Human Physiology

Hepatic portal system In human anatomy, the hepatic portal system is the system of veins comprised of the hepatic portal vein and its tributaries. It is also called the portal venous system, although it is not the only example of a portal venous system, and splanchnic veins, which is not synonymous with hepatic portal system and is imprecise (as it means visceral veins and not necessarily the veins of the abdominal viscera).

Function
The portal venous system is responsible for directing blood from parts of the gastrointestinal tract to the liver. Substances absorbed in the small intestine travel first to the liver for processing before continuing to the heart. Not all of the gastrointestinal tract is part of this system. The system extends from about the lower portion of the esophagus to the upper part of the anal canal. It also includes venous drainage from the spleen and pancreas. Many drugs that are absorbed through the GI tract are substantially metabolized by the liver before reaching general circulation. This is known as the first pass effect. As a consequence, certain drugs can only be taken via certain routes. For example, nitroglycerin cannot be swallowed because the liver would inactivate the medication, but it can be taken under the tongue or transdermal (through the skin) and thus is absorbed in a way that bypasses the portal venous system. Blood flow to the liver is unique in that it receives both oxygenated and deoxygenated blood. As a result, the partial pressure of oxygen (pO2) and perfusion pressure of portal blood are lower than in other organs of the body. Blood passes from branches of the portal vein through cavities between "plates" of hepatocytes called sinusoids. Blood also flows from branches of the hepatic artery and mixes in the sinusoids to supply the hepatocytes with oxygen. This mixture percolates through the sinusoids and collects in a central vein which drains into the hepatic vein. The hepatic vein subsequently drains into the inferior vena cava. Large veins that are considered part of the portal venous system are the: Hepatic portal vein Splenic vein Roughly, the portal venous system corresponds to areas supplied by the celiac trunk, the superior mesenteric artery, and the inferior mesenteric artery.

The hepatic portal system begins in the capillaries of the digestive organs and ends in the portal vein. Consequently, portal blood contains substances absorbed by the stomach and intestines. Portal blood is passed through the hepatic lobules where nutrients and toxins are absorbed, excreted or converted. Restriction of outflow through the hepatic portal system can lead to portal hypertension. Portal hypertension is most often associated with cirrhosis. Patients usually present with splenomegaly, ascites, GI bleeding and/or portal systemic encephalopathy. The consequences of portal hypertension are due to portal systemic anastomosis formed by the body as an attempt to bypass the obstructed liver circulation. These collateral vessels form along the falciform ligament, diaphragm, spleen, stomach and peritoneum. The collaterals find their way to the renal vein where blood drained from the digestive organs is let into the systemic circulation.

Blood Components

Normally, 7-8% of human body weight is from blood. In adults, this amounts

to 4-5 quarts of blood. This essential fluid carries out the critical functions of transporting oxygen and nutrients to our cells and getting rid of carbon dioxide, ammonia, and other waste products. In addition, it plays a vital role in our immune system and in maintaining a relatively constant body temperature. Blood is a highly specialized tissue composed of many different kinds of components. Four of the most important ones are red cells, white cells, platelets, and plasma. All humans produce these blood components-there are no populational or regional differences.

Red Cells
Red cells, or erythrocytes , are relatively large microscopic cells without nuclei. In this latter trait, they are similar to the primitive prokaryotic cells of bacteria. Red cells normally make up 40-50% of the total blood volume. They transport oxygen from the lungs to all of the living tissues of the body and carry away carbon dioxide. The red cells are produced continuously in our bone marrow from stem cellsat a rate of about 2-3 million cells per second. Hemoglobin is the gas transporting protein molecule that makes up 95% of a red cell. Each red cell has about 270,000,000 iron-rich hemoglobin molecules. People who are anemic generally have a deficiency in red cells. The red color of blood is primarily due to oxygenated red cells. Human fetal hemoglobin molecules differ from those produced by adults in the number of amino acid chains. Fetal hemoglobin has three chains, while adults produce only two. As a consequence, fetal hemoglobin molecules attract and transport relatively more oxygen to

the cells of the body.

White Cells
White cells, or leukocytes , exist in variable numbers and types but make up a very small part of blood's volume-normally only about 1% in healthy people. Leukocytes are not limited to blood. They occur elsewhere in the body as well, most notably in the spleen, liver, and lymph glands. Most are produced in our bone marrow from the same kind of stem cells that produce red blood cells. Others are produced in the thymus gland, which is at the base of the neck. Some white cells (called lymphocytes ) are the first responders for our immune system. They seek out, identify, and bind to alien protein on bacteria, viruses, and fungi so that they can be removed. Other white cells (called granulocytes and macrophages ) then arrive to surround and destroy the alien cells. They also have the function of getting rid of dead or dying blood cells as well as foreign matter such as dust and asbestos. Red cells remain viable for only about 4 months before they are removed from the blood and their components recycled in the spleen. Individual white cells usually only last 18-36 hours before they also are removed, though some types live as much as a year. The description of white cells presented here is a simplification. There are actually many specialized sub-types of them that participate in different ways in our immune responses.

Platelets
Platelets , or thrombocytes , are cell fragments without nuclei that work with blood clotting chemicals at the site of wounds. They do this by adhering to the walls of blood vessels, thereby plugging the rupture in thevascular wall. They also can release coagulating chemicals which cause clots to form in the blood that can plug up narrowed blood vessels. There are more than a dozen types of blood clotting factors and platelets that need to interact in the blood clotting process. Recent research has shown that platelets help fight infections by releasing proteins that kill invading bacteria and some other microorganisms. In addition, platelets stimulate the immune system. Individual platelets are about 1/3 the size of red cells. They have a lifespan of 9-10 days. Like the red and white blood cells, platelets are produced in bone marrow from stem cells.

Plasma

Plasma is the relatively clear liquid water (92+%), sugar, fat, protein and salt solution which carries the red cells, white cells, platelets, and some other chemicals. Normally, 55% of our blood's volume is made up of plasma. About 95% of it consists of water. As the heart pumps blood to cells throughout the body, plasma brings nourishment to them and removes the waste products ofmetabolism. Plasma also contains blood clotting factors, sugars, lipids, vitamins, minerals, hormones, enzymes, antibodies, and other proteins. It is likely that plasma contains some of every protein produced by the body--approximately 500 have been identified in human plasma so far.

Kidney Filteration How does the mammalian kidney produce urine? The kidney contains numerous functional units, called nephrons, which produce urine through a series of steps: glomerular filtration of the blood, tubular reabsorption of the glomerular filtrate, and tubular secretion of harmful substances.

