygen reserve in venous blood (even though venous blood is thought of as “deoxygenated”) from which the muscles can draw during physical exertion. Because the oxygen saturation is much lower at higher altitudes, this venous reserve is small, resulting in pathological symptoms of low blood oxygen levels. You may have heard that it is important to drink more water when traveling at higher altitudes than you are accustomed to. This is because your body will increase micturition (urination) at high altitudes to counteract the effects of lower oxygen levels. By removing fluids, blood plasma levels drop but not the total number of erythrocytes. In this way, the overall concentration of erythrocytes in the blood increases, which helps tissues obtain the oxygen they need. Acute mountain sickness (AMS), or altitude sickness, is a condition that results from acute exposure to high altitudes due to a low partial pressure of oxygen at high altitudes. AMS typically can occur at 2400 meters (8000 feet) above sea level. AMS is a result of low blood oxygen levels, as the body has acute difficulty adjusting to the low partial pressure of oxygen. In serious cases, AMS can cause pulmonary or cerebral edema. Symptoms of AMS include nausea, vomiting, fatigue, lightheadedness, drowsiness, feeling disoriented, increased pulse, and nosebleeds. The only treatment for AMS is descending to a lower altitude; however, pharmacologic treatments and supplemental oxygen can improve symptoms. AMS can be prevented by slowly ascending to the desired altitude, allowing the body to acclimate, as well as maintaining proper hydration.

Acclimatization

Especially in situations where the ascent occurs too quickly, traveling to areas of high altitude can cause AMS. Acclimatization is the process of adjustment that the respiratory system makes due to chronic exposure to a high altitude. Over a period of time, the body adjusts to accommodate the lower partial pressure of oxygen. The low partial pressure of oxygen at high altitudes results in a lower oxygen saturation level of hemoglobin in the blood. In turn, the tissue levels of oxygen are also lower. As a result, the kidneys are stimulated to produce the hormone erythropoietin (EPO), which stimulates the production of erythrocytes, resulting in a greater number of circulating erythrocytes in an individual at a high altitude over a long period. With more red blood cells, there is more hemoglobin to help transport the available oxygen. Even though there is low saturation of each hemoglobin molecule, there will be more hemoglobin present, and therefore more oxygen in the blood. Over time, this allows the person to partake in physical exertion without developing AMS.
]]> 2631 2618 3 0 Authors https://pressbooks.bccampus.ca/apdouglas2017/authors/ Mon, 10 Jul 2017 16:28:25 +0000 https://pressbooks.bccampus.ca/apdouglas2017/authors/ ]]> 8 0 0 0 Cover https://pressbooks.bccampus.ca/apdouglas2017/ Mon, 10 Jul 2017 16:28:25 +0000 https://pressbooks.bccampus.ca/apdouglas2017/cover/ ]]> 9 0 0 0 Table of Contents https://pressbooks.bccampus.ca/apdouglas2017/table-of-contents/ Mon, 10 Jul 2017 16:28:25 +0000 https://pressbooks.bccampus.ca/apdouglas2017/table-of-contents/ ]]> 10 0 0 0 About https://pressbooks.bccampus.ca/apdouglas2017/about/ Mon, 10 Jul 2017 16:28:25 +0000 https://pressbooks.bccampus.ca/apdouglas2017/about/ ]]> 11 0 0 0 Buy https://pressbooks.bccampus.ca/apdouglas2017/buy/ Mon, 10 Jul 2017 16:28:25 +0000 https://pressbooks.bccampus.ca/apdouglas2017/buy/ ]]> 12 0 0 0 Access Denied https://pressbooks.bccampus.ca/apdouglas2017/access-denied/ Mon, 10 Jul 2017 16:28:25 +0000 https://pressbooks.bccampus.ca/apdouglas2017/access-denied/ ]]> 13 0 0 0 Book Information https://pressbooks.bccampus.ca/apdouglas2017/?metadata=book-information Mon, 10 Jul 2017 16:28:25 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?metadata=book-information 17 0 1 0 © 1999-2016, Rice University. Except where otherwise noted, content created on this site is licensed under a Creative Commons Attribution License 4.0 License. Under this license, any user of this textbook or the textbook contents herein must provide proper attribution as follows: The OpenStax College name, OpenStax College logo, OpenStax College book covers, OpenStax CNX name, and OpenStax CNX logo are not subject to the creative commons license and may not be reproduced without the prior and express written consent of Rice University. For questions regarding this license, please contact partners@openstaxcollege.org.
  • If you use this textbook as a bibliographic reference, then you should cite it as follows:
    OpenStax, Anatomy & Physiology. OpenStax CNX. Feb 26, 2016 http://cnx.org/contents/14fb4ad7-39a1-4eee-ab6e-3ef2482e3e22@8.24.
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]]> Introduction https://pressbooks.bccampus.ca/apdouglas2017/?post_type=front-matter&p=1830 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=front-matter&p=1830 This is where you can write your introduction.

]]> 1830 0 1 0 Preface https://pressbooks.bccampus.ca/apdouglas2017/?post_type=front-matter&p=1833 Mon, 30 Nov -0001 00:00:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=front-matter&p=1833 Human Anatomy and Physiology is designed for the two-semester anatomy and physiology course taken by life science and allied health students. The textbook follows the scope and sequence of most Human Anatomy and Physiology courses, and its coverage and organization were informed by hundreds of instructors who teach the course. Instructors can customize the book, adapting it to the approach that works best in their classroom. The artwork for this textbook is aimed focusing student learning through a powerful blend of traditional depictions and instructional innovations. Color is used sparingly, to emphasize the most important aspects of any given illustration. Significant use of micrographs from the University of Michigan complement the illustrations, and provide the students with a meaningful alternate depiction of each concept. Finally, enrichment elements provide relevance and deeper context for students, particularly in the areas of health, disease, and information relevant to their intended careers.

Welcome to Anatomy and Physiology, an OpenStax resource. We created this textbook with several goals in mind: accessibility, customization, and student engagement—helping students reach high levels of academic scholarship. Instructors and students alike will find that this textbook offers a thorough introduction to the content in an accessible format.

About OpenStax

OpenStax is a nonprofit organization committed to improving student access to quality learning materials. Our free textbooks are developed and peer-reviewed by educators to ensure that they are readable, accurate, and organized in accordance with the scope and sequence requirements of today’s college courses. Unlike traditional textbooks, OpenStax resources live online and are owned by the community of educators using them. Through partnerships with companies and foundations committed to reducing costs for students, we are working to improve access to higher education for all. OpenStax is an initiative of Rice University and is made possible through the generous support of several philanthropic foundations.

About OpenStax’s Resources

OpenStax resources provide quality academic instruction. Three key features set our materials apart from others: 1) They can be easily customized by instructors for each class, 2) they are “living” resources that grow online through contributions from science educators, and 3) they are available for free or for a minimal cost.

Customization

OpenStax learning resources are conceived and written with flexibility in mind so that they can be customized for each course. Our textbooks provide a solid foundation on which instructors can build their own texts. Instructors can select the sections that are most relevant to their curricula and create a textbook that speaks directly to the needs of their students. Instructors are encouraged to expand on existing examples in the text by adding unique context via geographically localized applications and topical connections.

Anatomy and Physiology can be easily customized using our online platform (https://openstaxcollege.org/textbooks/anatomy-and-physiology/adapt). The text is arranged in a modular chapter format. Simply select the content most relevant to your syllabus and create a textbook that addresses the needs of your class. This customization feature will ensure that your textbook reflects the goals of your course.

Curation

To broaden access and encourage community curation, Anatomy and Physiology is “open source” under a Creative Commons Attribution (CC BY) license. Members of the scientific community are invited to submit examples, emerging research, and other feedback to enhance and strengthen the material, keeping it current and relevant for today’s students. You can submit your suggestions to info@openstaxcollege.org.

Cost

Our textbooks are available for free online, and in low-cost print and tablet editions.

About Anatomy and Physiology

Anatomy and Physiology is designed for the two-semester anatomy and physiology course taken by life science and allied health students. It supports effective teaching and learning, and prepares students for further learning and future careers. The text focuses on the most important concepts and aims to minimize distracting students with more minor details.

The development choices for this textbook were made with the guidance of hundreds of faculty who are deeply involved in teaching this course. These choices led to innovations in art, terminology, career orientation, practical applications, and multimedia-based learning, all with a goal of increasing relevance to students. We strove to make the discipline meaningful and memorable to students, so that they can draw from it a working knowledge that will enrich their future studies.

Coverage and Scope

The units of our Anatomy and Physiology textbook adhere to the scope and sequence followed by most two-semester courses nationwide.

Unit 1: Levels of Organization

Chapters 1–4 provide students with a basic understanding of human anatomy and physiology, including its language, the levels of organization, and the basics of chemistry and cell biology. These chapters provide a foundation for the further study of the body. They also focus particularly on how the body’s regions, important chemicals, and cells maintain homeostasis.

Chapter 1 An Introduction to the Human Body Chapter 2 The Chemical Level of Organization Chapter 3 The Cellular Level of Organization Chapter 4 The Tissue Level of Organization

Unit 2: Support and Movement

In Chapters 5–11, students explore the skin, the largest organ of the body, and examine the body’s skeletal and muscular systems, following a traditional sequence of topics. This unit is the first to walk students through specific systems of the body, and as it does so, it maintains a focus on homeostasis as well as those diseases and conditions that can disrupt it.

Chapter 5 The Integumentary System Chapter 6 Bone and Skeletal Tissue Chapter 7 The Axial Skeleton Chapter 8 The Appendicular Skeleton Chapter 9 Joints Chapter 10 Muscle Tissue Chapter 11 The Muscular System

Unit 3: Regulation, Integration, and Control

Chapters 12–17 help students answer questions about nervous and endocrine system control and regulation. In a break with the traditional sequence of topics, the special senses are integrated into the chapter on the somatic nervous system. The chapter on the neurological examination offers students a unique approach to understanding nervous system function using five simple but powerful diagnostic tests.

Chapter 12 Introduction to the Nervous System Chapter 13 The Anatomy of the Nervous System Chapter 14 The Somatic Nervous System Chapter 15 The Autonomic Nervous System Chapter 16 The Neurological Exam Chapter 17 The Endocrine System

Unit 4: Fluids and Transport

In Chapters 18–21, students examine the principal means of transport for materials needed to support the human body, regulate its internal environment, and provide protection. Chapter 18 Blood Chapter 19 The Cardiovascular System: The Heart Chapter 20 The Cardiovascular System: Blood Vessels and Circulation Chapter 21 The Lymphatic System and Immunity

Unit 5: Energy, Maintenance, and Environmental Exchange

In Chapters 22–26, students discover the interaction between body systems and the outside environment for the exchange of materials, the capture of energy, the release of waste, and the overall maintenance of the internal systems that regulate the exchange. The explanations and illustrations are particularly focused on how structure relates to function.

Chapter 22 The Respiratory System Chapter 23 The Digestive System Chapter 24 Nutrition and Metabolism Chapter 25 The Urinary System Chapter 26 Fluid, Electrolyte, and Acid–Base Balance

Unit 6: Human Development and the Continuity of Life

The closing chapters examine the male and female reproductive systems, describe the process of human development and the different stages of pregnancy, and end with a review of the mechanisms of inheritance.

Chapter 27 The Reproductive System Chapter 28 Development and Genetic Inheritance

Pedagogical Foundation and Features

Anatomy and Physiology is designed to promote scientific literacy. Throughout the text, you will find features that engage the students by taking selected topics a step further.

  • Homeostatic Imbalances discusses the effects and results of imbalances in the body.
  • Disorders showcases a disorder that is relevant to the body system at hand. This feature may focus on a specific disorder, or a set of related disorders.
  • Diseases showcases a disease that is relevant to the body system at hand.
  • Aging explores the effect aging has on a body’s system and specific disorders that manifest over time.
  • Career Connections presents information on the various careers often pursued by allied health students, such as medical technician, medical examiner, and neurophysiologist. Students are introduced to the educational requirements for and day-to-day responsibilities in these careers.
  • Everyday Connections tie anatomical and physiological concepts to emerging issues and discuss these in terms of everyday life. Topics include “Anabolic Steroids” and “The Effect of Second-Hand Tobacco Smoke.”
  • Interactive Links direct students to online exercises, simulations, animations, and videos to add a fuller context to core content and help improve understanding of the material. Many features include links to the University of Michigan’s interactive WebScopes, which allow students to zoom in on micrographs in the collection. These resources were vetted by reviewers and other subject matter experts to ensure that they are effective and accurate. We strongly urge students to explore these links, whether viewing a video or inputting data into a simulation, to gain the fullest experience and to learn how to search for information independently.

Dynamic, Learner-Centered Art

Our unique approach to visuals is designed to emphasize only the components most important in any given illustration. The art style is particularly aimed at focusing student learning through a powerful blend of traditional depictions and instructional innovations.

Much of the art in this book consists of black line illustrations. The strongest line is used to highlight the most important structures, and shading is used to show dimension and shape. Color is used sparingly to highlight and clarify the primary anatomical or functional point of the illustration. This technique is intended to draw students’ attention to the critical learning point in the illustration, without distraction from excessive gradients, shadows, and highlights. Full color is used when the structure or process requires it (for example, muscle diagrams and cardiovascular system illustrations).

[caption id="" align="aligncenter" width="300"]A color illustration of the pharynx. The Pharynx. By highlighting the most important portions of the illustration, the artwork helps students focus on the most important points, without overwhelming them.[/caption]

Micrographs

Micrograph magnifications have been calculated based on the objective provided with the image. If a micrograph was recorded at 40×, and the image was magnified an additional 2×, we calculated the final magnification of the micrograph to be 80×.

