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Seven
Alterations in the
Gastrointestinal System
CHAPTER
S
Autonomic Nervous System tructurally, the gastrointestinal tract is a long, hollow tube
Swallowing and Esophageal Motility with its lumen inside the body and its wall acting as an
Gastric Motility interface between the internal and external environments.
Small Intestinal Motility The wall does not normally allow harmful agents to enter the
Colonic Motility body, nor does it permit body uids and other materials to es-
cape. The process of digestion and absorption of nutrients re-
Defecation
quires an intact and healthy gastrointestinal tract epithelial
Hormonal and Secretory Function lining that can resist the effects of its own digestive secretions.
Gastrointestinal Hormones The process also involves movement of materials through the
Gastrointestinal Secretions gastrointestinal tract at a rate that facilitates absorption, and it
Salivary Secretions requires the presence of enzymes for the digestion and absorp-
Gastric Secretions tion of nutrients.
Intestinal Secretions As a matter of semantics, the gastrointestinal tract also is re-
ferred to as the digestive tract, the alimentary canal, and at times,
the gut. The intestinal portion also may be called the bowel. For
the purposes of this text, the salivary glands, the liver, and the
pancreas, which produce secretions that aid in digestion, are
considered accessory organs.
459
460 Unit Seven: Alterations in the Gastrointestinal System
the rst three parts of the gastrointestinal tract. The liver and
STRUCTURE AND ORGANIZATION pancreas are discussed in Chapter 28.
OF THE GASTROINTESTINAL TRACT
In the digestive tract, food and other materials move slowly Upper Gastrointestinal Tract
along its length as they are systematically broken down into
ions and molecules that can be absorbed into the body. In the The mouth forms the entryway into the gastrointestinal tract
large intestine, unabsorbed nutrients and wastes are collected for food; it contains the teeth, used in the mastication of food,
for later elimination. Although the gastrointestinal tract is lo- and the tongue and other structures needed to direct food to-
cated inside the body, it is a long, hollow tube, the lumen (i.e., ward the pharyngeal structures and the esophagus.
hollow center) of which is an extension of the external envi-
ronment. Nutrients do not become part of the internal envi- Esophagus
ronment until they have passed through the intestinal wall and The esophagus is a tube that connects the oropharynx with
have entered the blood or lymph channels. the stomach. The esophagus begins at the lower end of the
For simplicity and understanding, the digestive system can pharynx. It is a muscular, collapsible tube, approximately
be divided into four parts (Fig. 26-1). The upper partthe 25 cm (10 in) long, that lies behind the trachea. The muscu-
mouth, esophagus, and stomachacts as an intake source and lar walls of the upper third of the esophagus are skeletal-type
receptacle through which food passes and in which initial di- striated muscle; these muscle fibers are gradually replaced by
gestive processes take place. The middle portion consists of the smooth muscle fibers until, at the lower third of the esopha-
small intestinethe duodenum, jejunum, and ileum. Most di- gus, the muscle layer is entirely smooth muscle.
gestive and absorptive processes occur in the small intestine. The esophagus functions primarily as a conduit for passage
The lower segmentthe cecum, colon, and rectumserves as of food from the pharynx to the stomach, and the structures of
a storage channel for the efcient elimination of waste. The its walls are designed for this purpose: the smooth muscle lay-
fourth part consists of the accessory organsthe salivary ers provide the peristaltic movements needed to move food
glands, liver, and pancreas. These structures produce digestive along its length, and the epithelial layer secretes mucus, which
secretions that help dismantle foods and regulate the use and protects its surface and aids in lubricating food. There are
storage of nutrients. The discussion in this chapter focuses on sphincters at either end of the esophagus: an upper esophageal
Soft palate
Hard palate
Oral cavity Nasopharynx
Tongue
Sublingual gland Parotid gland
Submandibular Oropharynx
gland
Pharynx
Trachea
Esophagus
Stomach Duodenum
The stomach is a pouchlike structure that lies in the upper part
of the abdomen and serves as a food storage reservoir during
the early stages of digestion. Although the residual volume of
the stomach is only approximately 50 mL, it can increase to
almost 1000 mL before the intraluminal pressure begins to rise.