Glomerular Filtration

Blood flows from the afferent arteriole into the glomerulus, a tuft of fenestrated capillaries enclosed in the Bowmans capsule. Here 15 to 25 percent of the plasmas water and solutes are filtered through a single cell layer of the capillary walls, through a basement membrane, and into the lumen of the Bowmans capsule. The filtrate then flows into the renal tubule, to undergo tubular reabsorption.

The rate of glomerular filtration depends on three factors: the hydrostatic pressure difference between the capillaries and the Bowmans capsule (due to blood pressure), the colloid osmotic pressure, which opposes filtration, and the hydraulic permeability of the three-layered tissue separating the capillaries and the lumen of the Bowmans capsule. Overall, blood pressure in the body has a major effect on the glomerular filtration rate, because the amount of blood passing through the glomerulus determines how much and how fast the fluid can be filtered. Among the dangers of very low blood pressure, therefore, is the loss of kidney function. This is a primary reason for the use of inflatable "shock suits" on the lower body in cases of extreme blood loss from trauma. By reducing blood flow to the legs, blood pressure in

the trunk is kept higher, in an effort to maintain kidney function.

The body through endocrine responses can regulate the glomerular filtration rate. In the case of auto regulation, increased blood pressure stretches walls of the afferent arteriole, which responds by contracting - thereby reducing fluctuation of blood pressure in the glomerulus. A drop in blood pressure brings about a decrease in the glomerular filtration rate, which, in turn, results in a decrease in sodium ions in the filtrate. (The filtrate moves through the nephrons more slowly, allowing more sodium to be reabsorbed along the way.). This lower sodium level in the filtrate is detected by the macula densa, modified cells of the wall of the distal convoluted tubule that lie adjacent to the afferent and efferent arterioles of the glomerulus. In response to the low sodium, cells of the juxtaglomerular apparatus (JGA) release rennin. This triggers a series of biochemical reactions which bring about an increase of blood pressure, and thereby an increase in GFR. This series of reactions includes an increase in angiotensin II, which helps bring blood pressure back up by (1) causing vasoconstriction in arterioles throughout much of the body, and (2) promoting increased synthesis of antidiiuretic hormone (ADH), which increases resorption of water in the collecting ducts of the kidney, thereby increasing blood volume. Angiotensin II also promotes the release of aldosterone from adrenal cortex, which promotes retention of both sodium and water, thereby helping to bring blood pressure back up. The glomerular filtration rate can also be regulated by sympathetic nerve responses, which can result in the constriction or dilation of the afferent arterioles in times of great bodily stress. Vasoconstriction increases blood pressure, and therefore GFR, whereas vasodilation decreases blood pressure and GFR.

Tubular Reabsorption

As the glomerular filtrate moves through the nephron, it changes composition dramatically. About 99% of the water in the original filtrate is reabsorbed, and less than 1% of the original NaCl content appears in the final urine. How does this happen? First, the filtrate moves into the proximal tubule, where about 70% of the sodium ions are reabsorbed through active transport. Water and chloride ions follow passively. Glucose and amino acids are reabsorbed here. Approximately 75% of the glomerular filtrate is reabsorbed in the proximal tubule. This is aided by the so-called "brush border" of microvilli cells which function in increasing the surface area for reabsorption. After moving through the proximal tubule, the filtrate moves on to the descending limb of the Loop of Henle. The cells in this area have no brush border, and there is no active salt transport here. The cells have a low permeability to urea and salt, but are very permeable to water. As the filtrate descends the Loop of Henle, water diffuses out because of the high salt concentration in the surrounding tissue. This is part of the urine-concentrating system of the nephron. The Loop of Henle acts as a countercurrent multiplier. For this reason, mammals living in marine or desert environments have longer loops of Henle and can conserve more water by producing more concentrated urine. The filtrate moves along the Loop of Henle to the thin segment of the ascending limb, which is highly permeable to Na+ and Cl-, and impermeable to water and urea. As the filtrate moves up the thin segment, Na+ and Cl- diffuse out because there is a higher concentration in the filtrate than in the surrounding tissues. The filtrate then moves to the medullary thick ascending limb, which is involved in the active transport of Na+ and Cl- outward from the lumen into the interstitial space. This causes the fluid reaching the distal tubule to be hypoosmotic to the interstitial fluid, and allows for the passive transport of water out

of the tubule. The distal tubule functions in transporting K+, H+, and NH3 into the lumen, and Na+, Cl-, and HCO3- out. Transport of salts in this area is under endocrine control and adjusted according to osmotic conditions. The filtrate then moves into the collecting duct, which carries the fluid to the renal pelvis, to the ureters, and out of the body through the urethra. The epithelium of the collecting duct is permeable to water, but not salt or urea. The hormone ADH from the posterior pituitary gland controls the permeability o the collecting duct to water. This response causes the filtrate, which is hypoosmotic to the interstitial fluid at this point, to lose water by osmosis and therefore increase the concentration of salts and urea in the urine. At the bottom of the collecting duct, the epithelium is permeable to urea. The diffusion of some urea out of the filtrate and into the surrounding tissue helps produce the interstitial concentration gradient necessary for the diffusion of water out of the descending limb of the Loop of Henle. The urea also helps draw water out of the filtrate passing down the collecting duct, thereby enabling the kidney to excrete urine that is hypertonic to the general body fluids, a property that is important in water conservation.

Tubular Secretion

In several places along the nephron, substances that are not part of the initial filtrate (because the molecules are too big to be filtered through the glomerulus) are actively transported from the blood into the filtrate for elimination from the body in the urine. Some substances, such as toxins and drugs, are processed in the liver and conjugated with glucouronic acid. This marks them for removal from blood capillaries in the kidney and transport into the lumen of the nephron to become part of the filtrate.

How do the kidneys regulate pH?