Please note that, when viewing the textbook electronically, the micrograph magnification provided in the text does not take into account the size and magnification of the screen on your electronic device. There may be some variation

[caption id="" align="aligncenter" width="500"]A color illustration of the pharynx. Sebaceous Glands. These glands secrete oils that lubricate and protect the skin. LM × 400. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)[/caption]

Learning Resources

The following resources are (or will be) available in addition to main text:

  • PowerPoint slides: For each chapter, the illustrations are presented, one per slide, with their respective captions.
  • Pronunciation guide: A subset of the text’s key terms are presented with easy-to-follow phonetic transcriptions. For example, blastocyst is rendered as “blas'to-sist”

About Our Team

Senior Contributing Authors

J. Gordon Betts Tyler Junior College
Peter Desaix University of North Carolina at Chapel Hill
Eddie Johnson Central Oregon Community College
Jody E. Johnson Arapahoe Community College
Oksana Korol Aims Community College
Dean Kruse Portland Community College
Brandon Poe Springfield Technical Community College
James A. Wise Hampton University
Mark Womble Youngstown State University
Kelly A. Young California State University, Long Beach

Advisor

Robin J. Heyden

Contributing Authors

Kim Aaronson Aquarius Institute; Triton College
Lopamudra Agarwal Augusta Technical College
Gary Allen Dalhousie University
Robert Allison McLennan Community College
Heather Armbruster Southern Union State Community College
Timothy Ballard University of North Carolina Wilmington
Matthew Barlow Eastern New Mexico University
William Blaker Furman University
Julie Bowers East Tennessee State University
Emily Bradshaw Florida Southern College
Nishi Bryska University of North Carolina, Charlotte
Susan Caley Opsal Illinois Valley Community College
Boyd Campbell Southwest College of Naturopathic Medicine and Health Sciences
Ann Caplea Walsh University
Marnie Chapman University of Alaska, Sitka
Barbara Christie-Pope Cornell College
Kenneth Crane Texarkana College
Maurice Culver Florida State College at Jacksonville
Heather Cushman Tacoma Community College
Noelle Cutter Molloy College
Lynnette Danzl-Tauer Rock Valley College
Jane Davis Aurora University
AnnMarie DelliPizzi Dominican College
Susan Dentel Washtenaw Community College
Pamela Dobbins Shelton State Community College
Patty Dolan Pacific Lutheran University
Sondra Dubowsky McLennan Community College
Peter Dukehart Three Rivers Community College
Ellen DuPré Central College
Elizabeth DuPriest Warner Pacific College
Pam Elf University of Minnesota
Sharon Ellerton Queensborough Community College
Carla Endres Utah State University - College of Eastern Utah: San Juan Campus
Myriam Feldman Lake Washington Institute of Technology; Cascadia Community College
Greg Fitch Avila University
Lynn Gargan Tarant County College
Michael Giangrande Oakland Community College
Chaya Gopalan St. Louis College of Pharmacy
Victor Greco Chattahoochee Technical College
Susanna Heinze Skagit Valley College
Ann Henninger Wartburg College
Dale Horeth Tidewater Community College
Michael Hortsch University of Michigan
Rosemary Hubbard Marymount University
Mark Hubley Prince George's Community College
Branko Jablanovic College of Lake County
Norman Johnson University of Massachusetts Amherst
Mark Jonasson North Arkansas College
Jeff Keyte College of Saint Mary
William Kleinelp Middlesex County College
Leigh Kleinert Grand Rapids Community College
Brenda Leady University of Toledo
John Lepri University of North Carolina, Greensboro
Sarah Leupen University of Maryland, Baltimore County
Lihua Liang Johns Hopkins University
Robert Mallet University of North Texas Health Science Center
Bruce Maring Daytona State College
Elisabeth Martin College of Lake County
Natalie Maxwell Carl Albert State College, Sallisaw
Julie May William Carey University
Debra McLaughlin University of Maryland University College
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Shobhana Natarajan Brookhaven College
Phillip Nicotera St. Petersburg College
Mary Jane Niles University of San Francisco
Ikemefuna Nwosu Parkland College; Lake Land College
Betsy Ott Tyler Junior College
Ivan Paul John Wood Community College
Aaron Payette College of Southern Nevada
Scott Payne Kentucky Wesleyan College
Cameron Perkins South Georgia College
David Pfeiffer University of Alaska, Anchorage
Thomas Pilat Illinois Central College
Eileen Preston Tarrant County College
Mike Pyle Olivet Nazarene University
Robert Rawding Gannon University
Jason Schreer State University of New York at Potsdam
Laird Sheldahl Mt. Hood Community College
Brian Shmaefsky Lone Star College System
Douglas Sizemore Bevill State Community College
Susan Spencer Mount Hood Community College
Cynthia Standley University of Arizona
Robert Sullivan Marist College
Eric Sun Middle Georgia State College
Tom Swenson Ithaca College
Kathleen Tallman Azusa Pacific University
Rohinton Tarapore University of Pennsylvania
Elizabeth Tattersall Western Nevada College
Mark Thomas University of Northern Colorado
Janis Thompson Lorain County Community College
Rita Thrasher Pensacola State College
David Van Wylen St. Olaf College
Lynn Wandrey Mott Community College
Margaret Weck St. Louis College of Pharmacy
Kathleen Weiss George Fox University
Neil Westergaard Williston State College
David Wortham West Georgia Technical College
Umesh Yadav University of Texas Medical Branch
Tony Yates Oklahoma Baptist University
Justin York Glendale Community College
Cheri Zao North Idaho College
Elena Zoubina Bridgewater State University; Massasoit Community College
Shobhana Natarajan Alcon Laboratories, Inc.
Jennifer Barker Douglas College
Sarah McKinnon Douglas College

Special Thanks

OpenStax wishes to thank the Regents of University of Michigan Medical School for the use of their extensive micrograph collection. Many of the UM micrographs that appear in Anatomy and Physiology are interactive WebScopes, which students can explore by zooming in and out.

We also wish to thank the Open Learning Initiative at Carnegie Mellon University, with whom we shared and exchanged resources during the development of Anatomy and Physiology.

]]> 1833 0 2 0 1103 Chapter 1. An Introduction to the Human Body https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-1-an-introduction-to-the-human-body-2/ Thu, 13 Jul 2017 22:31:36 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-1-an-introduction-to-the-human-body-2/ 1834 0 1 0 1103 Chapter 2. The Chemical Level of Organization https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-2-the-chemical-level-of-organization-2/ Thu, 13 Jul 2017 22:32:49 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-2-the-chemical-level-of-organization-2/ 1901 0 2 0 1103 Chapter 3. The Cellular Level of Organization https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-3-the-cellular-level-of-organization-2/ Thu, 13 Jul 2017 22:33:37 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-3-the-cellular-level-of-organization-2/ 1936 0 3 0 1103 Chapter 4. The Tissue Level of Organization https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-4-the-tissue-level-of-organization-2/ Thu, 13 Jul 2017 22:34:09 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-4-the-tissue-level-of-organization-2/ 1977 0 4 0 8.5 Development of the Appendicular Skeleton https://pressbooks.bccampus.ca/apdouglas2017/chapter/8-5-development-of-the-appendicular-skeleton/ Wed, 02 Aug 2017 23:10:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2107

Learning Objectives

By the end of this section, you will be able to:
  • Describe the growth and development of the embryonic limb buds
  • Discuss the appearance of primary and secondary ossification centers

Embryologically, the appendicular skeleton arises from mesenchyme, a type of embryonic tissue that can differentiate into many types of tissues, including bone or muscle tissue. Mesenchyme gives rise to the bones of the upper and lower limbs, as well as to the pectoral and pelvic girdles. Development of the limbs begins near the end of the fourth embryonic week, with the upper limbs appearing first. Thereafter, the development of the upper and lower limbs follows similar patterns, with the lower limbs lagging behind the upper limbs by a few days.

Limb Growth

Each upper and lower limb initially develops as a small bulge called a limb bud, which appears on the lateral side of the early embryo. The upper limb bud appears near the end of the fourth week of development, with the lower limb bud appearing shortly after (Figure 1).
[caption id="" align="aligncenter" width="280"]This photograph shows a seven-week embryo which derived from an ectopic pregnancy. Figure 1. Embryo at Seven Weeks. Limb buds are visible in an embryo at the end of the seventh week of development (embryo derived from an ectopic pregnancy). (credit: Ed Uthman/flickr)[/caption]
Initially, the limb buds consist of a core of mesenchyme covered by a layer of ectoderm. The ectoderm at the end of the limb bud thickens to form a narrow crest called the apical ectodermal ridge. This ridge stimulates the underlying mesenchyme to rapidly proliferate, producing the outgrowth of the developing limb. As the limb bud elongates, cells located farther from the apical ectodermal ridge slow their rates of cell division and begin to differentiate. In this way, the limb develops along a proximal-to-distal axis.

During the sixth week of development, the distal ends of the upper and lower limb buds expand and flatten into a paddle shape. This region will become the hand or foot. The wrist or ankle areas then appear as a constriction that develops at the base of the paddle. Shortly after this, a second constriction on the limb bud appears at the future site of the elbow or knee. Within the paddle, areas of tissue undergo cell death, producing separations between the growing fingers and toes. Also during the sixth week of development, mesenchyme within the limb buds begins to differentiate into hyaline cartilage that will form models of the future limb bones.

The early outgrowth of the upper and lower limb buds initially has the limbs positioned so that the regions that will become the palm of the hand or the bottom of the foot are facing medially toward the body, with the future thumb or big toe both oriented toward the head. During the seventh week of development, the upper limb rotates laterally by 90 degrees, so that the palm of the hand faces anteriorly and the thumb points laterally. In contrast, the lower limb undergoes a 90-degree medial rotation, thus bringing the big toe to the medial side of the foot.

Ossification of Appendicular Bones

All of the girdle and limb bones, except for the clavicle, develop by the process of endochondral ossification. This process begins as the mesenchyme within the limb bud differentiates into hyaline cartilage to form cartilage models for future bones. By the twelfth week, a primary ossification center will have appeared in the diaphysis (shaft) region of the long bones, initiating the process that converts the cartilage model into bone. A secondary ossification center will appear in each epiphysis (expanded end) of these bones at a later time, usually after birth. The primary and secondary ossification centers are separated by the epiphyseal plate, a layer of growing hyaline cartilage. This plate is located between the diaphysis and each epiphysis. It continues to grow and is responsible for the lengthening of the bone. The epiphyseal plate is retained for many years, until the bone reaches its final, adult size, at which time the epiphyseal plate disappears and the epiphysis fuses to the diaphysis. (Seek additional content on ossification in the chapter on bone tissue).

Small bones, such as the phalanges, will develop only one secondary ossification center and will thus have only a single epiphyseal plate. Large bones, such as the femur, will develop several secondary ossification centers, with an epiphyseal plate associated with each secondary center. Thus, ossification of the femur begins at the end of the seventh week with the appearance of the primary ossification center in the diaphysis, which rapidly expands to ossify the shaft of the bone prior to birth. Secondary ossification centers develop at later times. Ossification of the distal end of the femur, to form the condyles and epicondyles, begins shortly before birth. Secondary ossification centers also appear in the femoral head late in the first year after birth, in the greater trochanter during the fourth year, and in the lesser trochanter between the ages of 9 and 10 years. Once these areas have ossified, their fusion to the diaphysis and the disappearance of each epiphyseal plate follow a reversed sequence. Thus, the lesser trochanter is the first to fuse, doing so at the onset of puberty (around 11 years of age), followed by the greater trochanter approximately 1 year later. The femoral head fuses between the ages of 14–17 years, whereas the distal condyles of the femur are the last to fuse, between the ages of 16–19 years. Knowledge of the age at which different epiphyseal plates disappear is important when interpreting radiographs taken of children. Since the cartilage of an epiphyseal plate is less dense than bone, the plate will appear dark in a radiograph image. Thus, a normal epiphyseal plate may be mistaken for a bone fracture.

The clavicle is the one appendicular skeleton bone that does not develop via endochondral ossification. Instead, the clavicle develops through the process of intramembranous ossification. During this process, mesenchymal cells differentiate directly into bone-producing cells, which produce the clavicle directly, without first making a cartilage model. Because of this early production of bone, the clavicle is the first bone of the body to begin ossification, with ossification centers appearing during the fifth week of development. However, ossification of the clavicle is not complete until age 25.

]]> 2107 2082 6 0 2.4 Inorganic Compounds Essential to Human Functioning https://pressbooks.bccampus.ca/apdouglas2017/chapter/2-4-inorganic-compounds-essential-to-human-functioning-2/ Wed, 02 Aug 2017 22:11:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2447

Learning Objectives

By the end of this section, you will be able to:
  • Compare and contrast inorganic and organic compounds
  • Identify the properties of water that make it essential to life
  • Explain the role of salts in body functioning
  • Distinguish between acids and bases, and explain their role in pH
  • Discuss the role of buffers in helping the body maintain pH homeostasis
The concepts you have learned so far in this chapter govern all forms of matter, and would work as a foundation for geology as well as biology. This section of the chapter narrows the focus to the chemistry of human life; that is, the compounds important for the body’s structure and function. In general, these compounds are either inorganic or organic.
  • An inorganic compound is a substance that does not contain both carbon and hydrogen. A great many inorganic compounds do contain hydrogen atoms, such as water (H2O) and the hydrochloric acid (HCl) produced by your stomach. In contrast, only a handful of inorganic compounds contain carbon atoms. Carbon dioxide (CO2) is one of the few examples.
  • An organic compound, then, is a substance that contains both carbon and hydrogen. Organic compounds are synthesized via covalent bonds within living organisms, including the human body. Recall that carbon and hydrogen are the second and third most abundant elements in your body. You will soon discover how these two elements combine in the foods you eat, in the compounds that make up your body structure, and in the chemicals that fuel your functioning.

The following section examines the three groups of inorganic compounds essential to life: water, salts, acids, and bases. Organic compounds are covered later in the chapter.

Water

As much as 70 percent of an adult’s body weight is water. This water is contained both within the cells and between the cells that make up tissues and organs. Its several roles make water indispensable to human functioning.

Water as a Lubricant and Cushion

Water is a major component of many of the body’s lubricating fluids. Just as oil lubricates the hinge on a door, water in synovial fluid lubricates the actions of body joints, and water in pleural fluid helps the lungs expand and recoil with breathing. Watery fluids help keep food flowing through the digestive tract, and ensure that the movement of adjacent abdominal organs is friction free.

Water also protects cells and organs from physical trauma, cushioning the brain within the skull, for example, and protecting the delicate nerve tissue of the eyes. Water cushions a developing fetus in the mother’s womb as well.

Water as a Heat Sink

A heat sink is a substance or object that absorbs and dissipates heat but does not experience a corresponding increase in temperature. In the body, water absorbs the heat generated by chemical reactions without greatly increasing in temperature. Moreover, when the environmental temperature soars, the water stored in the body helps keep the body cool. This cooling effect happens as warm blood from the body’s core flows to the blood vessels just under the skin and is transferred to the environment. At the same time, sweat glands release warm water in sweat. As the water evaporates into the air, it carries away heat, and then the cooler blood from the periphery circulates back to the body core.