The esophagus opens into the stomach through an opening Antrum
called the cardiac orice, so named because of its proximity to FIGURE 26-2 Structures of the stomach, showing the pace-
the heart. The part of the stomach that lies above and to the left maker area and the direction of chyme movement resulting from
of the cardiac orice is called the fundus, the central portion is peristaltic contractions.
462 Unit Seven: Alterations in the Gastrointestinal System
(splenic) exure. The descending colon extends from the colic Jejunum
exure to the rectum. The rectum extends from the sigmoid
colon to the anus. The anal canal passes between the two me-
dial borders of the levator ani muscles. Powerful sphincter
muscles guard against fecal incontinence.
Peritoneum Duodenum
Transverse
colon
Longitudinal
muscle
Mesentery
Greater
Circular omentum
muscle
Small intestine
Submucosa Parietal
peritoneum Uterus
Lumen of gut
Visceral
Mucosa peritoneum Rectum
(mucous Bladder
membrane)
The greater omentum protects the intestines from cold. It Enteric Nervous System
always contains some fat, which in obese persons can be a con- The intramural neurons (i.e., those contained within the wall
siderable amount. The omentum also controls the spread of in- of the gastrointestinal tract) consist of two networks: the
fection from gastrointestinal contents. In the case of infection, myenteric and submucosal plexuses. Both plexuses are aggre-
the omentum adheres to the inamed area so that the infection gates of ganglionic cells that extend along the length of the
is less likely to enter the peritoneal cavity. The lesser omentum gastrointestinal wall. The myenteric (Auerbachs) plexus is lo-
extends between the transverse ssure of the liver and the lesser cated between the circular muscle and longitudinal muscle
curvature of the stomach. layers, and the submucosal (Meissners) plexus between the
mucosal layer and the circular muscle layers (Fig. 26-6). The
In summary, the gastrointestinal tract is a long, hollow activity of the neurons in the myenteric and submucosal
tube, the lumen of which is an extension of the external envi- plexuses is regulated by local influences, input from the ANS,
ronment. The digestive tract can be divided into four parts: and by interconnecting fibers that transmit information
an upper part, consisting of the mouth, esophagus, and between the two plexuses.
stomach; a middle part, consisting of the small intestine; a The myenteric plexus consists mainly of a linear chain of
lower part, consisting of the cecum, colon, and rectum; and interconnecting neurons that extend the full length of the gas-
the accessory organs, consisting of the salivary glands, the trointestinal tract. Because it extends all the way down the in-
liver, and the pancreas. Throughout its length, except for the testinal wall and because it lies between the two muscle layers,
mouth, throat, and upper esophagus, the gastrointestinal it is concerned mainly with motility along the length of the gut.
tract is composed of ve layers: an inner mucosal layer, a sub- The submucosal plexus, which lies between the mucosal and
mucosal layer, a layer of circular smooth muscle bers, a layer circular muscle layers of the intestinal wall, is mainly con-
of longitudinal smooth muscle bers, and an outer serosal cerned with controlling the function of each segment of the in-
layer that forms the peritoneum and is continuous with the testinal tract. It integrates signals received from the mucosal
mesentery. layer into local control of motility, intestinal secretions, and
absorption of nutrients.