Regulation of pH is governed by the carbon dioxide/bicarbonate buffering system in the body, which consists of three steps: CO2 + H2O <==> H2CO3 <==> HCO3- + H+ CO2 + OH- + H+ <==> HCO3- + H+ HOH <==> OH- + H+ The excretion of acid by the kidney is one of the two major factors, which influence this system (the other being the excretion of carbon dioxide by the lungs). The excretion of hydrogen ions (acid) in the urine is primarily responsible for maintaining the plasma HCO3- concentration. Mammalian urine is mildly acidic, with a pH of about 6, and contains no bicarbonate. However, the initial glomerular filtrate has a high bicarbonate concentration and a low hydrogen ion concentration. Therefore, in the process of urine formation, acid must be added to the filtrate, and bicarbonate must be removed. Therefore, the excretion of H+ and the recovery of HCO3- are both important mechanisms by which the kidneys help the body regulate pH. Special cells in the distal tubule and collecting duct, called A-type cells and B-type cells, accomplish this

process. The A-type cells are acid-secreting cells that have a proton ATPase in the apical membrane and a Cl-/ HCO3- exchange system in the basolateral membrane. The cells also contain carbonic anhydrase, which hydrates carbon dioxide passing through the membrane to form protons and bicarbonate ions. The protons formed are pumped back into the lumen and can react with the bicarbonate in the filtrate to form carbon dioxide and water, which can diffuse back into the cell, and create an uptake of bicarbonate back into the blood. B-type cells, on the other hand, are base-secreting cells. They have a different form of chloride/bicarbonate exchanger in the apical membrane than the A-type cells, and secrete bicarbonate into the lumen of the tubule in exchange for chloride ions. Regulation of pH is accomplished then, by altering the activity of A and B-type cells, which determines whether bicarbonate is reabsorbed or secreted. Another mechanism used in pH regulation is the uptake of H+ by HPO4- and NH3 in the lumen to trap excess H+ in the filtrate. This occurs in order to bind H+ with something so that these protons will not move back into the epithelial cells and the blood, which would lower pH. Excretion and osmoregulation

The major problems that animals face with regard to osmoregulation

In most animals, the majority of cells are not in direct contact with the external environment but are bathed by an internal body fluid. Homeostatic mechanisms hamper changes in an animals body fluid, which both gives protection from harmful external environments and impedes quick exchange between intracellular compartments. The cells of the animal cannot survive much additional water gain or loss. Water continuously enters and leaves an animal cell across the plasma membrane, however, uptake and loss must balance. Animal cells swell and burst if there is a net uptake of water or shrivel and die it there is a net loss of water. Other problems associated with osmoregulation are body and environment temperatures. The enzyme activity in the body function between temperatures of 0o 40o Celsius. The way animals deal with temperature and regulating it is by way of water loss. So animals in hot environments need to limit the amount of water loss due to evaporation and respiration. The importance of water in temperature regulating leads to conflicts and compromises between physiological adaptations to environmental temperatures and osmotic stresses in terrestrial animals.

The major structures involved in osmoregulation and excretion

Organisms in different environments utilize different structures in osmoregulation and excretion. The major structures involved are the integument, the respiratory surface, the kidney, and the salt gland. All animals use at least one of these structures in their osmoregulatory processes. The common characteristic in structures such as gills, skin, kidneys and the integument are cells called transport epithelia, anatomically and functionally polarized cells which determine the osmoregulatory capabilities of the structure, through properties such as permeability to various solutes. The integumentfunctions in osmoregulation by acting as a barrier between the extracellular compartment and the environment to regulate water gain and loss, as well as solute flux. The permeablity of the integument to water and solutes varies from animal to animal.

Respiratory surfaces such as the alveoli of the lung, and gills in aquatic animals also serve in osmoregulation and excretion. Respiratory surfaces are the chief avenues for the excretion of carbon dioxide and metabolic water, as well as other gaseous wastes, in animals. The kidney is the main organ involved in maintaining water balance and excreting harmful substances in mammals. Elasmobranchs, marine birds, and some reptiles have a structure called a salt gland to secrete NaCl from their bodies. These animals require a lower internal NaCl concentration than the surrounding seawater, which causes a concentration gradient favoring the influx of salt. Therefore, they need a way to secrete it. The solution is provided by glands in the rectum of sharks and the skulls of marine birds and reptiles which produces a concentrated salt solution for secretion. The sodium ions are removed from the blood by these glands not by filtration, but by the sodium-transport mechanism (the sodium-potassium pump). This Na+/ K+ / ATPase activity allows for the movement of NaCl from the blood across the epithelium into the lumen of the salt gland for secretion. Interestingly, the shark rectal gland, bird nasal gland, fish gill, and the thick ascending Loop of Henle in the kidney all contain salt-secreting cells that transport NaCl by the same basic mechanism. Active transport produces an increase in the chloride concentration in the cytoplasm of epithelial cells. This results in the diffusion of chloride ions out of the cell across the apical surface. The build-up of chloride ions at the apical surface attracts sodium ions to diffuse between the cells (the paracellular route). Insects have a network of Malpighian tubules extending throughout much of the body cavity and attached to the alimentary canal between the midgut and the hindgut. The secretory cells which line the walls of these long, thin tubules secrete KCl, NaCl, and phosphate from the hemolymph (blood) into the lumen of the tubule. Smaller molecules, such as water, amino acids, and sugars diffuse down their concentration gradient and into the lumen. The fluid then flows along the tubule and into the gut. As the fluid passes through the hindgut, water and valuable ions are transported back into the hemolymph, leaving behind a concentrated waste for excretion from the body.

Why and how do organisms excrete metabolic wastes (particularly nitrogenous wastes)?

Waste products generated in metabolic processes are often toxic, and therefore must be eliminated before they can harm the organism. The major metabolic wastes produced by animals include carbon dioxide, metabolic water, and nitrogenous wastes. Small aquatic organisms are able to get rid of wastes by simple diffusion across membranes. More complex animals with circulatory systems rely on kidneys to filter wastes out of the blood and eliminate them from the body.

Carbon dioxide and metabolic water produced in respiration easily diffuse into the environment from respiratory surfaces. Nitrogenous waste excretion is more difficult, yet necessary. Elevated ammonia levels in the body can lead to convulsions, coma, and even death. This is because ammonium ions can substitute for potassium ions in ion-exchange mechanisms. Ammonia can also adversely affect metabolism and amino acid transport. An excessive amount of ammonia in the system elevates bodily pH, which causes changes in the tertiary structure of proteins, and thus cellular functions can be altered. There are three main types of nitrogenous wastes: ammonia, urea, and uric acid. The type of waste an animal excretes depends on its living environment, because nitrogenous waste excretion is accompanied by a certain amount of water loss. Ammonotelic (ammonia-excreting) animals generally live only in aquatic habitats, because ammonia is extremely toxic, and a large volume of water is required to maintain the excreted ammonia level lower than the body level. This is needed because ammonia excretion relies on passive diffusion, so a gradient is required between the organism and the environment in order for the ammonia to flow from high concentration to low concentration. Whereas most excretion of ammonia occurs across the gills of aquatic animals, mammals do excrete some ammonia in the urine. Amino groups are enzymatically transformed into glutamate, and then changed to glutamine in the liver. Glutamine can cross the kidney membranes (whereas amino acids can

not). In the kidney tubules, the glutamine is deaminated to ammonia and then excreted in the urine.