Water as a Component of Liquid Mixtures

A mixture is a combination of two or more substances, each of which maintains its own chemical identity. In other words, the constituent substances are not chemically bonded into a new, larger chemical compound. The concept is easy to imagine if you think of powdery substances such as flour and sugar; when you stir them together in a bowl, they obviously do not bond to form a new compound. The room air you breathe is a gaseous mixture, containing three discrete elements—nitrogen, oxygen, and argon—and one compound, carbon dioxide. There are three types of liquid mixtures, all of which contain water as a key component. These are solutions, colloids, and suspensions.

For cells in the body to survive, they must be kept moist in a water-based liquid called a solution. In chemistry, a liquid solution consists of a solvent that dissolves a substance called a solute. An important characteristic of solutions is that they are homogeneous; that is, the solute molecules are distributed evenly throughout the solution. If you were to stir a teaspoon of sugar into a glass of water, the sugar would dissolve into sugar molecules separated by water molecules. The ratio of sugar to water in the left side of the glass would be the same as the ratio of sugar to water in the right side of the glass. If you were to add more sugar, the ratio of sugar to water would change, but the distribution—provided you had stirred well—would still be even.

Water is considered the “universal solvent” and it is believed that life cannot exist without water because of this. Water is certainly the most abundant solvent in the body; essentially all of the body’s chemical reactions occur among compounds dissolved in water. Because water molecules are polar, with regions of positive and negative electrical charge, water readily dissolves ionic compounds and polar covalent compounds. Such compounds are referred to as hydrophilic, or “water-loving.” As mentioned above, sugar dissolves well in water. This is because sugar molecules contain regions of hydrogen-oxygen polar bonds, making it hydrophilic. Nonpolar molecules, which do not readily dissolve in water, are called hydrophobic, or “water-fearing.”

Concentrations of Solutes

Various mixtures of solutes and water are described in chemistry. The concentration of a given solute is the number of particles of that solute in a given space (oxygen makes up about 21 percent of atmospheric air). In the bloodstream of humans, glucose concentration is usually measured in milligram (mg) per deciliter (dL), and in a healthy adult averages about 100 mg/dL. Another method of measuring the concentration of a solute is by its molarilty—which is moles (M) of the molecules per liter (L). The mole of an element is its atomic weight, while a mole of a compound is the sum of the atomic weights of its components, called the molecular weight. An often-used example is calculating a mole of glucose, with the chemical formula C6H12O6. Using the periodic table, the atomic weight of carbon (C) is 12.011 grams (g), and there are six carbons in glucose, for a total atomic weight of 72.066 g. Doing the same calculations for hydrogen (H) and oxygen (O), the molecular weight equals 180.156g (the “gram molecular weight” of glucose). When water is added to make one liter of solution, you have one mole (1M) of glucose. This is particularly useful in chemistry because of the relationship of moles to “Avogadro’s number.” A mole of any solution has the same number of particles in it: 6.02 × 1023. Many substances in the bloodstream and other tissue of the body are measured in thousandths of a mole, or millimoles (mM).

A colloid is a mixture that is somewhat like a heavy solution. The solute particles consist of tiny clumps of molecules large enough to make the liquid mixture opaque (because the particles are large enough to scatter light). Familiar examples of colloids are milk and cream. In the thyroid glands, the thyroid hormone is stored as a thick protein mixture also called a colloid.

A suspension is a liquid mixture in which a heavier substance is suspended temporarily in a liquid, but over time, settles out. This separation of particles from a suspension is called sedimentation. An example of sedimentation occurs in the blood test that establishes sedimentation rate, or sed rate. The test measures how quickly red blood cells in a test tube settle out of the watery portion of blood (known as plasma) over a set period of time. Rapid sedimentation of blood cells does not normally happen in the healthy body, but aspects of certain diseases can cause blood cells to clump together, and these heavy clumps of blood cells settle to the bottom of the test tube more quickly than do normal blood cells.

The Role of Water in Chemical Reactions

Two types of chemical reactions involve the creation or the consumption of water: dehydration synthesis and hydrolysis.

  • In dehydration synthesis, one reactant gives up an atom of hydrogen and another reactant gives up a hydroxyl group (OH) in the synthesis of a new product. In the formation of their covalent bond, a molecule of water is released as a byproduct (Figure 1). This is also sometimes referred to as a condensation reaction.
  • In hydrolysis, a molecule of water disrupts a compound, breaking its bonds. The water is itself split into H and OH. One portion of the severed compound then bonds with the hydrogen atom, and the other portion bonds with the hydroxyl group.
These reactions are reversible, and play an important role in the chemistry of organic compounds (which will be discussed shortly).
[caption id="" align="aligncenter" width="520"]The top panel in this figure shows a dehydration-synthesis reaction, and the bottom panel shows a hydrolysis reaction. Figure 1. Dehydration Synthesis and Hydrolysis. Monomers, the basic units for building larger molecules, form polymers (two or more chemically-bonded monomers). (a) In dehydration synthesis, two monomers are covalently bonded in a reaction in which one gives up a hydroxyl group and the other a hydrogen atom. A molecule of water is released as a byproduct during dehydration reactions. (b) In hydrolysis, the covalent bond between two monomers is split by the addition of a hydrogen atom to one and a hydroxyl group to the other, which requires the contribution of one molecule of water.[/caption]
[caption id="attachment_3037" align="aligncenter" width="150"] Watch this amoeba sisters video to learn more about the properties of water![/caption] [caption id="attachment_2951" align="aligncenter" width="150"] Watch this CrashCourse video to learn more about the importance of water![/caption]

Salts

Recall that salts are formed when ions form ionic bonds. In these reactions, one atom gives up one or more electrons, and thus becomes positively charged, whereas the other accepts one or more electrons and becomes negatively charged. You can now define a salt as a substance that, when dissolved in water, dissociates into ions other than H+ or OH–. This fact is important in distinguishing salts from acids and bases, discussed next.

A typical salt, NaCl, dissociates completely in water (Figure 2). The positive and negative regions on the water molecule (the hydrogen and oxygen ends respectively) attract the negative chloride and positive sodium ions, pulling them away from each other. Again, whereas nonpolar and polar covalently bonded compounds break apart into molecules in solution, salts dissociate into ions. These ions are electrolytes; they are capable of conducting an electrical current in solution. This property is critical to the function of ions in transmitting nerve impulses and prompting muscle contraction.

[caption id="" align="aligncenter" width="480"]This figure shows a crystal lattice of sodium chloride interacting with water to form a hydrated sodium ion and a hydrated chloride ion. Figure 2. Dissociation of Sodium Chloride in Water. Notice that the crystals of sodium chloride dissociate not into molecules of NaCl, but into Na+ cations and Cl– anions, each completely surrounded by water molecules.[/caption]

Many other salts are important in the body. For example, bile salts produced by the liver help break apart dietary fats, and calcium phosphate salts form the mineral portion of teeth and bones.

Acids and Bases

Acids and bases, like salts, dissociate in water into electrolytes. Acids and bases can very much change the properties of the solutions in which they are dissolved.

Acids

An acid is a substance that releases hydrogen ions (H+) in solution (Figure 3a). Because an atom of hydrogen has just one proton and one electron, a positively charged hydrogen ion is simply a proton. This solitary proton is highly likely to participate in chemical reactions. Strong acids are compounds that release all of their H+ in solution; that is, they ionize completely. Hydrochloric acid (HCl), which is released from cells in the lining of the stomach, is a strong acid because it releases all of its H+ in the stomach’s watery environment. This strong acid aids in digestion and kills ingested microbes. Weak acids do not ionize completely; that is, some of their hydrogenionsremain bonded within a compound in solution. An example of a weak acid is vinegar, or acetic acid; it is called acetate after it gives up a proton.
Acids and Bases
(a) In aqueous solution, an acid dissociates into hydrogen ions (H+) and anions. Nearly every molecule of a strong acid dissociates, producing a high concentration of H+. (b) In aqueous solution, a base dissociates into hydroxyl ions (OH–) and cations. Nearly every molecule of a strong base dissociates, producing a high concentration of OH–.
[caption id="" align="aligncenter" width="420"]This figure shows four beakers containing different liquids. Figure 3. Acids and Bases. (a) In aqueous solution, an acid dissociates into hydrogen ions (H+) and anions. Nearly every molecule of a strong acid dissociates, producing a high concentration of H+. (b) In aqueous solution, a base dissociates into hydroxyl ions (OH–) and cations. Nearly every molecule of a strong base dissociates, producing a high concentration of OH–.[/caption]

Bases

A base is a substance that releases hydroxyl ions (OH–) in solution, or one that accepts H+ already present in solution (see Figure 3b). The hydroxyl ions (also known as hydroxide ions) or other basic substances combine with H+ present to form a water molecule, thereby removing H+ and reducing the solution’s acidity. Strong bases release most or all of their hydroxyl ions; weak bases release only some hydroxyl ions or absorb only a few H+. Food mixed with hydrochloric acid from the stomach would burn the small intestine, the next portion of the digestive tract after the stomach, if it were not for the release of bicarbonate (HCO3–), a weak base that attracts H+. Bicarbonate accepts some of the H+ protons, thereby reducing the acidity of the solution.

The Concept of pH

The relative acidity or alkalinity of a solution can be indicated by its pH. A solution’s pH is the negative, base-10 logarithm of the hydrogen ion (H+) concentration of the solution. As an example, a pH 4 solution has an H+ concentration that is ten times greater than that of a pH 5 solution. That is, a solution with a pH of 4 is ten times more acidic than a solution with a pH of 5. The concept of pH will begin to make more sense when you study the pH scale, like that shown in [link]. The scale consists of a series of increments ranging from 0 to 14. A solution with a pH of 7 is considered neutral—neither acidic nor basic. Pure water has a pH of 7. The lower the number below 7, the more acidic the solution, or the greater the concentration of H+. The concentration of hydrogen ions at each pH value is 10 times different than the next pH. For instance, a pH value of 4 corresponds to a proton concentration of 10–4 M, or 0.0001M, while a pH value of 5 corresponds to a proton concentration of 10–5 M, or 0.00001M. The higher the number above 7, the more basic (alkaline) the solution, or the lower the concentration of H+. Human urine, for example, is ten times more acidic than pure water, and HCl is 10,000,000 times more acidic than water.
[caption id="" align="aligncenter" width="320"]This figure shows a vertical arrow with the top half showing the basic scale and the bottom half showing the acidic scale. Different chemicals and their pH are also shown. Figure 4. The pH Scale[/caption]

Buffers

The pH of human blood normally ranges from 7.35 to 7.45, although it is typically identified as pH 7.4. At this slightly basic pH, blood can reduce the acidity resulting from the carbon dioxide (CO2) constantly being released into the bloodstream by the trillions of cells in the body. Homeostatic mechanisms (along with exhaling CO2 while breathing) normally keep the pH of blood within this narrow range. This is critical, because fluctuations—either too acidic or too alkaline—can lead to life-threatening disorders.

All cells of the body depend on homeostatic regulation of acid–base balance at a pH of approximately 7.4. The body therefore has several mechanisms for this regulation, involving breathing, the excretion of chemicals in urine, and the internal release of chemicals collectively called buffers into body fluids. A buffer is a solution of a weak acid and its conjugate base. A buffer can neutralize small amounts of acids or bases in body fluids. For example, if there is even a slight decrease below 7.35 in the pH of a bodily fluid, the buffer in the fluid—in this case, acting as a weak base—will bind the excess hydrogen ions. In contrast, if pH rises above 7.45, the buffer will act as a weak acid and contribute hydrogen ions.

Homeostatic Imbalances

Acids and Bases Excessive acidity of the blood and other body fluids is known as acidosis. Common causes of acidosis are situations and disorders that reduce the effectiveness of breathing, especially the person’s ability to exhale fully, which causes a buildup of CO2 (and H+) in the bloodstream. Acidosis can also be caused by metabolic problems that reduce the level or function of buffers that act as bases, or that promote the production of acids. For instance, with severe diarrhea, too much bicarbonate can be lost from the body, allowing acids to build up in body fluids. In people with poorly managed diabetes (ineffective regulation of blood sugar), acids called ketones are produced as a form of body fuel. These can build up in the blood, causing a serious condition called diabetic ketoacidosis. Kidney failure, liver failure, heart failure, cancer, and other disorders also can prompt metabolic acidosis.

In contrast, alkalosis is a condition in which the blood and other body fluids are too alkaline (basic). As with acidosis, respiratory disorders are a major cause; however, in respiratory alkalosis, carbon dioxide levels fall too low. Lung disease, aspirin overdose, shock, and ordinary anxiety can cause respiratory alkalosis, which reduces the normal concentration of H+.

Metabolic alkalosis often results from prolonged, severe vomiting, which causes a loss of hydrogen and chloride ions (as components of HCl). Medications also can prompt alkalosis. These include diuretics that cause the body to lose potassium ions, as well as antacids when taken in excessive amounts, for instance by someone with persistent heartburn or an ulcer.

]]> 2447 2429 3 0 2.5 Organic Compounds Essential to Human Functioning https://pressbooks.bccampus.ca/apdouglas2017/chapter/2-5-organic-compounds-essential-to-human-functioning-2/ Wed, 02 Aug 2017 22:48:28 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2461

Learning Objectives

By the end of this section, you will be able to:
  • Identify four types of organic molecules essential to human functioning
  • Explain the chemistry behind carbon’s affinity for covalently bonding in organic compounds
  • Provide examples of three types of carbohydrates, and identify the primary functions of carbohydrates in the body
  • Discuss four types of lipids important in human functioning
  • Describe the structure of proteins, and discuss their importance to human functioning
  • Identify the building blocks of nucleic acids, and the roles of DNA, RNA, and ATP in human functioning

Organic compounds typically consist of groups of carbon atoms covalently bonded to hydrogen, usually oxygen, and often other elements as well. Created by living things, they are found throughout the world, in soils and seas, commercial products, and every cell of the human body. The four types most important to human structure and function are carbohydrates, lipids, proteins, and nucleotides. Before exploring these compounds, you need to first understand the chemistry of carbon.