Intramural plexus neurons also communicate with receptors
in the mucosal and muscle layers. Mechanoreceptors monitor
the stretch and distention of the gastrointestinal tract wall, and
INNERVATION AND MOTILITY chemoreceptors monitor the chemical composition (i.e., osmo-
lality, pH, and digestive products of protein and fat metabo-
The motility of the gastrointestinal tract propels food prod- lism) of its contents. These receptors can communicate directly
ucts and fluids along its length, from mouth to anus, in a with ganglionic cells in the intramural plexuses or with visceral
manner that facilitates digestion and absorption. Except in afferent bers that inuence ANS control of gastrointestinal
the pharynx and upper third of the esophagus, smooth mus- function.
cle provides the contractile force for gastrointestinal motility
(the actions of smooth muscle are discussed in Chapter 1). Autonomic Nervous System
The rhythmic movements of the digestive tract are self- The gastrointestinal tract is innervated by both the sympa-
perpetuating, much like the activity of the heart, and are in- thetic and parasympathetic nervous systems. Parasympathetic
fluenced by local, humoral (i.e., blood-borne), and neural in- innervation to the stomach, small intestine, cecum, ascending
fluences. The ability to initiate impulses is a property of the colon, and transverse colon occurs by way of the vagus nerve
smooth muscle itself. Impulses are conducted from one mus- (Fig. 26-7). The remainder of the colon is innervated by para-
cle fiber to another. sympathetic fibers that exit the sacral segments of the spinal
The smooth muscle movements of the gastrointestinal tract cord by way of the pelvic nerve. Preganglionic parasympa-
are tonic and rhythmic. The tonic movements are continuous thetic fibers can synapse with intramural plexus neurons,
movements that last for minutes or even hours. Tonic contrac- or they can act directly on intestinal smooth muscle. Most
tions occur at sphincters. The rhythmic movements consist of parasympathetic fibers are excitatory. Numerous vagovagal
intermittent contractions that are responsible for mixing and reflexes influence motility and secretions of the digestive
moving food along the digestive tract. Peristaltic movements are tract.
rhythmic propulsive movements that occur when the smooth Sympathetic innervation of the gastrointestinal tract occurs
muscle layer constricts, forming a contractile band that forces through the thoracic chain of sympathetic ganglia and the
the intraluminal contents forward. During peristalsis, the seg- celiac, superior mesenteric, and inferior mesenteric ganglia.
ment that lies distal to, or ahead of, the contracted portion re- Sympathetic control of gastrointestinal function is largely me-
laxes, and the contents move forward with ease. Normal peri- diated by altering the activity of neurons in the intramural
stalsis always moves in the direction from the mouth toward plexuses. The sympathetic nervous system exerts several effects
the anus. on gastrointestinal function. It controls mucus secretion by the
mucosal glands, reduces motility by inhibiting the activity of
Innervation intramural plexus neurons, enhances sphincter function, and
increases the vascular smooth muscle tone of the blood vessels
Gastrointestinal function is controlled by the enteric nervous sys- that supply the gastrointestinal tract. The sympathetic bers
tem, which lies entirely within the wall of the gastrointestinal that supply the lower esophageal, pyloric, and internal and
tract, and by the parasympathetic and sympathetic divisions of external anal sphincters are largely excitatory, but their role in
the autonomic nervous system (ANS). controlling these sphincters is poorly understood.
464 Unit Seven: Alterations in the Gastrointestinal System
Epithelium
Lamina
Mucous Villi
propria
membrane
Muscularis
mucosae
Submucous
plexus
Gland in Submucosa
mucous
membrane Circular
Muscular
Longitudinal coat
Gland in
submucosa
Mesentery
FIGURE 26-6 Diagram of the four main layers of the wall of the digestive tube: mucosa,
submucosa, muscular, and serosa (below the diaphragm).
swallowing is initiated. The soft palate is pulled upward, the control of gastric emptying. Afferent receptor bers synapse
palatopharyngeal folds are pulled together so that food does with the neurons in the intramural plexus or trigger intrinsic re-
not enter the nasopharynx, the vocal cords are pulled together, exes by means of vagal or sympathetic pathways that partici-
and the epiglottis is moved so that it covers the larynx. Res- pate in extrinsic reexes.