Although ammonia excretion is present in some forms in mammals, the major nitrogenous waste excreted is urea. Urea is less toxic than ammonia, and requires less water for elimination. Therefore, ureotelic (urea-excreting) animals are most often (but not exclusively) terrestrial. A downside to urea excretion is that urea synthesis requires energy, in the form of ATP. Vertebrates synthesize urea in the liver using the ornithine-urea cycle. Teleosts and invertebrates produce urea from uric acid via the uricolytic pathway.

Birds, reptiles, and most terrestrial arthropods often are subject to very limited water availability, so even urea excretion is not possible. Therefore, these uricotelic (uric acid-excreting) animals synthesize uric acid, which requires even less water than urea for elimination. The ability to produce uric acid, which is relatively insoluble, is quite important to birds and reptiles prior to hatching. Nitrogenous wastes can be safely stored within the egg in the form of uric acid, whereas a build-up of either ammonia or urea would be deadly.

Human Vertebral System

In vertebrate animals, the flexible column extending from neck to tail, made of a series of bones, the vertebrae. The major function of the vertebral column is protection of the spinal cord; it also provides stiffening for the body and attachment for the pectoral and pelvic girdles and many muscles. In humans an additional function is to transmit body weight in walking and standing. Each vertebra, in higher vertebrates, consists of a ventral body, or centrum, surmounted by a Yshaped neural arch. The arch extends a spinous process (projection) downward and backward that may be felt as a series of bumps down the back, and two transverse processes, one to either side, which provide attachment for muscles and ligaments. Together the centrum and neural arch surround an opening, the vertebral foramen, through which the spinal cord passes. The centrums are separated by cartilaginous intervertebral disks, which help cushion shock in locomotion. Vertebrae in lower vertebrates are more complex, and the relationships of their parts to those of higher animals are often unclear. In primitive chordates (e.g., amphioxus, lampreys) a rodlike structure, the notochord, stiffens the body and helps protect the overlying spinal cord. The notochord appears in the embryos of all vertebrates in the space later occupied by the vertebral bodiesin some fish it remains throughout life, surrounded by spool-shaped centrums; in other vertebrates it is lost in the developed animal. In primitive chordates the spinal cord is protected dorsally by segmented cartilagesthese foreshadow the development of the neural arch of true vertebrae. Fish have trunk and caudal (tail) vertebrae; in land vertebrates with legs, the vertebral column becomes further subdivided into regions in which the vertebrae have different shapes and functions. Crocodilians and lizards, birds, and mammals demonstrate five regions: (1) cervical, in the neck, (2) thoracic, in the chest, which articulates with the ribs, (3) lumbar, in the lower back, more robust than the other vertebrae, (4) sacral, often fused to form a sacrum, which articulates with the pelvic girdle, (5) caudal, in the tail. The atlas and axis vertebrae, the top two cervicals, form a freely movable joint with the skull. The numbers of vertebrae in each region and in total vary with the species. Snakes have the greatest number, all very similar in type. In turtles some vertebrae may be fused to the shell (carapace); in birds all but the cervical vertebrae are usually fused into a rigid structure, which

lends support in flight. Most mammals have seven cervical vertebrae; size rather than number account for the variations in neck length in different species. Whales show several specializationsthe cervical vertebrae may be either much reduced or much increased in number, and the sacrum is missing. Humans have 7 cervical, 12 thoracic, 5 lumbar, 5 fused sacral, and 3 to 5 fused caudal vertebrae (together called the coccyx). The vertebral column is characterized by a variable number of curves. In quadrupeds the column is curved in a single arc (the highest portion occurring at the middle of the back), which functions somewhat like a bow spring in locomotion. In humans this primary curve is modified by three more: (1) a sacral curve, in which the sacrum curves backward and helps support the abdominal organs, (2) an anterior cervical curve, which develops soon after birth as the head is raised, and (3) a lumbar curve, also anterior, which develops as the child sits and walks. The lumbar curve is a permanent characteristic only of humans and their bipedal forebears, though a temporary lumbar curve appears in other primates in the sitting position. The cervical curve disappears in humans when the head is bent forward but appears in other animals as the head is raised. Human skeletal system The vertebral column is not actually a column but rather a sort of spiral spring in the form of the letter S. The newborn child has a relatively straight backbone. The development of the curvatures occurs as the supporting functions of the vertebral column in humansi.e., holding up the trunk, keeping the head erect, serving as an anchor for the extremitiesare developed. The S-curvature enables the vertebral column to absorb the shocks of walking on hard surfaces; a straight column would conduct the jarring shocks directly from the pelvic girdle to the head. The curvature meets the problem of the weight of the viscera. In an erect animal with a straight column, the column would be pulled forward by the viscera. Additional space for the viscera is provided by the concavities of the thoracic and pelvic regions. Weight distribution of the entire body is also effected by the S-curvature. The upper sector to a large extent carries the head; the central sector carries the thoracic viscera, the organs and structures in the chest; and the lower sector carries the abdominal viscera. If the column were straight, the weight load would increase from the head downward and be relatively great at the base. Lastly, the S-curvature protects the vertebral column from breakage. The doubly bent spring arrangement is far less vulnerable to fracture than would be a straight column. The protective function of the skeleton is perhaps most conspicuous in relation to the central nervous system, although it is equally important for the heart and lungs and some other organs. A high degree of protection for the nervous system is made possible by the relatively small amount of motion and expansion needed by the component parts of this system and by certain physiological adaptations relating to circulation, to thecerebrospinal fluid, and to the meninges, the coverings of the brain and spinal cord. The brain itself is snugly enclosed within the boxlike cranium. Sharing in the protection afforded by the cranium is the pituitary gland, or hypophysis. In human anatomy, the vertebral column (backbone or spine) is a column usually consisting of 24 vertebrae,[1] the sacrum, intervertebral discs, and the coccyx situated in the dorsal aspect of the torso, separated by spinal discs. It houses the spinal cord in its spinal canal.
Curves

Viewed laterally the vertebral column presents several curves, which correspond to the different regions of the column, and are called cervical, thoracic, lumbar, and pelvic. The cervical curve, convex forward, begins at the apex of the odontoid (tooth-like) process, and ends at the middle of the second thoracic vertebra; it is the least marked of all the curves. The thoracic curve, concave forward, begins at the middle of the second and ends at the middle of the twelfth thoracic vertebra. Its most prominent point behind corresponds to the spinous process of the seventh thoracic vertebra. This curve is known as a tt curve. The lumbar curve is more marked in the female than in the male; it begins at the middle of the last thoracic vertebra, and ends at the sacrovertebral angle. It is convex anteriorly, the convexity of the lower three vertebrae being much greater than that of the upper two. This curve is described as a lordotic curve. The pelvic curve begins at the sacrovertebral articulation, and ends at the point of the coccyx; its concavity is directed downward and forward. The thoracic and pelvic curves are termed primary curves, because they alone are present during fetal life. The cervical and lumbar curves are compensatory or secondary, and are developed after birth, the former when the child is able to hold up its head (at three or four months) and to sit upright (at nine months), the latter at twelve or eighteen months, when the child begins to walk.
Names of individual vertebrae