The Chemistry of Carbon

What makes organic compounds ubiquitous is the chemistry of their carbon core. Recall that carbon atoms have four electrons in their valence shell, and that the octet rule dictates that atoms tend to react in such a way as to complete their valence shell with eight electrons. Carbon atoms do not complete their valence shells by donating or accepting four electrons. Instead, they readily share electrons via covalent bonds.

Commonly, carbon atoms share with other carbon atoms, often forming a long carbon chain referred to as a carbon skeleton. When they do share, however, they do not share all their electrons exclusively with each other. Rather, carbon atoms tend to share electrons with a variety of other elements, one of which is always hydrogen. Carbon and hydrogen groupings are called hydrocarbons. If you study the figures of organic compounds in the remainder of this chapter, you will see several with chains of hydrocarbons in one region of the compound.

Many combinations are possible to fill carbon’s four “vacancies.” Carbon may share electrons with oxygen or nitrogen or other atoms in a particular region of an organic compound. Moreover, the atoms to which carbon atoms bond may also be part of a functional group. A functional group is a group of atoms linked by strong covalent bonds and tending to function in chemical reactions as a single unit. You can think of functional groups as tightly knit “cliques” whose members are unlikely to be parted. Five functional groups are important in human physiology; these are the hydroxyl, carboxyl, amino, methyl and phosphate groups (Table 1).
Functional Groups Important in Human Physiology
Functional group Structural formula Importance
Hydroxyl —O—H Hydroxyl groups are polar. They are components of all four types of organic compounds discussed in this chapter. They are involved in dehydration synthesis and hydrolysis reactions.
Carboxyl O—C—OH Carboxyl groups are found within fatty acids, amino acids, and many other acids.
Amino —N—H2 Amino groups are found within amino acids, the building blocks of proteins.
Methyl —C—H3 Methyl groups are found within amino acids.
Phosphate —P—O42– Phosphate groups are found within phospholipids and nucleotides.

Carbon’s affinity for covalent bonding means that many distinct and relatively stable organic molecules nevertheless readily form larger, more complex molecules. Any large molecule is referred to as macromolecule (macro- = “large”), and the organic compounds in this section all fit this description. However, some macromolecules are made up of several “copies” of single units called monomer (mono- = “one”; -mer = “part”). Like beads in a long necklace, these monomers link by covalent bonds to form long polymers (poly- = “many”). There are many examples of monomers and polymers among the organic compounds.

Monomers form polymers by engaging in dehydration synthesis (see Chapter 2.4 Figure 1). As was noted earlier, this reaction results in the release of a molecule of water. Each monomer contributes: One gives up a hydrogen atom and the other gives up a hydroxyl group. Polymers are split into monomers by hydrolysis (-lysis = “rupture”). The bonds between their monomers are broken, via the donation of a molecule of water, which contributes a hydrogen atom to one monomer and a hydroxyl group to the other.

Carbohydrates

The term carbohydrate means “hydrated carbon.” Recall that the root hydro- indicates water. A carbohydrate is a molecule composed of carbon, hydrogen, and oxygen; in most carbohydrates, hydrogen and oxygen are found in the same two-to-one relative proportions they have in water. In fact, the chemical formula for a “generic” molecule of carbohydrate is (CH2O)n.

Carbohydrates are referred to as saccharides, a word meaning “sugars.” Three forms are important in the body. Monosaccharides are the monomers of carbohydrates. Disaccharides (di- = “two”) are made up of two monomers. Polysaccharides are the polymers, and can consist of hundreds to thousands of monomers.

Monosaccharides

A monosaccharide is a monomer of carbohydrates. Five monosaccharides are important in the body. Three of these are the hexose sugars, so called because they each contain six atoms of carbon. These are glucose, fructose, and galactose, shown in Figure 1a. The remaining monosaccharides are the two pentose sugars, each of which contains five atoms of carbon. They are ribose and deoxyribose, shown in Figure 2b.

[caption id="" align="aligncenter" width="420"]This figure shows the structure of glucose, fructose, galactose, deoxyribose, and ribose. Figure 1. Five Important Monosaccharides.[/caption]

Disaccharides

A disaccharide is a pair of monosaccharides. Disaccharides are formed via dehydration synthesis, and the bond linking them is referred to as a glycosidic bond (glyco- = “sugar”). Three disaccharides (shown in Figure 2) are important to humans. These are sucrose, commonly referred to as table sugar; lactose, or milk sugar; and maltose, or malt sugar. As you can tell from their common names, you consume these in your diet; however, your body cannot use them directly. Instead, in the digestive tract, they are split into their component monosaccharides via hydrolysis.

[caption id="" align="aligncenter" width="420"]This figure shows the structure of sucrose, lactose, and maltose. Figure 2. Three Important Disaccharides. All three important disaccharides form by dehydration synthesis.[/caption]

Polysaccharides

Polysaccharides can contain a few to a thousand or more monosaccharides. Three are important to the body (Figure 3):

  • Starches are polymers of glucose. They occur in long chains called amylose or branched chains called amylopectin, both of which are stored in plant-based foods and are relatively easy to digest.
  • Glycogen is also a polymer of glucose, but it is stored in the tissues of animals, especially in the muscles and liver. It is not considered a dietary carbohydrate because very little glycogen remains in animal tissues after slaughter; however, the human body stores excess glucose as glycogen, again, in the muscles and liver.
  • Cellulose, a polysaccharide that is the primary component of the cell wall of green plants, is the component of plant food referred to as “fiber”. In humans, cellulose/fiber is not digestible; however, dietary fiber has many health benefits. It helps you feel full so you eat less, it promotes a healthy digestive tract, and a diet high in fiber is thought to reduce the risk of heart disease and possibly some forms of cancer.
[caption id="" align="aligncenter" width="420"]This figure shows the structure of starch, glycogen, and cellulose. Figure 3. Three Important Polysaccharides. Three important polysaccharides are starches, glycogen, and fiber.[/caption]

Functions of Carbohydrates

The body obtains carbohydrates from plant-based foods. Grains, fruits, and legumes and other vegetables provide most of the carbohydrate in the human diet, although lactose is found in dairy products.

Although most body cells can break down other organic compounds for fuel, all body cells can use glucose. Moreover, nerve cells (neurons) in the brain, spinal cord, and through the peripheral nervous system, as well as red blood cells, can use only glucose for fuel. In the breakdown of glucose for energy, molecules of adenosine triphosphate, better known as ATP, are produced. Adenosine triphosphate (ATP) is composed of a ribose sugar, an adenine base, and three phosphate groups. ATP releases free energy when its phosphate bonds are broken, and thus supplies ready energy to the cell. More ATP is produced in the presence of oxygen (O2) than in pathways that do not use oxygen. The overall reaction for the conversion of the energy in glucose to energy stored in ATP can be written:

C6H12O6 + 6 O2 → 6 CO2 + 6 H2O + ATP

In addition to being a critical fuel source, carbohydrates are present in very small amounts in cells’ structure. For instance, some carbohydrate molecules bind with proteins to produce glycoproteins, and others combine with lipids to produce glycolipids, both of which are found in the membrane that encloses the contents of body cells.

Lipids

A lipid is one of a highly diverse group of compounds made up mostly of hydrocarbons. The few oxygen atoms they contain are often at the periphery of the molecule. Their nonpolar hydrocarbons make all lipids hydrophobic. In water, lipids do not form a true solution, but they may form an emulsion, which is the term for a mixture of solutions that do not mix well.

Triglycerides

A triglyceride is one of the most common dietary lipid groups, and the type found most abundantly in body tissues. This compound, which is commonly referred to as a fat, is formed from the synthesis of two types of molecules (Figure 4):

  • A glycerol backbone at the core of triglycerides, consists of three carbon atoms.
  • Three fatty acids, long chains of hydrocarbons with a carboxyl group and a methyl group at opposite ends, extend from each of the carbons of the glycerol.
[caption id="" align="aligncenter" width="550"]This image shows the reaction for the formation of triglycerides. Figure 4. Triglycerides. Triglycerides are composed of glycerol attached to three fatty acids via dehydration synthesis. Notice that glycerol gives up a hydrogen atom, and the carboxyl groups on the fatty acids each give up a hydroxyl group.[/caption]

Triglycerides form via dehydration synthesis. Glycerol gives up hydrogen atoms from its hydroxyl groups at each bond, and the carboxyl group on each fatty acid chain gives up a hydroxyl group. A total of three water molecules are thereby released.

Fatty acid chains that have no double carbon bonds anywhere along their length and therefore contain the maximum number of hydrogen atoms are called saturated fatty acids. These straight, rigid chains pack tightly together and are solid or semi-solid at room temperature (Figure 5a). Butter and lard are examples, as is the fat found on a steak or in your own body. In contrast, fatty acids with one double carbon bond are kinked at that bond (Figure 5b). These monounsaturated fatty acids are therefore unable to pack together tightly, and are liquid at room temperature. Polyunsaturated fatty acids contain two or more double carbon bonds, and are also liquid at room temperature. Plant oils such as olive oil typically contain both mono- and polyunsaturated fatty acids.

[caption id="" align="aligncenter" width="380"]This diagram shows the chain structures of a saturated and an unsaturated fatty acid. Figure 5. Fatty Acid Shapes. The level of saturation of a fatty acid affects its shape. (a) Saturated fatty acid chains are straight. (b) Unsaturated fatty acid chains are kinked.[/caption]

Whereas a diet high in saturated fatty acids increases the risk of heart disease, a diet high in unsaturated fatty acids is thought to reduce the risk. This is especially true for the omega-3 unsaturated fatty acids found in cold-water fish such as salmon. These fatty acids have their first double carbon bond at the third hydrocarbon from the methyl group (referred to as the omega end of the molecule).

Finally, trans fatty acids found in some processed foods, including some stick and tub margarines, are thought to be even more harmful to the heart and blood vessels than saturated fatty acids. Trans fats are created from unsaturated fatty acids (such as corn oil) when chemically treated to produce partially hydrogenated fats.

As a group, triglycerides are a major fuel source for the body. When you are resting or asleep, a majority of the energy used to keep you alive is derived from triglycerides stored in your fat (adipose) tissues. Triglycerides also fuel long, slow physical activity such as gardening or hiking, and contribute a modest percentage of energy for vigorous physical activity. Dietary fat also assists the absorption and transport of the nonpolar fat-soluble vitamins A, D, E, and K. Additionally, stored body fat protects and cushions the body’s bones and internal organs, and acts as insulation to retain body heat.

Fatty acids are also components of glycolipids, which are sugar-fat compounds found in the cell membrane. Lipoproteins are compounds in which the hydrophobic triglycerides are packaged in protein envelopes for transport in body fluids.

Phospholipids

As its name suggests, a phospholipid is a bond between the glycerol component of a lipid and a phosphorous molecule. In fact, phospholipids are similar in structure to triglycerides. However, instead of having three fatty acids, a phospholipid is generated from a diglyceride, a glycerol with just two fatty acid chains (Figure 6). The third binding site on the glycerol is taken up by the phosphate group, which in turn is attached to a polar “head” region of the molecule. Recall that triglycerides are nonpolar and hydrophobic. This still holds for the fatty acid portion of a phospholipid compound. However, the head of a phospholipid contains charges on the phosphate groups, as well as on the nitrogen atom. These charges make the phospholipid head hydrophilic. Therefore, phospholipids are said to have hydrophobic tails, containing the neutral fatty acids, and hydrophilic heads, containing the charged phosphate groups and nitrogen atom.

[caption id="" align="aligncenter" width="600"]This figure shows the chemical structure of different lipids. Figure 6. Other Important Lipids. (a) Phospholipids are composed of two fatty acids, glycerol, and a phosphate group. (b) Sterols are ring-shaped lipids. Shown here is cholesterol. (c) Prostaglandins are derived from unsaturated fatty acids. Prostaglandin E2 (PGE2) includes hydroxyl and carboxyl groups.[/caption]

Steroids

A steroid compound (referred to as a sterol) has as its foundation a set of four hydrocarbon rings bonded to a variety of other atoms and molecules (see Figure 6b). Although both plants and animals synthesize sterols, the type that makes the most important contribution to human structure and function is cholesterol, which is synthesized by the liver in humans and animals and is also present in most animal-based foods. Like other lipids, cholesterol’s hydrocarbons make it hydrophobic; however, it has a polar hydroxyl head that is hydrophilic. Cholesterol is an important component of bile acids, compounds that help emulsify dietary fats. In fact, the word root chole- refers to bile. Cholesterol is also a building block of many hormones, signaling molecules that the body releases to regulate processes at distant sites. Finally, like phospholipids, cholesterol molecules are found in the cell membrane, where their hydrophobic and hydrophilic regions help regulate the flow of substances into and out of the cell.

Prostaglandins

Like a hormone, a prostaglandin is one of a group of signaling molecules, but prostaglandins are derived from unsaturated fatty acids (see Figure 6c). One reason that the omega-3 fatty acids found in fish are beneficial is that they stimulate the production of certain prostaglandins that help regulate aspects of blood pressure and inflammation, and thereby reduce the risk for heart disease. Prostaglandins also sensitize nerves to pain. One class of pain-relieving medications called nonsteroidal anti-inflammatory drugs (NSAIDs) works by reducing the effects of prostaglandins.

Proteins

You might associate proteins with muscle tissue, but in fact, proteins are critical components of all tissues and organs. A protein is an organic molecule composed of amino acids linked by peptide bonds. Proteins include the keratin in the epidermis of skin that protects underlying tissues, the collagen found in the dermis of skin, in bones, and in the meninges that cover the brain and spinal cord. Proteins are also components of many of the body’s functional chemicals, including digestive enzymes in the digestive tract, antibodies, the neurotransmitters that neurons use to communicate with other cells, and the peptide-based hormones that regulate certain body functions (for instance, growth hormone). While carbohydrates and lipids are composed of hydrocarbons and oxygen, all proteins also contain nitrogen (N), and many contain sulfur (S), in addition to carbon, hydrogen, and oxygen.