piration is inhibited, and the bolus is moved backward into the Disorders of gastric motility can occur when the rate is too
esophagus by constrictive movements of the pharynx. Although slow or too fast (see Chapter 27). A rate that is too slow leads
the striated muscles of the pharynx are involved in the second to gastric retention. It can be caused by obstruction or gastric
stage of swallowing, it is an involuntary stage. atony. Gastric atony can occur as a complication of visceral
The third phase of swallowing is the esophageal stage. As food neuropathies in diabetes mellitus. Surgical procedures that dis-
enters the esophagus and stretches its walls, local and central rupt vagal activity also can result in gastric atony. Abnormally
nervous system reexes that initiate peristalsis are triggered. fast emptying occurs in the dumping syndrome, which is a con-
There are two types of peristalsis: primary and secondary. Pri- sequence of certain types of gastric operations. This condition is
mary peristalsis is controlled by the swallowing center in characterized by the rapid dumping of highly acidic and hyper-
the brain stem and begins when food enters the esophagus. osmotic gastric secretions into the duodenum and jejunum.
Secondary peristalsis is partially mediated by smooth muscle
bers in the esophagus and occurs when primary peristalsis is Small Intestinal Motility
inadequate to move food through the esophagus. Peristalsis be-
gins at the site of distention and moves downward. Before the The small intestine is the major site for the digestion and ab-
peristaltic wave reaches the stomach, the lower esophageal sorption of food; its movements are mixing and propulsive.
sphincter relaxes to allow the bolus of food to enter the stom- Regular peristaltic movements begin in the duodenum near the
ach. The pressure in the lower esophageal sphincter normally entry sites of the common duct and the main hepatic duct. A
is greater than that in the stomach, an important factor in pre- series of local pacemakers maintains the frequency of intes-
venting the reux of gastric contents. tinal contraction. The peristaltic movements (approximately
12 per minute in the jejunum) become less frequent as they
Gastric Motility move further from the pylorus, becoming approximately 9 per
minute in the ileum.
The stomach serves as a reservoir for ingested solids and liq- The peristaltic contractions produce segmentation waves
uids. Motility of the stomach results in the churning and grind- and propulsive movements of the small intestine. With seg-
ing of solid foods and regulates the emptying of the gastric con- mentation waves, slow contractions of circular muscle occlude
tents, or chyme, into the duodenum. Peristaltic mixing and the lumen and drive the contents forward and backward. Most
churning contractions begin in a pacemaker area in the middle of the contractions that produce segmentation waves are local
of the stomach and move toward the antrum (see Fig. 26-2). events involving only 1 to 4 cm at a time. They function mainly
They occur at a frequency of three to ve contractions per to mix the chyme with the digestive enzymes from the pancreas
minute, each with a duration of 2 to 20 seconds. As the peri- and to ensure adequate exposure of all parts of the chyme to
staltic wave approaches the antrum, it speeds up, and the en- the mucosal surface of the intestine, where absorption takes
tire terminal 5 to 10 cm of the antrum contracts, occluding the place. The frequency of segmenting activity increases after a
pyloric opening. Contraction of the antrum reverses the move- meal, presumably stimulated by receptors in the stomach and
ment of the chyme, returning the larger particles to the body of intestine.
the stomach for further churning and kneading. Because the Propulsive movements occur with synchronized activity in
pylorus is contracted during antral contraction, the gastric con- a section 10 to 20 cm long. They are accomplished by contrac-
tents are emptied into the duodenum between contractions. tion of the proximal portion of the intestine with the sequen-
Although the pylorus does not contain a true anatomic tial relaxation of its distal, or anal, portion. After material has
sphincter, it does function as a physiologic sphincter to prevent been propelled to the ileocecal junction by peristaltic move-
the backow of gastric contents and allow them to ow into the ment, stretching of the distal ileum produces a local reex that
duodenum at a rate commensurate with the ability of the duo- relaxes the sphincter and allows uid to squirt into the cecum.