Individual vertebrae named according to region and position, from superior to inferior

Cervical 7 vertebrae (C1-C7) o C1 is known as "atlas" and supports the head, C2 is known as "axis" o Possesses bifid spinous processes, which is absent in C7 o Small-bodied Thoracic 12 vertebrae (T1-T12) o Distinguished by the presence of costal facets for the articulation of the heads of ribs o Body is intermediate in size between the cervical and lumbar vertebrae Lumbar 5 vertebrae (L1-L5) o Has a large body o Does not have costal facets nor transverse process foramina Sacral 5 (fused) vertebrae (S1-S5) Coccygeal 4 (3-5) (fused) vertebrae (Tailbone)

Gaseous Exchange Breathing consists of two phases, inspiration and expiration. During inspiration, the diaphragm and the intercostal muscles contract. The diaphragm moves downwards increasing the volume of the thoracic (chest) cavity, and the intercostal muscles pull the ribs up expanding the rib cage and further increasing this volume. This increase of volume lowers the air pressure in the alveoli to below atmospheric pressure. Because air always flows from a region of high pressure to a region of lower pressure, it rushes in through the respiratory tract and into the alveoli. This is called negative pressure breathing, changing the pressure inside the lungs relative to the pressure of the outside atmosphere. In contrast to inspiration, during expiration the diaphragm and intercostal muscles relax. This returns the thoracic cavity to it's original volume, increasing the air pressure in the lungs, and

forcing the air out. External Respiration When a breath is taken, air passes in through the nostrils, through the nasal passages, into the pharynx, through the larynx, down the trachea, into one of the main bronchi, then into smaller broncial tubules, through even smaller bronchioles, and into a microscopic air sac called an alveolus. It is here that external respiration occurs. Simply put, it is the exchange of oxygen and carbon dioxide between the air and the blood in the lungs. Blood enters the lungs via the pulmonary arteries. It then proceeds through arterioles and into the alveolar capillaries. Oxygen and carbon dioxide are exchanged between blood and the air. This blood then flows out of the alveolar capillaries, through vaneuoles, and back to the heart via the pulmonary veins. For an explanation as to why gasses are exchanged here, see partial pressure. Gas Transport If 100mL of plasma is exposed to an atmosphere with a pO2 of 100mm Hg, only 0.3mL of oxygen would be absorbed. However, if 100mL of blood is exposed to the same atmosphere, about 19mL of oxygen would be absorbed. This is due to the presence of hemoglobin, the main means of oxygen transport in the body. The respiratory pigment hemoglobin is made up of an iron-containing porphyron, haem, combined with the protein globin. Each iron atom in haem is attached to four pyrole groups by covalent bonds. A fifth covalent bond of the iron is attached to the globin part of the molecule and the sixth covalent bond is available for combination with oxygen. There are four iron atoms in each heamoglobin molecule and therefore four heam groups. Oxygen Transport In the loading and unloading of oxygen, there is a cooperation between these four haem groups. When oxygen binds to one of the groups, the others change shape slighty and their attraction to oxygen increases. The loading of the first oxygen, results in the rapid loading of the next three (forming oxyhaemoglobin). At the other end, when one group unloads it's oxygen, the other three rapidly unload as their groups change shape again having less attraction for oxygen. This method of cooperative binding and release can be seen in the dissociation curve for hemoglobin. Over the range of oxygen concentrations where the curve has a steep slope, the slightest change in concentration will cause hemoglobin to load or unload a substantial amount of oxygen. Notice that the steep part of the curve corresponds to the range of oxygen concentrations found in the tissues. When the cells in a particular location begin to work harder, e.g. during exercise, oxygen concentration dips in that location, as the oxygen is used in cellular respiration. Because of the cooperation between the haem groups, this slight change in concentration is enough to cause a large increase in the amount of oxygen unloaded. As with all proteins, hemoglobins shape shift is sensitive to a variety of environmental conditions. A drop in pH lowers the attraction of hemoglobin to oxygen, an effect known as the Bohr shift. Because carbon dioxide reacts with water to produce carbonic acid, an active tissue will lower the pH of it's surroundings and encourage hemoglobin to give up extra oxygen, to be used in cellular respiration. Hemoglobin a notable molecule for it's ability to transport oxygens from regions of supply to regions of demand. Carbon Dioxide Transport - Out of the carbon dioxide released from respiring cells, 7% dissolves into the plasma, 23% binds to the multiple amino groups of hemoglobin (Caroxyhaemoglobin), and 70% is carried as bicarbonate ions. Carbon dioxide created by

respiring cells diffuses into the blood plasma and then into the red blood cells, where most of it is converted to bicarbonate ions. It first reacts with water forming carbonic acid, which then breaks down into H+ and CO3-. Most of the hydrogen ions that are produced attach to hemoglobin or other proteins. In this Internal Respiration The body tissues need the oxygen and have to get rid of the carbon dioxide, so the blood carried throughout the body exchanges oxygen and carbon dioxide with the body's tissues. Internal respiration is basically the exchange of gasses between the blood in the capillaries and the body's cells. Additional readings. Gaseous Exchange In The Body The exchange of Oxygen (O2) and Carbon Dioxide (CO2) between alveolar air and pulmonary blood occurs via passive diffusion. This is governed by the behavior of gases as described by Dalton's Law and Henry's Law. Gaseous exchange in the body occurs in two places: 1. Between the air in the alveoli of the lungs and the blood in pulmonary capillaries (External respiration). 2. Between the systemic capillaries and tissue cells (Internal respiration). External Respiration This process results in the conversion of deoxygenated blood from the right side of the heart to oxygenated blood returning to the left side of the heart. Gases are exchanged by diffusion according to the differences in their partial pressures. The now deoxygenated blood returns to the heart and is pumped to the lungs where the process of external respiration begins again. The partial pressure of O2 in alveolar air is 105mmHg, while the resting partial pressure of O2 in deoxygenated blood is ~40mmHg. Due to this difference O2 diffuses down its concentration gradient from the alveolar air into the deoxygenated blood until equilibrium is reached, the result being that the blood becomes oxygenated. The partial pressure of CO2 in alveolar air is 40mmHg, while in deoxygenated blood it is 45 mmHg. CO2 therefore diffuses in the opposite direction to O2, again down its concentration gradient. The result being that CO2 is removed from the blood and exhaled. The now deoxygenated blood returns to the heart and is pumped to the lungs where the process of external respiration begins again. The rate of gas exchange during external respiration depends on several factors: Partial pressure difference of the gases Surface area available for gas exchange Diffusion distance Solubility and molecular weight of the gases Internal Respiration The process of internal respiration is much the same as external respiration, only the site in which it occurs is different. Internal respiration results in the conversion of oxygenated blood (from the capillaries) to deoxygenated blood. Once again the gases are exchanged in accordance with their partial pressures. The partial pressure of O2 in capillary blood is ~100mmHg, whilst in the tissues it is ~ 40mmHg. Due to this difference O2 diffuses from the blood, through the interstitial fluid into the tissue cells until the partial pressure of O2 in the blood decreases to ~40mmHg.