Microstructure of Proteins

Proteins are polymers made up of nitrogen-containing monomers called amino acids. An amino acid is a molecule composed of an amino group and a carboxyl group, together with a variable side chain. Just 20 different amino acids contribute to nearly all of the thousands of different proteins important in human structure and function. Body proteins contain a unique combination of a few dozen to a few hundred of these 20 amino acid monomers. All 20 of these amino acids share a similar structure (Figure 7). All consist of a central carbon atom to which the following are bonded:

  • a hydrogen atom
  • an alkaline (basic) amino group NH2 (see Table 1)
  • an acidic carboxyl group COOH (see Table 1)
  • a variable group
[caption id="" align="aligncenter" width="320"]This figure shows the structure of an amino acid. Figure 8. Structure of an Amino Acid[/caption]

Notice that all amino acids contain both an acid (the carboxyl group) and a base (the amino group) (amine = “nitrogen-containing”). For this reason, they make excellent buffers, helping the body regulate acid–base balance. What distinguishes the 20 amino acids from one another is their variable group, which is referred to as a side chain or an R-group. This group can vary in size and can be polar or nonpolar, giving each amino acid its unique characteristics. For example, the side chains of two amino acids—cysteine and methionine—contain sulfur. Sulfur does not readily participate in hydrogen bonds, whereas all other amino acids do. This variation influences the way that proteins containing cysteine and methionine are assembled.

Amino acids join via dehydration synthesis to form protein polymers (Figure 8). The unique bond holding amino acids together is called a peptide bond. A peptide bond is a covalent bond between two amino acids that forms by dehydration synthesis. A peptide, in fact, is a very short chain of amino acids. Strands containing fewer than about 100 amino acids are generally referred to as polypeptides rather than proteins.

[caption id="" align="aligncenter" width="280"]This figure shows the formation of a peptide bond, highlighted in blue. Figure 8.Peptide Bond. Different amino acids join together to form peptides, polypeptides, or proteins via dehydration synthesis. The bonds between the amino acids are peptide bonds.[/caption]

The body is able to synthesize most of the amino acids from components of other molecules; however, nine cannot be synthesized and have to be consumed in the diet. These are known as the essential amino acids.

Free amino acids available for protein construction are said to reside in the amino acid pool within cells. Structures within cells use these amino acids when assembling proteins. If a particular essential amino acid is not available in sufficient quantities in the amino acid pool, however, synthesis of proteins containing it can slow or even cease.

Shape of Proteins

Just as a fork cannot be used to eat soup and a spoon cannot be used to spear meat, a protein’s shape is essential to its function. A protein’s shape is determined, most fundamentally, by the sequence of amino acids of which it is made (Figure 9a). The sequence is called the primary structure of the protein.

[caption id="" align="aligncenter" width="480"]This figure shows the secondary structure of peptides. The top panel shows a straight chain, the middle panel shows an alpha-helix and a beta sheet. The bottom panel shows the tertiary structure and fully folded protein. Figure 9. The Shape of Proteins. (a) The primary structure is the sequence of amino acids that make up the polypeptide chain. (b) The secondary structure, which can take the form of an alpha-helix or a beta-pleated sheet, is maintained by hydrogen bonds between amino acids in different regions of the original polypeptide strand. (c) The tertiary structure occurs as a result of further folding and bonding of the secondary structure. (d) The quaternary structure occurs as a result of interactions between two or more tertiary subunits. The example shown here is hemoglobin, a protein in red blood cells which transports oxygen to body tissues.[/caption]

Although some polypeptides exist as linear chains, most are twisted or folded into more complex secondary structures that form when bonding occurs between amino acids with different properties at different regions of the polypeptide. The most common secondary structure is a spiral called an alpha-helix. If you were to take a length of string and simply twist it into a spiral, it would not hold the shape. Similarly, a strand of amino acids could not maintain a stable spiral shape without the help of hydrogen bonds, which create bridges between different regions of the same strand (see Figure 9b). Less commonly, a polypeptide chain can form a beta-pleated sheet, in which hydrogen bonds form bridges between different regions of a single polypeptide that has folded back upon itself, or between two or more adjacent polypeptide chains.

The secondary structure of proteins further folds into a compact three-dimensional shape, referred to as the protein’s tertiary structure (see Figure 9c). In this configuration, amino acids that had been very distant in the primary chain can be brought quite close via hydrogen bonds or, in proteins containing cysteine, via disulfide bonds. A disulfide bond is a covalent bond between sulfur atoms in a polypeptide. Often, two or more separate polypeptides bond to form an even larger protein with a quaternary structure (see Figure 9d). The polypeptide subunits forming a quaternary structure can be identical or different. For instance, hemoglobin, the protein found in red blood cells is composed of four tertiary polypeptides, two of which are called alpha chains and two of which are called beta chains.

When they are exposed to extreme heat, acids, bases, and certain other substances, proteins will denature. Denaturation is a change in the structure of a molecule through physical or chemical means. Denatured proteins lose their functional shape and are no longer able to carry out their jobs. An everyday example of protein denaturation is the curdling of milk when acidic lemon juice is added.

The contribution of the shape of a protein to its function can hardly be exaggerated. For example, the long, slender shape of protein strands that make up muscle tissue is essential to their ability to contract (shorten) and relax (lengthen). As another example, bones contain long threads of a protein called collagen that acts as scaffolding upon which bone minerals are deposited. These elongated proteins, called fibrous proteins, are strong and durable and typically hydrophobic.

In contrast, globular proteins are globes or spheres that tend to be highly reactive and are hydrophilic. The hemoglobin proteins packed into red blood cells are an example (see Figure 9d); however, globular proteins are abundant throughout the body, playing critical roles in most body functions. Enzymes, introduced earlier as protein catalysts, are examples of this. The next section takes a closer look at the action of enzymes.

Proteins Function as Enzymes

If you were trying to type a paper, and every time you hit a key on your laptop there was a delay of six or seven minutes before you got a response, you would probably get a new laptop. In a similar way, without enzymes to catalyze chemical reactions, the human body would be nonfunctional. It functions only because enzymes function.

Enzymatic reactions—chemical reactions catalyzed by enzymes—begin when substrates bind to the enzyme. A substrate is a reactant in an enzymatic reaction. This occurs on regions of the enzyme known as active sites (Figure 10). Any given enzyme catalyzes just one type of chemical reaction. This characteristic, called specificity, is due to the fact that a substrate with a particular shape and electrical charge can bind only to an active site corresponding to that substrate.

[caption id="" align="aligncenter" width="520"]This image shows the steps in which an enzyme can act. The substrate is shown binding to the enzyme, forming a product, and the detachment of the product. Figure 10. Steps in an Enzymatic Reaction. (a) Substrates approach active sites on enzyme. (b) Substrates bind to active sites, producing an enzyme–substrate complex. (c) Changes internal to the enzyme–substrate complex facilitate interaction of the substrates. (d) Products are released and the enzyme returns to its original form, ready to facilitate another enzymatic reaction.[/caption]

Binding of a substrate produces an enzyme–substrate complex. It is likely that enzymes speed up chemical reactions in part because the enzyme–substrate complex undergoes a set of temporary and reversible changes that cause the substrates to be oriented toward each other in an optimal position to facilitate their interaction. This promotes increased reaction speed. The enzyme then releases the product(s), and resumes its original shape. The enzyme is then free to engage in the process again, and will do so as long as substrate remains.

Other Functions of Proteins

Advertisements for protein bars, powders, and shakes all say that protein is important in building, repairing, and maintaining muscle tissue, but the truth is that proteins contribute to all body tissues, from the skin to the brain cells. Also, certain proteins act as hormones, chemical messengers that help regulate body functions, For example, growth hormone is important for skeletal growth, among other roles.

As was noted earlier, the basic and acidic components enable proteins to function as buffers in maintaining acid–base balance, but they also help regulate fluid–electrolyte balance. Proteins attract fluid, and a healthy concentration of proteins in the blood, the cells, and the spaces between cells helps ensure a balance of fluids in these various “compartments.” Moreover, proteins in the cell membrane help to transport electrolytes in and out of the cell, keeping these ions in a healthy balance. Like lipids, proteins can bind with carbohydrates. They can thereby produce glycoproteins or proteoglycans, both of which have many functions in the body.

The body can use proteins for energy when carbohydrate and fat intake is inadequate, and stores of glycogen and adipose tissue become depleted. However, since there is no storage site for protein except functional tissues, using protein for energy causes tissue breakdown, and results in body wasting.

Nucleotides

The fourth type of organic compound important to human structure and function are the nucleotides (Figure 11). A nucleotide is one of a class of organic compounds composed of three subunits:

  • one or more phosphate groups
  • a pentose sugar: either deoxyribose or ribose
  • a nitrogen-containing base: adenine, cytosine, guanine, thymine, or uracil

Nucleotides can be assembled into nucleic acids (DNA or RNA) or the energy compound adenosine triphosphate.

[caption id="" align="aligncenter" width="520"]This figure shows the structure of nucleotides. Figure 11. Nucleotides. (a) The building blocks of all nucleotides are one or more phosphate groups, a pentose sugar, and a nitrogen-containing base. (b) The nitrogen-containing bases of nucleotides. (c) The two pentose sugars of DNA and RNA.[/caption]

Nucleic Acids

The nucleic acids differ in their type of pentose sugar. Deoxyribonucleic acid (DNA) is nucleotide that stores genetic information. DNA contains deoxyribose (so-called because it has one less atom of oxygen than ribose) plus one phosphate group and one nitrogen-containing base. The “choices” of base for DNA are adenine, cytosine, guanine, and thymine. Ribonucleic acid (RNA) is a ribose-containing nucleotide that helps manifest the genetic code as protein. RNA contains ribose, one phosphate group, and one nitrogen-containing base, but the “choices” of base for RNA are adenine, cytosine, guanine, and uracil.

The nitrogen-containing bases adenine and guanine are classified as purines. A purine is a nitrogen-containing molecule with a double ring structure, which accommodates several nitrogen atoms. The bases cytosine, thymine (found in DNA only) and uracil (found in RNA only) are pyramidines. A pyramidine is a nitrogen-containing base with a single ring structure

Bonds formed by dehydration synthesis between the pentose sugar of one nucleic acid monomer and the phosphate group of another form a “backbone,” from which the components’ nitrogen-containing bases protrude. In DNA, two such backbones attach at their protruding bases via hydrogen bonds. These twist to form a shape known as a double helix (Figure 12). The sequence of nitrogen-containing bases within a strand of DNA form the genes that act as a molecular code instructing cells in the assembly of amino acids into proteins. Humans have almost 22,000 genes in their DNA, locked up in the 46 chromosomes inside the nucleus of each cell (except red blood cells which lose their nuclei during development). These genes carry the genetic code to build one’s body, and are unique for each individual except identical twins.

[caption id="" align="aligncenter" width="320"]This figure shows a double helix. Figure 12. DNA. In the DNA double helix, two strands attach via hydrogen bonds between the bases of the component nucleotides.[/caption]

In contrast, RNA consists of a single strand of sugar-phosphate backbone studded with bases. Messenger RNA (mRNA) is created during protein synthesis to carry the genetic instructions from the DNA to the cell’s protein manufacturing plants in the cytoplasm, the ribosomes.

Adenosine Triphosphate

The nucleotide adenosine triphosphate (ATP), is composed of a ribose sugar, an adenine base, and three phosphate groups (Figure 13). ATP is classified as a high energy compound because the two covalent bonds linking its three phosphates store a significant amount of potential energy. In the body, the energy released from these high energy bonds helps fuel the body’s activities, from muscle contraction to the transport of substances in and out of cells to anabolic chemical reactions.

[caption id="" align="aligncenter" width="420"]This figure shows the structure of ATP. Figure 13. Structure of Adenosine Triphosphate (ATP).[/caption]

When a phosphate group is cleaved from ATP, the products are adenosine diphosphate (ADP) and inorganic phosphate (Pi). This hydrolysis reaction can be written:

ATP + H2O → ADP + Pi + energy

Removal of a second phosphate leaves adenosine monophosphate (AMP) and two phosphate groups. Again, these reactions also liberate the energy that had been stored in the phosphate-phosphate bonds. They are reversible, too, as when ADP undergoes phosphorylation. Phosphorylation is the addition of a phosphate group to an organic compound, in this case, resulting in ATP. In such cases, the same level of energy that had been released during hydrolysis must be reinvested to power dehydration synthesis.

Cells can also transfer a phosphate group from ATP to another organic compound. For example, when glucose first enters a cell, a phosphate group is transferred from ATP, forming glucose phosphate (C6H12O6—P) and ADP. Once glucose is phosphorylated in this way, it can be stored as glycogen or metabolized for immediate energy.

[caption id="attachment_3052" align="aligncenter" width="150"] Watch this amoeba sisters video to learn more about biomolecules![/caption]
]]> 2461 2429 4 0 22.7 Embryonic Development of the Respiratory System https://pressbooks.bccampus.ca/apdouglas2017/chapter/22-7-embryonic-development-of-the-respiratory-system/ Wed, 02 Aug 2017 22:04:34 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2633

Learning Objectives

By the end of this section, you will be able to:
  • Create a timeline of the phases of respiratory development in the fetus
  • Propose reasons for fetal breathing movements
  • Explain how the lungs become inflated after birth

Development of the respiratory system begins early in the fetus. It is a complex process that includes many structures, most of which arise from the endoderm. Towards the end of development, the fetus can be observed making breathing movements. Until birth, however, the mother provides all of the oxygen to the fetus as well as removes all of the fetal carbon dioxide via the placenta.

Time Line

The development of the respiratory system begins at about week 4 of gestation. By week 28, enough alveoli have matured that a baby born prematurely at this time can usually breathe on its own. The respiratory system, however, is not fully developed until early childhood, when a full complement of mature alveoli is present.

Weeks 4–7

Respiratory development in the embryo begins around week 4. Ectodermal tissue from the anterior head region invaginates posteriorly to form olfactory pits, which fuse with endodermal tissue of the developing pharynx. An olfactory pit is one of a pair of structures that will enlarge to become the nasal cavity. At about this same time, the lung bud forms. The lung bud is a dome-shaped structure composed of tissue that bulges from the foregut. The foregut is endoderm just inferior to the pharyngeal pouches. The laryngotracheal bud is a structure that forms from the longitudinal extension of the lung bud as development progresses. The portion of this structure nearest the pharynx becomes the trachea, whereas the distal end becomes more bulbous, forming bronchial buds. A bronchial bud is one of a pair of structures that will eventually become the bronchi and all other lower respiratory structures (Figure 1).

[caption id="" align="aligncenter" width="440"]This flowchart shows the embryonic development of the respiratory system and correlates the gestational age to the appearance of new features. Figure 1. Development of the Lower Respiratory System.[/caption]

Weeks 7–16

Bronchial buds continue to branch as development progresses until all of the segmental bronchi have been formed. Beginning around week 13, the lumens of the bronchi begin to expand in diameter. By week 16, respiratory bronchioles form. The fetus now has all major lung structures involved in the airway.