denum to accept them. This is important because the regurgi- Motility disturbances of the small bowel are common, and
tation of bile salts and duodenal contents can damage the mu- auscultation of the abdomen can be used to assess bowel
cosal surface of the antrum and lead to gastric ulcers. Likewise, activity. Inammatory changes increase motility. In many in-
the duodenal mucosa can be damaged by the rapid inux of stances, it is not certain whether changes in motility occur be-
highly acid gastric contents. cause of inammation or occur secondary to toxins and un-
Like other parts of the gastrointestinal tract, the stomach is absorbed materials. Delayed passage of materials in the small
richly innervated by the enteric nervous system and its con- intestine also can be a problem. Transient interruption of in-
nections with the sympathetic and parasympathetic nervous testinal motility often occurs after gastrointestinal surgery. In-
systems. Axons from the intramural plexuses innervate the tubation with suction often is required to remove the accumu-
smooth muscles and glands of the stomach. Parasympathetic lating intestinal contents and gases until activity is resumed.
innervation is provided by the vagus nerve and sympathetic
innervation by the celiac ganglia. The emptying of the stom- Colonic Motility
ach is regulated by hormonal and neural mechanisms. The hor-
mones cholecystokinin and gastric inhibitory peptide, which The storage function of the colon dictates that movements in
are thought to control gastric emptying, are released in response this section of the gut are different from those in the small in-
to the pH and the osmolar and fatty acid composition of the testine. Movements in the colon are of two types. First are the
chyme. Local and central circuitry are involved in the neural segmental mixing movements, called haustrations, so named
466 Unit Seven: Alterations in the Gastrointestinal System
Cholecystokinin Duodenum, jejunum Products of protein digestion and long Stimulates contraction of the gallbladder;
chain fatty acids stimulates secretion of pancreatic enzymes;
slows gastric emptying
Gastrin Antrum of the stomach, Vagal stimulation; epinephrine; neutral Stimulates secretion of gastric acid and pep-
duodenum amino acids; calcium-containing uids sinogen; increases gastric blood ow; stimu-
such as milk; and alcohol. Secretion is lates gastric smooth muscle contraction;
inhibited by acid contents in the an- stimulates growth of gastric, small intestine,
trum of the stomach (below pH 2.5) and colon mucosa
Secretin Duodenum Acid pH or chyme entering duodenum Stimulates secretion of bicarbonate-containing
(below pH 3.0) solution by pancreas and liver
Gastric pits
Mucosa Mucous
cell
Parietal or
oxyntic cell
Peptic or
chief cell
are concentrated at the site where the contents from the stom- tract secretions. Neural, humoral, and local mechanisms con-
ach and secretions from the liver and pancreas enter the duo- tribute to the control of these secretions. The parasympa-
denum. These glands secrete large amounts of alkaline mucus thetic nervous system increases secretion, and sympathetic
that protect the duodenum from the acid content in the gastric activity exerts an inhibitory effect. In addition to secreting
chyme and from the action of the digestive enzymes. The ac- uids containing digestive enzymes, the gastrointestinal tract
tivity of Brunners glands is strongly inuenced by ANS activ- produces and secretes hormones, such as gastrin, secretin,
ity. For example, sympathetic stimulation causes a marked de- and cholecystokinin, that contribute to the control of
crease in mucus production, leaving this area more susceptible gastrointestinal function.
to irritation.
In addition to mucus, the intestinal mucosa produces two
other types of secretions. The rst is a serous uid (pH 6.5 to
7.5) secreted by specialized cells (i.e., crypts of Lieberkhn) in
the intestinal mucosal layer. This uid, which is produced at DIGESTION AND ABSORPTION
the rate of 2000 mL/day, acts as a vehicle for absorption. The
second type of secretion consists of surface enzymes that aid Digestion and absorption occur mainly in the small intestine.
absorption. These enzymes are the peptidases, or enzymes that The stomach is a poor absorptive structure, and only a few
separate amino acids, and the disaccharidases, or enzymes that lipid-soluble substances, including alcohol, are absorbed from
split sugars. the stomach.