CO2 again diffuses in the opposite direction. The partial pressure of CO2 in the tissue cells is 45mmHg, whilst in the blood it is 40mmHg. Therefore, CO2 diffuses from the tissue cells through the interstitial fluid into the blood until its partial pressure in the blood reaches ~45mmHg. The deoxygenated blood returns to the heart from where it is pumped to the lungs and the process of external respiration begins again. Gaseous Exchange Erythrocyte (red blood cells) in the mammalian body are filled with a globular protein called Haemoglobin. This molecule consists of 4 polypeptide chains each with an Iron Haem group in the centre. This Haem group is very important for gaseous exchange. Once the Erythrocyte, laden with Oxygen molecules, reaches some tissues which have an Oxygen deficit, the following reaction occurs: CO2 + H2O <> H2CO3 <> HCO3- + H + (<> = a reversible reaction) This equation is catalysed by the highly efficient enzyme called Carbonic Anhydrase. The CO2 is absorbed into the red blood cell then in this equation it is reacted with water to make firstly to Hydrogen Carbonate and then to Hydrogen Carbonate- ions and H+ ions. The HCO3- ions are pumped out of the cell and are replaced with Cl- ions to keep the charges balanced. The next stage in this process is as follows: H+ + HbO8 <> HHb+ + 4O2 Here I have used Hb for Haemoglobin for simplicity because Hemoglobin is actually many thousands of molecules long. This stage of the reaction involves the Hydrogen ions bonding with HbO8 (Oxygen bonded with Haemoglobin). The Hydrogen ions displace the Oxygen molecules from the Haem groups and allow the Oxygen molecules to come into solution in the red blood cell. These then diffuse through the phospholipid bilayer membrane of the red blood cell and move into the surrounding tissues where Oxygen is needed leaving the Hydrogen bonded with the Haemoglobin acting as a spectator ion, as it is not actually used for anything. Once this reaction has occurred there will be Oxygen in the tissues and the Carbon Dioxide which was in the tissues will now be carried back to the lungs to be exchanged for more Oxygen. This reaction is entirely reversible and the opposite reaction happens at the lung to absorb new Oxygen.

Enzymes are very efficient catalysts for biochemical reactions. They speed up reactions by providing an alternative reaction pathway of lower activation energy. Like all catalysts, enzymes take part in the reaction - that is how they provide an alternative reaction pathway. But they do not undergo permanent changes and so remain unchanged at the end of the reaction. They can only alter the rate of reaction, not the position of the equilibrium. Most chemical catalysts catalyse a wide range of reactions. They are not usually very selective. In contrast enzymes are usually highly selective, catalysing specific reactions only. This specificity is due to the shapes of the enzyme molecules. Many enzymes consist of a protein and a non-protein (called the cofactor). The proteins in enzymes are usually globular. The intra- and intermolecular bonds that hold proteins in their secondary and tertiary structures are disrupted by changes in temperature and pH. This affects shapes and so the catalytic activity of an enzyme is pH and temperature sensitive. Cofactors may be:

organic groups that are permanently bound to the enzyme (prosthetic groups) cations - positively charged metal ions (activators), which temporarily bind to the active site of the enzyme, giving an intense positive charge to the enzyme's protein organic molecules, usually vitamins or made from vitamins ( coenzymes), which are not permanently bound to the enzyme molecule, but combine with the enzymesubstrate complex temporarily.

How Enzymes Work For two molecules to react they must collide with one another. They must collide in the right direction (orientation) and with sufficient energy. Sufficient energy means that between them they have enough energy to overcome the energy barrier to reaction. This is called the activation energy. Enzymes have an active site. This is part of the molecule that has just the right shape and functional groups to bind to one of the reacting molecules. The reacting molecule that binds to the enzyme is called the substrate. An enzyme-catalysed reaction takes a different 'route'. The enzyme and substrate form a reaction intermediate. Its formation has a lower activation energy than the reaction between reactants without a catalyst. A simplified picture Route A reactant 1 + reactant 2 ----------- product Route B reactant 1 + enzyme ------------intermediate intermediate + reactant 2 ---------------- product + enzyme So the enzyme is used to form a reaction intermediate, but when this reacts with another reactant the enzyme reforms. Lock and Key Hypothesis This is the simplest model to represent how an enzyme works. The substrate simply fits into the active site to form a reaction intermediate.

Induced fit hypothesis In this model the enzyme molecule changes shape as the substrate molecules gets close. The change in shape is 'induced' by the approaching substrate molecule. This more sophisticated model relies on the fact that molecules are flexible because single covalent bonds are free to rotate.

Factors affecting catalytic activity of enzymes Temperature

As the temperature rises, reacting molecules have more and more kinetic energy. This increases the chances of a successful collision and so the rate increases. There is a certain temperature at which an enzyme's catalytic activity is at its greatest (see graph). This optimal temperature is usually around human body temperature (37.5 oC) for the enzymes in human cells. Above this temperature the enzyme structure begins to break down (denature) since at higher temperatures intra- and intermolecular bonds are broken as the enzyme molecules gain even more kinetic energy. ph