Weeks 16–24

Once the respiratory bronchioles form, further development includes extensive vascularization, or the development of the blood vessels, as well as the formation of alveolar ducts and alveolar precursors. At about week 19, the respiratory bronchioles have formed. In addition, cells lining the respiratory structures begin to differentiate to form type I and type II pneumocytes. Once type II cells have differentiated, they begin to secrete small amounts of pulmonary surfactant. Around week 20, fetal breathing movements may begin.

Weeks 24–Term

Major growth and maturation of the respiratory system occurs from week 24 until term. More alveolar precursors develop, and larger amounts of pulmonary surfactant are produced. Surfactant levels are not generally adequate to create effective lung compliance until about the eighth month of pregnancy. The respiratory system continues to expand, and the surfaces that will form the respiratory membrane develop further. At this point, pulmonary capillaries have formed and continue to expand, creating a large surface area for gas exchange. The major milestone of respiratory development occurs at around week 28, when sufficient alveolar precursors have matured so that a baby born prematurely at this time can usually breathe on its own. However, alveoli continue to develop and mature into childhood. A full complement of functional alveoli does not appear until around 8 years of age.

Fetal “Breathing”

Although the function of fetal breathing movements is not entirely clear, they can be observed starting at 20–21 weeks of development. Fetal breathing movements involve muscle contractions that cause the inhalation of amniotic fluid and exhalation of the same fluid, with pulmonary surfactant and mucus. Fetal breathing movements are not continuous and may include periods of frequent movements and periods of no movements. Maternal factors can influence the frequency of breathing movements. For example, high blood glucose levels, called hyperglycemia, can boost the number of breathing movements. Conversely, low blood glucose levels, called hypoglycemia, can reduce the number of fetal breathing movements. Tobacco use is also known to lower fetal breathing rates. Fetal breathing may help tone the muscles in preparation for breathing movements once the fetus is born. It may also help the alveoli to form and mature. Fetal breathing movements are considered a sign of robust health.

Birth

Prior to birth, the lungs are filled with amniotic fluid, mucus, and surfactant. As the fetus is squeezed through the birth canal, the fetal thoracic cavity is compressed, expelling much of this fluid. Some fluid remains, however, but is rapidly absorbed by the body shortly after birth. The first inhalation occurs within 10 seconds after birth and not only serves as the first inspiration, but also acts to inflate the lungs. Pulmonary surfactant is critical for inflation to occur, as it reduces the surface tension of the alveoli. Preterm birth around 26 weeks frequently results in severe respiratory distress, although with current medical advancements, some babies may survive. Prior to 26 weeks, sufficient pulmonary surfactant is not produced, and the surfaces for gas exchange have not formed adequately; therefore, survival is low.

Disorders of the…

Respiratory System: Respiratory Distress Syndrome Respiratory distress syndrome (RDS) primarily occurs in infants born prematurely. Up to 50 percent of infants born between 26 and 28 weeks and fewer than 30 percent of infants born between 30 and 31 weeks develop RDS. RDS results from insufficient production of pulmonary surfactant, thereby preventing the lungs from properly inflating at birth. A small amount of pulmonary surfactant is produced beginning at around 20 weeks; however, this is not sufficient for inflation of the lungs. As a result, dyspnea occurs and gas exchange cannot be performed properly. Blood oxygen levels are low, whereas blood carbon dioxide levels and pH are high.

The primary cause of RDS is premature birth, which may be due to a variety of known or unknown causes. Other risk factors include gestational diabetes, cesarean delivery, second-born twins, and family history of RDS. The presence of RDS can lead to other serious disorders, such as septicemia (infection of the blood) or pulmonary hemorrhage. Therefore, it is important that RDS is immediately recognized and treated to prevent death and reduce the risk of developing other disorders.

Medical advances have resulted in an improved ability to treat RDS and support the infant until proper lung development can occur. At the time of delivery, treatment may include resuscitation and intubation if the infant does not breathe on his or her own. These infants would need to be placed on a ventilator to mechanically assist with the breathing process. If spontaneous breathing occurs, application of nasal continuous positive airway pressure (CPAP) may be required. In addition, pulmonary surfactant is typically administered. Death due to RDS has been reduced by 50 percent due to the introduction of pulmonary surfactant therapy. Other therapies may include corticosteroids, supplemental oxygen, and assisted ventilation. Supportive therapies, such as temperature regulation, nutritional support, and antibiotics, may be administered to the premature infant as well.

]]> 2633 2618 4 0 28.4 Maternal Changes During Pregnancy, Labor, and Birth https://pressbooks.bccampus.ca/apdouglas2017/chapter/28-4-maternal-changes-during-pregnancy-labor-and-birth/ Wed, 02 Aug 2017 22:04:57 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2812

Learning Objectives

By the end of this section, you will be able to:
  • Explain how estrogen, progesterone, and hCG are involved in maintaining pregnancy
  • List the contributors to weight gain during pregnancy
  • Describe the major changes to the maternal digestive, circulatory, and integumentary systems during pregnancy
  • Summarize the events leading to labor
  • Identify and describe each of the three stages of childbirth

A full-term pregnancy lasts approximately 270 days (approximately 38.5 weeks) from conception to birth. Because it is easier to remember the first day of the last menstrual period (LMP) than to estimate the date of conception, obstetricians set the due date as 284 days (approximately 40.5 weeks) from the LMP. This assumes that conception occurred on day 14 of the woman’s cycle, which is usually a good approximation. The 40 weeks of an average pregnancy are usually discussed in terms of three trimesters, each approximately 13 weeks. During the second and third trimesters, the pre-pregnancy uterus—about the size of a fist—grows dramatically to contain the fetus, causing a number of anatomical changes in the mother (Figure 1).

[caption id="" align="aligncenter" width="300"]This figure shows a woman’s body and marks the size of the uterus as it grows throughout pregnancy. Figure 1. Size of Uterus throughout Pregnancy. The uterus grows throughout pregnancy to accommodate the fetus.[/caption]

Effects of Hormones

Virtually all of the effects of pregnancy can be attributed in some way to the influence of hormones—particularly estrogens, progesterone, and hCG. During weeks 7–12 from the LMP, the pregnancy hormones are primarily generated by the corpus luteum. Progesterone secreted by the corpus luteum stimulates the production of decidual cells of the endometrium that nourish the blastocyst before placentation. As the placenta develops and the corpus luteum degenerates during weeks 12–17, the placenta gradually takes over as the endocrine organ of pregnancy.

The placenta converts weak androgens secreted by the maternal and fetal adrenal glands to estrogens, which are necessary for pregnancy to progress. Estrogen levels climb throughout the pregnancy, increasing 30-fold by childbirth. Estrogens have the following actions:

  • They suppress FSH and LH production, effectively preventing ovulation. (This function is the biological basis of hormonal birth control pills.)
  • They induce the growth of fetal tissues and are necessary for the maturation of the fetal lungs and liver.
  • They promote fetal viability by regulating progesterone production and triggering fetal synthesis of cortisol, which helps with the maturation of the lungs, liver, and endocrine organs such as the thyroid gland and adrenal gland.
  • They stimulate maternal tissue growth, leading to uterine enlargement and mammary duct expansion and branching.
Relaxin, another hormone secreted by the corpus luteum and then by the placenta, helps prepare the mother’s body for childbirth. It increases the elasticity of the symphysis pubis joint and pelvic ligaments, making room for the growing fetus and allowing expansion of the pelvic outlet for childbirth. Relaxin also helps dilate the cervix during labor.

The placenta takes over the synthesis and secretion of progesterone throughout pregnancy as the corpus luteum degenerates. Like estrogen, progesterone suppresses FSH and LH. It also inhibits uterine contractions, protecting the fetus from preterm birth. This hormone decreases in late gestation, allowing uterine contractions to intensify and eventually progress to true labor. The placenta also produces hCG. In addition to promoting survival of the corpus luteum, hCG stimulates the male fetal gonads to secrete testosterone, which is essential for the development of the male reproductive system.

The anterior pituitary enlarges and ramps up its hormone production during pregnancy, raising the levels of thyrotropin, prolactin, and adrenocorticotropic hormone (ACTH). Thyrotropin, in conjunction with placental hormones, increases the production of thyroid hormone, which raises the maternal metabolic rate. This can markedly augment a pregnant woman’s appetite and cause hot flashes. Prolactin stimulates enlargement of the mammary glands in preparation for milk production. ACTH stimulates maternal cortisol secretion, which contributes to fetal protein synthesis. In addition to the pituitary hormones, increased parathyroid levels mobilize calcium from maternal bones for fetal use.

Weight Gain

The second and third trimesters of pregnancy are associated with dramatic changes in maternal anatomy and physiology. The most obvious anatomical sign of pregnancy is the dramatic enlargement of the abdominal region, coupled with maternal weight gain. This weight results from the growing fetus as well as the enlarged uterus, amniotic fluid, and placenta. Additional breast tissue and dramatically increased blood volume also contribute to weight gain (Table 2). Surprisingly, fat storage accounts for only approximately 2.3 kg (5 lbs) in a normal pregnancy and serves as a reserve for the increased metabolic demand of breastfeeding.

During the first trimester, the mother does not need to consume additional calories to maintain a healthy pregnancy. However, a weight gain of approximately 0.45 kg (1 lb) per month is common. During the second and third trimesters, the mother’s appetite increases, but it is only necessary for her to consume an additional 300 calories per day to support the growing fetus. Most women gain approximately 0.45 kg (1 lb) per week.
Contributors to Weight Gain During Pregnancy (Table 2)
Component Weight (kg) Weight (lb)
Fetus 3.2–3.6 7–8
Placenta and fetal membranes 0.9–1.8 2–4
Amniotic fluid 0.9–1.4 2–3
Breast tissue 0.9–1.4 2–3
Blood 1.4 4
Fat 0.9–4.1 3–9
Uterus 0.9–2.3 2–5
Total 10–16.3 22–36

Changes in Organ Systems During Pregnancy

As the woman’s body adapts to pregnancy, characteristic physiologic changes occur. These changes can sometimes prompt symptoms often referred to collectively as the common discomforts of pregnancy.

Digestive and Urinary System Changes

Nausea and vomiting, sometimes triggered by an increased sensitivity to odors, are common during the first few weeks to months of pregnancy. This phenomenon is often referred to as “morning sickness,” although the nausea may persist all day. The source of pregnancy nausea is thought to be the increased circulation of pregnancy-related hormones, specifically circulating estrogen, progesterone, and hCG. Decreased intestinal peristalsis may also contribute to nausea. By about week 12 of pregnancy, nausea typically subsides.

A common gastrointestinal complaint during the later stages of pregnancy is gastric reflux, or heartburn, which results from the upward, constrictive pressure of the growing uterus on the stomach. The same decreased peristalsis that may contribute to nausea in early pregnancy is also thought to be responsible for pregnancy-related constipation as pregnancy progresses.

The downward pressure of the uterus also compresses the urinary bladder, leading to frequent urination. The problem is exacerbated by increased urine production. In addition, the maternal urinary system processes both maternal and fetal wastes, further increasing the total volume of urine.

Circulatory System Changes

Blood volume increases substantially during pregnancy, so that by childbirth, it exceeds its preconception volume by 30 percent, or approximately 1–2 liters. The greater blood volume helps to manage the demands of fetal nourishment and fetal waste removal. In conjunction with increased blood volume, the pulse and blood pressure also rise moderately during pregnancy. As the fetus grows, the uterus compresses underlying pelvic blood vessels, hampering venous return from the legs and pelvic region. As a result, many pregnant women develop varicose veins or hemorrhoids.

Respiratory System Changes

During the second half of pregnancy, the respiratory minute volume (volume of gas inhaled or exhaled by the lungs per minute) increases by 50 percent to compensate for the oxygen demands of the fetus and the increased maternal metabolic rate. The growing uterus exerts upward pressure on the diaphragm, decreasing the volume of each inspiration and potentially causing shortness of breath, or dyspnea. During the last several weeks of pregnancy, the pelvis becomes more elastic, and the fetus descends lower in a process called lightening. This typically ameliorates dyspnea.

The respiratory mucosa swell in response to increased blood flow during pregnancy, leading to nasal congestion and nose bleeds, particularly when the weather is cold and dry. Humidifier use and increased fluid intake are often recommended to counteract congestion.

Integumentary System Changes

The dermis stretches extensively to accommodate the growing uterus, breast tissue, and fat deposits on the thighs and hips. Torn connective tissue beneath the dermis can cause striae (stretch marks) on the abdomen, which appear as red or purple marks during pregnancy that fade to a silvery white color in the months after childbirth.

An increase in melanocyte-stimulating hormone, in conjunction with estrogens, darkens the areolae and creates a line of pigment from the umbilicus to the pubis called the linea nigra (Figure 2). Melanin production during pregnancy may also darken or discolor skin on the face to create a chloasma, or “mask of pregnancy.”

[caption id="" align="aligncenter" width="225"]This photo shows a dark line below a woman’s navel. Figure 2. Linea Nigra. The linea nigra, a dark medial line running from the umbilicus to the pubis, forms during pregnancy and persists for a few weeks following childbirth. The linea nigra shown here corresponds to a pregnancy that is 22 weeks along.[/caption]

Physiology of Labor

Childbirth, or parturition, typically occurs within a week of a woman’s due date, unless the woman is pregnant with more than one fetus, which usually causes her to go into labor early. As a pregnancy progresses into its final weeks, several physiological changes occur in response to hormones that trigger labor.

First, recall that progesterone inhibits uterine contractions throughout the first several months of pregnancy. As the pregnancy enters its seventh month, progesterone levels plateau and then drop. Estrogen levels, however, continue to rise in the maternal circulation (Figure 3). The increasing ratio of estrogen to progesterone makes the myometrium (the uterine smooth muscle) more sensitive to stimuli that promote contractions (because progesterone no longer inhibits them). Moreover, in the eighth month of pregnancy, fetal cortisol rises, which boosts estrogen secretion by the placenta and further overpowers the uterine-calming effects of progesterone. Some women may feel the result of the decreasing levels of progesterone in late pregnancy as weak and irregular peristaltic Braxton Hicks contractions, also called false labor. These contractions can often be relieved with rest or hydration.