The large intestine usually secretes only mucus. ANS activ- Digestion is the process of dismantling foods into their con-
ity strongly inuences mucus production in the bowel, as in stituent parts. Digestion requires hydrolysis, enzyme cleavage,
other parts of the digestive tract. During intense parasympa- and fat emulsication. Hydrolysis is breakdown of a com-
thetic stimulation, mucus secretion may increase to the point pound that involves a chemical reaction with water. The im-
that the stool contains large amounts of obvious mucus. portance of hydrolysis to digestion is evidenced by the amount
Although the bowel normally does not secrete water or elec- of water (7 to 8 L) that is secreted into the gastrointestinal tract
trolytes, these substances are lost in large quantities when the daily. The intestinal mucosa is impermeable to most large mol-
bowel becomes irritated or inamed. ecules. Most proteins, fats, and carbohydrates must be broken
down into smaller particles before they can be absorbed. Al-
though some digestion of carbohydrates and proteins begins in
In summary, the secretions of the gastrointestinal tract the stomach, digestion takes place mainly in the small intes-
include saliva, gastric juices, bile, and pancreatic and intesti- tine. The breakdown of fats to free fatty acids and monoglyc-
nal secretions. Each day, more than 7000 mL of uid is se- erides takes place entirely in the small intestine. The liver, with
creted into the digestive tract; all but 50 to 200 mL of this its production of bile, and the pancreas, which supplies a num-
uid is reabsorbed. Water, derived from the extracellular uid ber of digestive enzymes, play important roles in digestion.
compartment, is the major component of gastrointestinal Absorption is the process of moving nutrients and other ma-
terials from the external environment of the gastrointestinal
Chapter 26: Structure and Function of the Gastrointestinal System 469
tract into the internal environment. Absorption is accomplished Enterocyte being extruded
by active transport and diffusion. The absorptive function of from a villus
the large intestine focuses mainly on water reabsorption. A
number of substances require a specic carrier or transport sys-
tem. For example, vitamin B12 is not absorbed in the absence of Enterocyte
intrinsic factor, which is secreted by the parietal cells of the
stomach. Transport of amino acids and glucose occurs mainly
in the presence of sodium. Water is absorbed passively along
an osmotic gradient.
The distinguishing characteristic of the small intestine is its Vein
large surface area, which in the adult is estimated to be ap- Lacteal
proximately 250 m2. Anatomic features that contribute to this
Artery
enlarged surface area are the circular folds that extend into the
lumen of the intestine and the villi, which are nger-like pro-
jections of mucous membrane, numbering as many as 25,000,
that line the entire small intestine (Fig. 26-9). Each villus is Crypt of
equipped with an arrangement of blood vessels for the ab- Lieberkhn
sorption of uid and dissolved material into the portal blood
and a central lacteal for absorption into the lymph (Fig. 26-10).
Fats rely largely on the lymphatics for absorption.
Each villus is covered with cells called enterocytes that con-
tribute to the absorptive and digestive functions of the small
bowel, and goblet cells that provide mucus. The crypts of FIGURE 26-10 A single villus from the small intestine.
Lieberkhn are glandular structures that open into the spaces
between the villi. The enterocytes have a life span of approxi-
mately 4 to 5 days; their replacement cells differentiate from
progenitor cells located in the area of the crypts. The maturing ecules diffuse through the membrane or are actively trans-
enterocytes migrate up the villus and eventually are extruded ported across the mucosal surface to enter the blood or, in the
from the tip. case of fatty acids, the lacteal. These molecules are then trans-
The enterocytes secrete enzymes that aid in the digestion of ported through the portal vein or lymphatics into the systemic
carbohydrates and proteins. These enzymes are called brush circulation.
border enzymes because they adhere to the border of the villus
structures. In this way they have access to the carbohydrates Carbohydrates
and protein molecules as they come in contact with the ab-
sorptive surface of the intestine. This mechanism of secretion Carbohydrates must be broken down into monosaccharides, or
places the enzymes where they are needed and eliminates the single sugars, before they can be absorbed from the small in-
need to produce enough enzymes to mix with the entire con- testine. The average daily intake of carbohydrate in the
tents that ll the lumen of the small bowel. The digested mol- American diet is approximately 350 to 400 g. Starch makes up
approximately 50% of this total, sucrose (i.e., table sugar) ap-
proximately 30%, lactose (i.e., milk sugar) approximately 6%,
and maltose approximately 1.5%.