Each enzyme works within quite a small pH range. There is a pH at which its activity is greatest (the optimal pH). This is because changes in pH can make and break intra- and intermolecular bonds, changing the shape of the enzyme and, therefore, its effectiveness. he rate of an enzyme-catalysed reaction depends on the concentrations of enzyme and substrate. As the concentration of either is increased the rate of reaction increases (see graphs). For a given enzyme concentration, the rate of reaction increases with increasing substrate concentration up to a point, above which any further increase in substrate concentration produces no significant change in reaction rate. This is because the active sites of the enzyme molecules at any given moment are virtually saturated with substrate. The enzyme/substrate complex has to dissociate before the active sites are free to accommodate more substrate. Provided that the substrate concentration is high and that temperature and pH are kept constant, the rate of reaction is proportional to the enzyme concentration. Inhibition of enzyme activity Some substances reduce or even stop the catalytic activity of enzymes in biochemical reactions. They block or distort the active site. These chemicals are called inhibitors, because they inhibit reaction. Inhibitors that occupy the active site and prevent a substrate molecule from binding to the enzyme are said to be active site-directed (or competitive, as they 'compete' with the substrate for the active site). Inhibitors that attach to other parts of the enzyme molecule, perhaps distorting its shape, are said to be non-active site-directed (or non competitive). Immobilized enzymes Enzymes are widely used commercially, for example in the detergent, food and brewing industries. Protease enzymes are used in 'biological' washing powders to speed up the breakdown of proteins in stains like blood and egg. Pectinase is used to produce and clarify fruit juices. Problems using enzymes commercially include:

they are water soluble which makes them hard to recover some products can inhibit the enzyme activity (feedback inhibition)

Enzymes can be immobilized by fixing them to a solid surface. This has a number of commercial advantages:

the enzyme is easily removed the enzyme can be packed into columns and used over a long period speedy separation of products reduces feedback inhibition thermal stability is increased allowing higher temperatures to be used higher operating temperatures increase rate of reaction

There are four principal methods of immobilization currently in use:

covalent bonding to a solid support adsorption onto an insoluble substance entrapment within a gel encapsulation behind a selectively permeable membrane

How Do Enzymes Function? Enzymes are "biological catalysts." "Biological" means the substance in question is produced or is derived from some living organism. "Catalyst" denotes a substance that has the ability to increase the rate of a chemical reaction, and is not changed or destroyed by the chemical reaction that it accelerates. Generally speaking, catalysts are specific in nature as to the type of reaction they can catalyze. Enzymes, as a subclass of catalysts, are very specific in nature. Each enzyme can act to catalyze only very select chemical reactions and only with very select substances. An enzyme has been described as a "key" which can "unlock" complex compounds. An enzyme, as the key, must have a certain structure or multi-dimensional shape that matches a specific section of the "substrate" (a substrate is the compound or substance which undergoes the change). Once these two components come together, certain chemical bonds within the substrate molecule change much as a lock is released, and just like the key in this illustration, the enzyme is free to execute its duty once again. Many chemical reactions do proceed but at such a slow rate that their progress would seem to be imperceptible at normally encountered environmental temperature. Consider for example, the oxidation of glucose or other sugars to useable energy by animals and plants. For a living organism to derive heat and other energy from sugar, the sugar must be oxidized (combined with oxygen) or metabolically "burned" However, in a living system, the oxidation of sugar must meet an additional condition; that oxidation of sugar must proceed essentially at normal body temperature. Obviously, sugar surrounded by sufficient oxygen would not oxidize very rapidly at this temperature. In conjunction with a series of enzymes created by the living organism, however, this reaction does proceed quite rapidly at temperatures up to 100F (38C). Therefore, enzymes allow the living organism to make use of the potential energy contained in sugar and other food substances. Enzymes or biological catalysts allow reactions that are necessary to sustain life proceed

relatively quickly at the normal environmental temperatures. Enzymes often increase the rate of a chemical reaction between 10 and 20 million times what the speed of reaction would be when left uncatalyzed (at a given temperature). Nutrients locked in certain organics are complex macromolecules, or in hard-to-digest matrices may be released or predigested by a high degree of heat or concentrated acid treatment. In an alternative manner, specific enzymes can promote the pre-digestion of certain complex nutrients and facilitate the release of highly digestible nutrients in organics during processing without the need of excessive heat or rigorous chemical treatment.

Transmission of Nerve Impulses The transmission of a nerve impulse along a neuron from one end to the other occurs as a result of chemical changes across the membrane of the neuron. The membrane of an unstimulated neuron is polarizedthat is, there is a difference in electrical charge between the outside and inside of the membrane. The inside is negative with respect to the outside

Polarization is established by maintaining an excess of sodium ions (Na+) on the outside and an excess of potassium ions (K+) on the inside. A certain amount of Na+ and K+ is always leaking across the membrane through leakage channels, but Na+/K+ pumps in the membrane actively restore the ions to the appropriate side. Other ions, such as large, negatively charged proteins and nucleic acids, reside within the cell. It is these large, negatively charged ions that contribute to the overall negative charge on the inside of the cell membrane as compared to the outside. In addition to crossing the membrane through leakage channels, ions may also cross through gated channels. Gated channels open in response to neurotransmitters, changes in membrane potential, or other stimuli. The following events characterize the transmission of a nerve impulse Resting potential. The resting potential describes the unstimulated, polarized state of a neuron (at about 70 millivolts). Graded potential. A graded potential is a change in the resting potential of the plasma membrane in the response to a stimulus. A graded potential occurs when the stimulus causes Na+ or K+ gated channels to open. If Na+ channels open, positive sodium ions enter, and the membrane depolarizes (becomes more positive). If the stimulus opens K+ channels, then positive potassium ions exit across the membrane and the membrane hyperpolarizes (becomes more negative). A graded potential is a local event that does not travel far from its origin. Graded potentials occur in cell bodies and dendrites. Light, heat, mechanical pressure, and chemicals, such as neurotransmitters, are examples of stimuli that may generate a graded potential (depending upon the neuron). Action potential. Unlike a graded potential, an action potential is capable of traveling long distances. If a depolarizing graded potential is sufficiently large, Na+ channels in the trigger zone open. In response, Na+ on the outside of the membrane becomes depolarized (as in a graded potential). If the stimulus is strong enough that is, if it is above a certain threshold leveladditional Na+ gates open, increasing the flow of Na+ even more, causing an action potential, or complete depolarization (from 70 to about +30 millivol ts). This, in turn, stimulates neighboring Na+ gates, farther down the axon, to open. In this manner, the action potential travels down the length of the axon as opened Na+ gates stimulate

neighboring Na+ gates to open. The action potential is an all-or-nothing event: When the stimulus fails to produce depolarization that exceeds the threshold value, no action potential results, but when threshold potential is exceeded, complete depolarization occurs. Repolarization. In response to the inflow of Na+, K+ channels open, this time allowing K+ on the inside to rush out of the cell. The movement of K+ out of the cell causes repolarization by restoring the original membrane polarization. Unlike the resting potential, however, in repolarization the K+ are on the outside and the Na+ are on the inside. Soon after the K+ gates open, the Na+ gates close.