[caption id="" align="aligncenter" width="400"]A graph hormone concentration versus week of pregnancy shows how three hormones vary throughout pregnancy. Figure 3. Hormones Initiating Labor. A positive feedback loop of hormones works to initiate labor.[/caption]
A common sign that labor will be short is the so-called “bloody show.” During pregnancy, a plug of mucus accumulates in the cervical canal, blocking the entrance to the uterus. Approximately 1–2 days prior to the onset of true labor, this plug loosens and is expelled, along with a small amount of blood.

Meanwhile, the posterior pituitary has been boosting its secretion of oxytocin, a hormone that stimulates the contractions of labor. At the same time, the myometrium increases its sensitivity to oxytocin by expressing more receptors for this hormone. As labor nears, oxytocin begins to stimulate stronger, more painful uterine contractions, which—in a positive feedback loop—stimulate the secretion of prostaglandins from fetal membranes. Like oxytocin, prostaglandins also enhance uterine contractile strength. The fetal pituitary also secretes oxytocin, which increases prostaglandins even further. Given the importance of oxytocin and prostaglandins to the initiation and maintenance of labor, it is not surprising that, when a pregnancy is not progressing to labor and needs to be induced, a pharmaceutical version of these compounds (called pitocin) is administered by intravenous drip.

Finally, stretching of the myometrium and cervix by a full-term fetus in the vertex (head-down) position is regarded as a stimulant to uterine contractions. The sum of these changes initiates the regular contractions known as true labor, which become more powerful and more frequent with time. The pain of labor is attributed to myometrial hypoxia during uterine contractions.

Stages of Childbirth

The process of childbirth can be divided into three stages: cervical dilation, expulsion of the newborn, and afterbirth (Figure 4).

Cervical Dilation

For vaginal birth to occur, the cervix must dilate fully to 10 cm in diameter—wide enough to deliver the newborn’s head. The dilation stage is the longest stage of labor and typically takes 6–12 hours. However, it varies widely and may take minutes, hours, or days, depending in part on whether the mother has given birth before; in each subsequent labor, this stage tends to be shorter.

[caption id="" align="aligncenter" width="420"]This multi-part figure shows the different stages of childbirth. The top panel shows dilation, the middle panel shows birth and the bottom panel shows afterbirth delivery. Figure 4. Stages of Childbirth. The stages of childbirth include Stage 1, early cervical dilation; Stage 2, full dilation and expulsion of the newborn; and Stage 3, delivery of the placenta and associated fetal membranes. (The position of the newborn’s shoulder is described relative to the mother.)[/caption]

True labor progresses in a positive feedback loop in which uterine contractions stretch the cervix, causing it to dilate and efface, or become thinner. Cervical stretching induces reflexive uterine contractions that dilate and efface the cervix further. In addition, cervical dilation boosts oxytocin secretion from the pituitary, which in turn triggers more powerful uterine contractions. When labor begins, uterine contractions may occur only every 3–30 minutes and last only 20–40 seconds; however, by the end of this stage, contractions may occur as frequently as every 1.5–2 minutes and last for a full minute.

Each contraction sharply reduces oxygenated blood flow to the fetus. For this reason, it is critical that a period of relaxation occur after each contraction. Fetal distress, measured as a sustained decrease or increase in the fetal heart rate, can result from severe contractions that are too powerful or lengthy for oxygenated blood to be restored to the fetus. Such a situation can be cause for an emergency birth with vacuum, forceps, or surgically by Caesarian section.

The amniotic membranes rupture before the onset of labor in about 12 percent of women; they typically rupture at the end of the dilation stage in response to excessive pressure from the fetal head entering the birth canal.

Expulsion Stage

The expulsion stage begins when the fetal head enters the birth canal and ends with birth of the newborn. It typically takes up to 2 hours, but it can last longer or be completed in minutes, depending in part on the orientation of the fetus. The vertex presentation known as the occiput anterior vertex is the most common presentation and is associated with the greatest ease of vaginal birth. The fetus faces the maternal spinal cord and the smallest part of the head (the posterior aspect called the occiput) exits the birth canal first.

In fewer than 5 percent of births, the infant is oriented in the breech presentation, or buttocks down. In a complete breech, both legs are crossed and oriented downward. In a frank breech presentation, the legs are oriented upward. Before the 1960s, it was common for breech presentations to be delivered vaginally. Today, most breech births are accomplished by Caesarian section.

Vaginal birth is associated with significant stretching of the vaginal canal, the cervix, and the perineum. Until recent decades, it was routine procedure for an obstetrician to numb the perineum and perform an episiotomy, an incision in the posterior vaginal wall and perineum. The perineum is now more commonly allowed to tear on its own during birth. Both an episiotomy and a perineal tear need to be sutured shortly after birth to ensure optimal healing. Although suturing the jagged edges of a perineal tear may be more difficult than suturing an episiotomy, tears heal more quickly, are less painful, and are associated with less damage to the muscles around the vagina and rectum.

Upon birth of the newborn’s head, an obstetrician will aspirate mucus from the mouth and nose before the newborn’s first breath. Once the head is birthed, the rest of the body usually follows quickly. The umbilical cord is then double-clamped, and a cut is made between the clamps. This completes the second stage of childbirth.

Afterbirth

The delivery of the placenta and associated membranes, commonly referred to as the afterbirth, marks the final stage of childbirth. After expulsion of the newborn, the myometrium continues to contract. This movement shears the placenta from the back of the uterine wall. It is then easily delivered through the vagina. Continued uterine contractions then reduce blood loss from the site of the placenta. Delivery of the placenta marks the beginning of the postpartum period—the period of approximately 6 weeks immediately following childbirth during which the mother’s body gradually returns to a non-pregnant state. If the placenta does not birth spontaneously within approximately 30 minutes, it is considered retained, and the obstetrician may attempt manual removal. If this is not successful, surgery may be required.

It is important that the obstetrician examines the expelled placenta and fetal membranes to ensure that they are intact. If fragments of the placenta remain in the uterus, they can cause postpartum hemorrhage. Uterine contractions continue for several hours after birth to return the uterus to its pre-pregnancy size in a process called involution, which also allows the mother’s abdominal organs to return to their pre-pregnancy locations. Breastfeeding facilitates this process.

Although postpartum uterine contractions limit blood loss from the detachment of the placenta, the mother does experience a postpartum vaginal discharge called lochia. This is made up of uterine lining cells, erythrocytes, leukocytes, and other debris. Thick, dark, lochia rubra (red lochia) typically continues for 2–3 days, and is replaced by lochia serosa, a thinner, pinkish form that continues until about the tenth postpartum day. After this period, a scant, creamy, or watery discharge called lochia alba (white lochia) may continue for another 1–2 weeks.

]]> 2812 2785 4 0 28.5 Adjustments of the Infant at Birth and Postnatal Stages https://pressbooks.bccampus.ca/apdouglas2017/chapter/28-5-adjustments-of-the-infant-at-birth-and-postnatal-stages/ Wed, 02 Aug 2017 22:05:07 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2814

Learning Objectives

By the end of this section, you will be able to:
  • Discuss the importance of an infant’s first breath
  • Explain the closing of the cardiac shunts
  • Describe thermoregulation in the newborn
  • Summarize the importance of intestinal flora in the newborn
From a fetal perspective, the process of birth is a crisis. In the womb, the fetus was snuggled in a soft, warm, dark, and quiet world. The placenta provided nutrition and oxygen continuously. Suddenly, the contractions of labor and vaginal childbirth forcibly squeeze the fetus through the birth canal, limiting oxygenated blood flow during contractions and shifting the skull bones to accommodate the small space. After birth, the newborn’s system must make drastic adjustments to a world that is colder, brighter, and louder, and where he or she will experience hunger and thirst. The neonatal period (neo- = “new”; -natal = “birth”) spans the first to the thirtieth day of life outside of the uterus.

Respiratory Adjustments

Although the fetus “practices” breathing by inhaling amniotic fluid in utero, there is no air in the uterus and thus no true opportunity to breathe. (There is also no need to breathe because the placenta supplies the fetus with all the oxygenated blood it needs.) During gestation, the partially collapsed lungs are filled with amniotic fluid and exhibit very little metabolic activity. Several factors stimulate newborns to take their first breath at birth. First, labor contractions temporarily constrict umbilical blood vessels, reducing oxygenated blood flow to the fetus and elevating carbon dioxide levels in the blood. High carbon dioxide levels cause acidosis and stimulate the respiratory center in the brain, triggering the newborn to take a breath.

The first breath typically is taken within 10 seconds of birth, after mucus is aspirated from the infant’s mouth and nose. The first breaths inflate the lungs to nearly full capacity and dramatically decrease lung pressure and resistance to blood flow, causing a major circulatory reconfiguration. Pulmonary alveoli open, and alveolar capillaries fill with blood. Amniotic fluid in the lungs drains or is absorbed, and the lungs immediately take over the task of the placenta, exchanging carbon dioxide for oxygen by the process of respiration.

Circulatory Adjustments

The process of clamping and cutting the umbilical cord collapses the umbilical blood vessels. In the absence of medical assistance, this occlusion would occur naturally within 20 minutes of birth because the Wharton’s jelly within the umbilical cord would swell in response to the lower temperature outside of the mother’s body, and the blood vessels would constrict. Natural occlusion has occurred when the umbilical cord is no longer pulsating. For the most part, the collapsed vessels atrophy and become fibrotic remnants, existing in the mature circulatory system as ligaments of the abdominal wall and liver. The ductus venosus degenerates to become the ligamentum venosum beneath the liver. Only the proximal sections of the two umbilical arteries remain functional, taking on the role of supplying blood to the upper part of the bladder (Figure 1).

[caption id="" align="aligncenter" width="500"]This figure illustrates the circulatory system in a newborn. The left image in both panels shows the blood circulation before birth and the right image shows the blood circulation after birth. Figure 1. Neonatal Circulatory System. A newborn’s circulatory system reconfigures immediately after birth. The three fetal shunts have been closed permanently, facilitating blood flow to the liver and lungs.[/caption]

The newborn’s first breath is vital to initiate the transition from the fetal to the neonatal circulatory pattern. Inflation of the lungs decreases blood pressure throughout the pulmonary system, as well as in the right atrium and ventricle. In response to this pressure change, the flow of blood temporarily reverses direction through the foramen ovale, moving from the left to the right atrium, and blocking the shunt with two flaps of tissue. Within 1 year, the tissue flaps usually fuse over the shunt, turning the foramen ovale into the fossa ovalis. The ductus arteriosus constricts as a result of increased oxygen concentration, and becomes the ligamentum arteriosum. Closing of the ductus arteriosus ensures that all blood pumped to the pulmonary circuit will be oxygenated by the newly functional neonatal lungs.

Thermoregulatory Adjustments

The fetus floats in warm amniotic fluid that is maintained at a temperature of approximately 98.6°F with very little fluctuation. Birth exposes newborns to a cooler environment in which they have to regulate their own body temperature. Newborns have a higher ratio of surface area to volume than adults. This means that their body has less volume throughout which to produce heat, and more surface area from which to lose heat. As a result, newborns produce heat more slowly and lose it more quickly. Newborns also have immature musculature that limits their ability to generate heat by shivering. Moreover, their nervous systems are underdeveloped, so they cannot quickly constrict superficial blood vessels in response to cold. They also have little subcutaneous fat for insulation. All these factors make it harder for newborns to maintain their body temperature.

Newborns, however, do have a special method for generating heat: nonshivering thermogenesis, which involves the breakdown of brown adipose tissue, or brown fat, which is distributed over the back, chest, and shoulders. Brown fat differs from the more familiar white fat in two ways:

  • It is highly vascularized. This allows for faster delivery of oxygen, which leads to faster cellular respiration.
  • It is packed with a special type of mitochondria that are able to engage in cellular respiration reactions that produce less ATP and more heat than standard cellular respiration reactions.

The breakdown of brown fat occurs automatically upon exposure to cold, so it is an important heat regulator in newborns. During fetal development, the placenta secretes inhibitors that prevent metabolism of brown adipose fat and promote its accumulation in preparation for birth.

Gastrointestinal and Urinary Adjustments

In adults, the gastrointestinal tract harbors bacterial flora—trillions of bacteria that aid in digestion, produce vitamins, and protect from the invasion or replication of pathogens. In stark contrast, the fetal intestine is sterile. The first consumption of breast milk or formula floods the neonatal gastrointestinal tract with beneficial bacteria that begin to establish the bacterial flora.

The fetal kidneys filter blood and produce urine, but the neonatal kidneys are still immature and inefficient at concentrating urine. Therefore, newborns produce very dilute urine, making it particularly important for infants to obtain sufficient fluids from breast milk or formula.

Homeostatic Imbalances

Homeostasis in the Newborn: Apgar Score In the minutes following birth, a newborn must undergo dramatic systemic changes to be able to survive outside the womb. An obstetrician, midwife, or nurse can estimate how well a newborn is doing by obtaining an Apgar score. The Apgar score was introduced in 1952 by the anesthesiologist Dr. Virginia Apgar as a method to assess the effects on the newborn of anesthesia given to the laboring mother. Healthcare providers now use it to assess the general wellbeing of the newborn, whether or not analgesics or anesthetics were used.

Five criteria—skin color, heart rate, reflex, muscle tone, and respiration—are assessed, and each criterion is assigned a score of 0, 1, or 2. Scores are taken at 1 minute after birth and again at 5 minutes after birth. Each time that scores are taken, the five scores are added together. High scores (out of a possible 10) indicate the baby has made the transition from the womb well, whereas lower scores indicate that the baby may be in distress.

The technique for determining an Apgar score is quick and easy, painless for the newborn, and does not require any instruments except for a stethoscope. A convenient way to remember the five scoring criteria is to apply the mnemonic APGAR, for “appearance” (skin color), “pulse” (heart rate), “grimace” (reflex), “activity” (muscle tone), and “respiration.”

Of the five Apgar criteria, heart rate and respiration are the most critical. Poor scores for either of these measurements may indicate the need for immediate medical attention to resuscitate or stabilize the newborn. In general, any score lower than 7 at the 5-minute mark indicates that medical assistance may be needed. A total score below 5 indicates an emergency situation. Normally, a newborn will get an intermediate score of 1 for some of the Apgar criteria and will progress to a 2 by the 5-minute assessment. Scores of 8 or above are normal.

]]> 2814 2785 5 0 28.6 Lactation https://pressbooks.bccampus.ca/apdouglas2017/chapter/28-6-lactation/ Wed, 02 Aug 2017 22:05:17 +0000 https://pressbooks.bccampus.ca/apdouglas2017/?post_type=chapter&p=2816

Learning Objectives

By the end of this section, you will be able to:
  • Describe the structure of the lactating breast
  • Summarize the process of lactation
  • Explain how the composition of breast milk changes during the first days of lactation and in the course of a single feeding
Lactation is the process by which milk is synthesized and secreted from the mammary glands of the postpartum female breast in response to an infant sucking at the nipple. Breast milk provides ideal nutrition and passive immunity for the infant, encourages mild uterine contractions to return the uterus to its pre-pregnancy size (i.e., involution), and induces a substantial metabolic increase in the mother, consuming the fat reserves stored during pregnancy.

Structure of the Lactating Breast

Mammary glands are modified sweat glands. The non-pregnant and non-lactating female breast is composed primarily of adipose and collagenous tissue, with mammary glands making up a very minor proportion of breast volume. The mammary gland is composed of milk-transporting lactiferous ducts, which expand and branch extensively during pregnancy in response to estrogen, growth hormone, cortisol, and prolactin. Moreover, in response to progesterone, clusters of breast alveoli bud from the ducts and expand outward toward the chest wall. Breast alveoli are balloon-like structures lined with milk-secreting cuboidal cells, or lactocytes, that are surrounded by a net of contractile myoepithelial cells. Milk is secreted from the lactocytes, fills the alveoli, and is squeezed into the ducts. Clusters of alveoli that drain to a common duct are called lobules; the lactating female has 12–20 lobules organized radially around the nipple. Milk drains from lactiferous ducts into lactiferous sinuses that meet at 4 to 18 perforations in the nipple, called nipple pores. The small bumps of the areola (the darkened skin around the nipple) are called Montgomery glands. They secrete oil to cleanse the nipple opening and prevent chapping and cracking of the nipple during breastfeeding.

The Process of Lactation

The pituitary hormone prolactin is instrumental in the establishment and maintenance of breast milk supply. It also is important for the mobilization of maternal micronutrients for breast milk.

Near the fifth week of pregnancy, the level of circulating prolactin begins to increase, eventually rising to approximately 10–20 times the pre-pregnancy concentration. We noted earlier that, during pregnancy, prolactin and other hormones prepare the breasts anatomically for the secretion of milk. The level of prolactin plateaus in late pregnancy, at a level high enough to initiate milk production. However, estrogen, progesterone, and other placental hormones inhibit prolactin-mediated milk synthesis during pregnancy. It is not until the placenta is expelled that this inhibition is lifted and milk production commences.

After childbirth, the baseline prolactin level drops sharply, but it is restored for a 1-hour spike during each feeding to stimulate the production of milk for the next feeding. With each prolactin spike, estrogen and progesterone also increase slightly.

When the infant suckles, sensory nerve fibers in the areola trigger a neuroendocrine reflex that results in milk secretion from lactocytes into the alveoli. The posterior pituitary releases oxytocin, which stimulates myoepithelial cells to squeeze milk from the alveoli so it can drain into the lactiferous ducts, collect in the lactiferous sinuses, and discharge through the nipple pores. It takes less than 1 minute from the time when an infant begins suckling (the latent period) until milk is secreted (the let-down). Figure 1 summarizes the positive feedback loop of the let-down reflex.

[caption id="" align="aligncenter" width="550"]This figure shows the process of let down reflex, the process in which the brain receives sensory impulses to release the hormones necessary for producing and discharging milk to the suckling newborn. Figure 1. Let-Down Reflex. A positive feedback loop ensures continued milk production as long as the infant continues to breastfeed.[/caption]

The prolactin-mediated synthesis of milk changes with time. Frequent milk removal by breastfeeding (or pumping) will maintain high circulating prolactin levels for several months. However, even with continued breastfeeding, baseline prolactin will decrease over time to its pre-pregnancy level. In addition to prolactin and oxytocin, growth hormone, cortisol, parathyroid hormone, and insulin contribute to lactation, in part by facilitating the transport of maternal amino acids, fatty acids, glucose, and calcium to breast milk.

Changes in the Composition of Breast Milk

In the final weeks of pregnancy, the alveoli swell with colostrum, a thick, yellowish substance that is high in protein but contains less fat and glucose than mature breast milk (Table 3). Before childbirth, some women experience leakage of colostrum from the nipples. In contrast, mature breast milk does not leak during pregnancy and is not secreted until several days after childbirth.

*Cow’s milk should never be given to an infant. Its composition is not suitable and its proteins are difficult for the infant to digest.
Compositions of Human Colostrum, Mature Breast Milk, and Cow’s Milk (g/L) (Table 3)
Human colostrum Human breast milk Cow’s milk*
Total protein 23 11 31
Immunoglobulins 19 0.1 1
Fat 30 45 38
Lactose 57 71 47
Calcium 0.5 0.3 1.4
Phosphorus 0.16 0.14 0.90
Sodium 0.50 0.15 0.41

Colostrum is secreted during the first 48–72 hours postpartum. Only a small volume of colostrum is produced—approximately 3 ounces in a 24-hour period—but it is sufficient for the newborn in the first few days of life. Colostrum is rich with immunoglobulins, which confer gastrointestinal, and also likely systemic, immunity as the newborn adjusts to a nonsterile environment.

After about the third postpartum day, the mother secretes transitional milk that represents an intermediate between mature milk and colostrum. This is followed by mature milk from approximately postpartum day 10 (see Table 3). As you can see in the accompanying table, cow’s milk is not a substitute for breast milk. It contains less lactose, less fat, and more protein and minerals. Moreover, the proteins in cow’s milk are difficult for an infant’s immature digestive system to metabolize and absorb.

The first few weeks of breastfeeding may involve leakage, soreness, and periods of milk engorgement as the relationship between milk supply and infant demand becomes established. Once this period is complete, the mother will produce approximately 1.5 liters of milk per day for a single infant, and more if she has twins or triplets. As the infant goes through growth spurts, the milk supply constantly adjusts to accommodate changes in demand. A woman can continue to lactate for years, but once breastfeeding is stopped for approximately 1 week, any remaining milk will be reabsorbed; in most cases, no more will be produced, even if suckling or pumping is resumed.

Mature milk changes from the beginning to the end of a feeding. The early milk, called foremilk, is watery, translucent, and rich in lactose and protein. Its purpose is to quench the infant’s thirst. Hindmilk is delivered toward the end of a feeding. It is opaque, creamy, and rich in fat, and serves to satisfy the infant’s appetite.

During the first days of a newborn’s life, it is important for meconium to be cleared from the intestines and for bilirubin to be kept low in the circulation. Recall that bilirubin, a product of erythrocyte breakdown, is processed by the liver and secreted in bile. It enters the gastrointestinal tract and exits the body in the stool. Breast milk has laxative properties that help expel meconium from the intestines and clear bilirubin through the excretion of bile. A high concentration of bilirubin in the blood causes jaundice. Some degree of jaundice is normal in newborns, but a high level of bilirubin—which is neurotoxic—can cause brain damage. Newborns, who do not yet have a fully functional blood–brain barrier, are highly vulnerable to the bilirubin circulating in the blood. Indeed, hyperbilirubinemia, a high level of circulating bilirubin, is the most common condition requiring medical attention in newborns. Newborns with hyperbilirubinemia are treated with phototherapy because UV light helps to break down the bilirubin quickly.
]]> 2816 2785 6 0 1103 Chapter 5. The Integumentary System https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-5-the-integumentary-system-2/ Thu, 13 Jul 2017 22:34:53 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-5-the-integumentary-system-2/ 2005 0 5 0 1103 Chapter 6. Bone Tissue and the Skeletal System https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-6-bone-tissue-and-the-skeletal-system-2/ Thu, 13 Jul 2017 22:35:09 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-6-bone-tissue-and-the-skeletal-system-2/ 2025 0 6 0 1103 Chapter 7. Axial Skeleton https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-7-axial-skeleton-2/ Thu, 13 Jul 2017 22:35:32 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-7-axial-skeleton-2/ 2052 0 7 0 1103 Chapter 8. The Appendicular Skeleton https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-8-the-appendicular-skeleton-2/ Thu, 13 Jul 2017 22:36:02 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-8-the-appendicular-skeleton-2/ 2082 0 8 0 1103 Chapter 9. Joints https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-9-joints-2/ Thu, 13 Jul 2017 22:36:21 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-9-joints-2/ 2108 0 9 0 1103 Chapter 10. Muscle Tissue https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-10-muscle-tissue-2/ Thu, 13 Jul 2017 22:36:51 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-10-muscle-tissue-2/ 2135 0 10 0 1103 Chapter 11. The Muscular System https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-11-the-muscular-system-2/ Thu, 13 Jul 2017 22:37:13 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-11-the-muscular-system-2/ 2162 0 11 0 1103 Chapter 18. The Cardiovascular System: Blood https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-18-the-cardiovascular-system-blood-2/ Thu, 13 Jul 2017 22:38:00 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-18-the-cardiovascular-system-blood-2/ 2194 0 12 0 1103 Chapter 19. The Cardiovascular System: The Heart https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-19-the-cardiovascular-system-the-heart-2/ Thu, 13 Jul 2017 22:38:37 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-19-the-cardiovascular-system-the-heart-2/ 2222 0 13 0 1103 Chapter 20. The Cardiovascular System: Blood Vessels and Circulation https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-20-the-cardiovascular-system-blood-vessels-and-circulation-2/ Thu, 13 Jul 2017 22:39:38 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-20-the-cardiovascular-system-blood-vessels-and-circulation-2/ 2261 0 14 0 1103 Chapter 21. The Lymphatic and Immune System https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-21-the-lymphatic-and-immune-system-2/ Thu, 13 Jul 2017 22:41:04 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-21-the-lymphatic-and-immune-system-2/ 2310 0 15 0 1103 Chapter 22. The Respiratory System https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-22-the-respiratory-system-2/ Thu, 13 Jul 2017 22:41:51 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-22-the-respiratory-system-2/ 2352 0 16 0 1103 Chapter 26. Fluid, Electrolyte, and Acid-Base Balance https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-26-fluid-electrolyte-and-acid-base-balance/ Thu, 13 Jul 2017 22:50:16 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-26-fluid-electrolyte-and-acid-base-balance/ 2395 0 18 0 1103 Chapter 24. Metabolism and Nutrition https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-24-metabolism-and-nutrition/ Thu, 13 Jul 2017 22:50:23 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-chapter-24-metabolism-and-nutrition/ 2405 0 17 0 1103 Review https://pressbooks.bccampus.ca/apdouglas2017/part/1103-review/ Thu, 13 Jul 2017 22:50:24 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1103-review/ 2408 0 19 0 1203 Chapter 1. An Introduction to the Human Body https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-1-an-introduction-to-the-human-body/ Thu, 13 Jul 2017 22:50:27 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-1-an-introduction-to-the-human-body/ 2420 0 20 0 1203 Chapter 2. The Chemical Level of Organization https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-2-the-chemical-level-of-organization/ Thu, 13 Jul 2017 22:50:32 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-2-the-chemical-level-of-organization/ 2429 0 21 0 1203 Chapter 3. The Cellular Level of Organization https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-3-the-cellular-level-of-organization/ Thu, 13 Jul 2017 22:51:19 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-3-the-cellular-level-of-organization/ 2462 0 22 0 1203 Chapter 4. The Tissue Level of Organization https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-4-the-tissue-level-of-organization/ Thu, 13 Jul 2017 22:51:21 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-4-the-tissue-level-of-organization/ 2467 0 23 0 1203 Chapter 12. The Nervous System and Nervous Tissue https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-12-the-nervous-system-and-nervous-tissue/ Thu, 13 Jul 2017 22:51:41 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-12-the-nervous-system-and-nervous-tissue/ 2477 0 24 0 1203 Chapter 13. The Central Nervous System https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-13-the-central-nervous-system/ Thu, 13 Jul 2017 22:52:09 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-13-the-central-nervous-system/ 2512 0 25 0 1203 Chapter 14. The Somatic Nervous System https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-14-the-somatic-nervous-system/ Thu, 13 Jul 2017 22:52:31 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-14-the-somatic-nervous-system/ 2535 0 26 0 1203 Chapter 15. The Autonomic Nervous System https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-15-the-autonomic-nervous-system/ Thu, 13 Jul 2017 22:53:19 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-15-the-autonomic-nervous-system/ 2569 0 27 0 1203 Chapter 17. The Endocrine System https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-17-the-endocrine-system/ Thu, 13 Jul 2017 22:53:42 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-17-the-endocrine-system/ 2585 0 28 0 1203 Chapter 20. The Cardiovascular System: Blood Vessels and Circulation https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-20-the-cardiovascular-system-blood-vessels-and-circulation/ Thu, 13 Jul 2017 22:54:04 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-20-the-cardiovascular-system-blood-vessels-and-circulation/ 2615 0 29 0 1203 Chapter 22. The Respiratory System https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-22-the-respiratory-system/ Thu, 13 Jul 2017 22:54:07 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-22-the-respiratory-system/ 2618 0 30 0 1203 Chapter 24. Metabolism and Nutrition https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-24-metabolism-and-nutrition/ Thu, 13 Jul 2017 22:54:22 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-24-metabolism-and-nutrition/ 2634 0 32 0 1203 Chapter 23. The Digestive System https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-23-the-digestive-system/ Thu, 13 Jul 2017 22:55:08 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-23-the-digestive-system/ 2664 0 31 0 1203 Chapter 25. The Urinary System https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-25-the-urinary-system/ Thu, 13 Jul 2017 22:55:45 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-25-the-urinary-system/ 2711 0 33 0 1203 Chapter 26. Fluid, Electrolyte, and Acid-Base Balance https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-26-fluid-electrolyte-and-acid-base-balance/ Thu, 13 Jul 2017 22:56:16 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-26-fluid-electrolyte-and-acid-base-balance/ 2738 0 34 0 1203 Chapter 27. The Reproductive System https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-27-the-reproductive-system/ Thu, 13 Jul 2017 22:56:39 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-27-the-reproductive-system/ 2763 0 35 0 1203 Chapter 28. Development and Inheritance https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-28-development-and-inheritance/ Thu, 13 Jul 2017 22:56:55 +0000 https://pressbooks.bccampus.ca/apdouglas2017/part/1203-chapter-28-development-and-inheritance/ 2785 0 36 0