Hyper polarization. By the time the K+ channels close, more K+ have moved out of the cell than is actually necessary to establish the original polarized potential. Thus, the membrane becomes hyperpolarized (about 80 millivolts). Refractory period. With the passage of the action potential, the cell membrane is in an unusual state of affairs. The membrane is polarized, but the Na+ and K+ are on the wrong sides of the membrane. During this refractory period, the axon will not respond to a new stimulus. To reestablish the original distribution of these ions, the Na+ and K+ are returned to their resting potential location by Na+/K+ pumps in the cell membrane. Once these ions are completely returned to their resting potential location, the neuron is ready for another stimulus.

The Synapse The junction across which a nerve impulse passes from an axon terminal to a neuron, muscle cell, or gland cell. A synapse, or synaptic cleft, is the gap that separates adjacent neurons or a neuron and a muscle. Transmission of an impulse across a synapse, from presynaptic cell to postsynaptic cell, may be electrical or chemical. In electrical synapses, the action potential travels along the membranes of gap junctions, small tubes of cytoplasm along the membranes of gap junctions, small tubes of cytoplasm that allow the transfer of ions between adjacent cells. In chemical synapses, action potentials are transferred across the synapse by the diffusion of chemicals, as follows: Calcium (Ca2+) gates open. When an action potential reaches the end of an axon, the depolarization of the membrane causes gated channels to open that allow Ca2+ to enter. Synaptic vesicles release neurotransmitter. The influx of Ca2+ into the terminal end of the axon causes synaptic vesicles to merge with the presynaptic membrane, releasing a neurotransmitter into the synaptic cleft. Neurotransmitter binds with postsynaptic receptors. The neurotransmitter diffuses across the synaptic cleft and binds with specialized protein receptors on the postsynaptic membrane. Different proteins are receptors for different neurotransmitters. The postsynaptic membrane is excited or inhibited. Depending upon the kind of neurotransmitter and the kind of membrane receptor, there are two possible outcomes for the postsynaptic membrane, both of which are graded potentials. oIf positive ion gates open (which allow more Na+ and Ca2+ to enter than K+ to exit), the

membrane becomes depolarized, which results in an excitatory postsynaptic potential (EPSP). If the threshold potential is exceeded, an action potential is generated. oIf K+ or chlorine ion (Cl) gates open (allowing K+ to exit or Cl to enter), the membran e becomes more polarized (hyperpolarized), which results in an inhibitory postsynaptic potential (IPSP). As a result, it becomes more difficult to generate an action potential on this membrane. The neurotransmitter is degraded and recycled. After the neurotransmitter binds to the postsynaptic membrane receptors, it is either transported back to and reabsorbed by the secreting neuron, or it is broken down by enzymes in the synaptic cleft. For example, the common neurotransmitter acetylcholine is broken down by cholinesterase. Reabsorbed and degraded neurotransmitters are recycled by the presynaptic cell. Here are some of the common neurotransmitters and the kinds of activity they generate: Acetylcholine (ACh) is commonly secreted at neuromuscular junctions, the gaps between motor neurons and muscle cells, where it stimulates muscles to contract (by opening gated positive ion channels). At other kinds of junctions, it typically produces an inhibitory postsynaptic potential. Epinephrine, norepinephrine (NE), dopamine, and serotonin are derived from amino acids and are secreted mostly between neurons of the CNS. Gamma aminobutyric acid (GABA) is usually an inhibitory neurotransmitter (opening gated Cl channels) among neurons in the brain.

Regulation of Digestion The activities of the digestive system are regulated by both hormones and neural reflexes. Four important hormones and their effects upon target cells follow. Gastrin is produced by enteroendocrine cells of the stomach mucosa. Effects include

oStimulation of gastric juice (especially HCl) secretion by gastric glands. oStimulation of smooth muscle contraction in the stomach, small intestine, and large intestine, which increases gastric and intestinal motility. oRelaxation of the pyloric sphincter, which promotes gastric emptying into the small intestine.

Secretin is produced by the enteroendocrine cells of the duodenal mucosa. Effects include

oStimulation of bicarbonate secretion by the pancreas, which neutralizes the acidity of chyme when released into the duodenum. oStimulation of bile production by the liver. oInhibition of gastric juice secretions and gastric motility, which, in turn, slows digestion in the stomach and retards gastric emptying.

Cholecystokinin (CCK) is produced by enteroendocrine cells of the duodenal mucosa. Effects include

oStimulation of bile release by the gallbladder. oStimulation of pancreatic juice secretion. oRelaxation of the hepatopancreatic ampulla, which allows flow of bile and pancreatic juices into the duodenum.

Gastric inhibitory peptide (GIP) is produced by enteroendocrine cells of the duodenal mucosa and causes the inhibition of gastric juice secretion and gastric motility, which, in turn, slows digestion in the stomach and retards gastric emptying.

The second regulatory agent of the digestive system is the nervous system. Stimuli that influence digestive activities may originate in the head, the stomach, or the small intestine. Based on these sites, there are three phases of digestive regulation: The cephalic phase comprises those stimuli that originate from the head: sight, smell, taste, or thoughts of food, as well as emotional states. In response, the following reflexes are initiated:

oNeural response. Stimuli that arouse digestion are relayed to the hypothalamus, which, in turn, initiates nerve impulses in the parasympathetic vagus nerve. These impulses innervate nerve networks of the GI tract (enteric nervous system), which promote contraction of smooth muscle (which causes peristalsis) and secretion of gastric juice. Stimuli that repress digestion (emotions of fear or anxiety, for example) innervate sympathetic fibers that suppress muscle contraction and secretion. oGeneral effects. The stomach prepares for the digestion of proteins.

The gastric phase describes those stimuli that originate from the stomach. These stimuli include distention of the stomach (which activates stretch receptors), low acidity (high pH), and the presence of peptides. In response, the following reflexes are initiated:

oNeural response. Gastric juice secretion and smooth muscle contraction are promoted. oHormonal response. Gastrin production is promoted. oGeneral effects. The stomach and small intestine prepare for the digestion of chyme, and gastric emptying is promoted.

The intestinal phase describes stimuli originating in the small intestine. These include distention of the duodenum, high acidity (low pH), and the presence of chyme (especially fatty acids and carbohydrates). In response, the following reflexes are initiated:

oNeural response. Gastric secretion and gastric motility are inhibited (enterogastric reflex). Intestinal secretions, smooth muscle contraction, and bile and pancreatic juice production are promoted. oHormonal response. Production of secretin, CCK, and GIP is promoted. oGeneral effects. Stomach emptying is retarded to allow adequate time for digestion (especially fats) in the small intestine. Intestinal digestion and motility are promoted.