Digestion of starch begins in the mouth with the action of
amylase. Pancreatic secretions also contain an amylase. Amy-
lase breaks down starch into several disaccharides, including
maltose, isomaltose, and -dextrins. The brush border en-
Villus
zymes convert the disaccharides into monosaccharides that can
be absorbed (Table 26-3). Sucrose yields glucose and fructose,
Crypt of
..
Lieberkuhn
lactose is converted to glucose and galactose, and maltose is
converted to two glucose molecules. When the disaccharides
Mucosal are not broken down to monosaccharides, they cannot be ab-
muscle
sorbed but remain as osmotically active particles in the con-
Submucosa
tents of the digestive system, causing diarrhea. Persons who are
decient in lactase, the enzyme that breaks down lactose, ex-
perience diarrhea when they drink milk or eat dairy products.
Fructose is transported across the intestinal mucosa by fa-
Circular cilitated diffusion, which does not require energy expenditure.
muscle Longitudinal
muscle Lymph
Serosa
In this case, fructose moves along a concentration gradient.
node Glucose and galactose are transported by way of a sodium-
FIGURE 26-9 The mucous membrane of the small intestine. dependent carrier system that uses adenosine triphosphate and
Note the numerous villi on a circular fold. the Na+/K+-ATPase pump as an energy source (Fig. 26-11).
470 Unit Seven: Alterations in the Gastrointestinal System
Water absorption from the intestine is linked to absorption of Fat that is not absorbed in the intestine is excreted in the
osmotically active particles, such as glucose and sodium. It fol- stool. Steatorrhea is the term used to describe fatty stools. It usu-
lows that an important consideration in facilitating the trans- ally indicates that there is 20 g or more of fat in a 24-hour stool
port of water across the intestine (and decreasing diarrhea) sample. Normally, a chemical test is done on a 72-hour stool
after temporary disruption in bowel function is to include collection, during which time the diet is restricted to 80 to
sodium and glucose in the uids that are taken. 100 g of fat per day.
Fats Proteins
The average adult eats approximately 60 to 100 g of fat daily, Proteins from the diet must be broken into amino acids to be
principally as triglycerides containing long-chain fatty acids. absorbed. Protein digestion begins in the stomach with the ac-
These triglycerides are broken down by pancreatic lipase. Bile tion of pepsin. Pepsinogen, the enzyme precursor of pepsin, is
salts act as a carrier system for the fatty acids and fat-soluble secreted by the chief cells in response to a meal and acid pH.
vitamins A, D, E, and K by forming micelles, which transport Acid in the stomach is required for the conversion of pepsino-
these substances to the surface of intestinal villi, where they are gen to pepsin. Pepsin is inactivated when it enters the intestine
absorbed by the lacteal. The major site of fat absorption is the by the alkaline pH.
upper jejunum. Medium-chain triglycerides, with 6 to 10 car- Proteins are broken down further by pancreatic enzymes,
bon atoms in their structures, are absorbed better than longer- such as trypsin, chymotrypsin, carboxypeptidase, and elastase.
chain fatty acids because they are more completely broken As with pepsin, the pancreatic enzymes are secreted as precur-
down by pancreatic lipase and they form micelles more easily. sor molecules. Trypsinogen, which lacks enzymatic activity, is
Because they are easily absorbed, medium-chain triglycerides activated by an enzyme located on the brush border cells of the
often are used in the treatment of persons with malabsorption duodenal enterocytes. Activated trypsin activates additional
syndrome. The absorption of vitamins A, D, E, and K, which are trypsinogen molecules and other pancreatic precursor prote-
fat-soluble vitamins, requires bile salts. olytic enzymes. The amino acids are liberated intramurally or
on the surface of the villi by brush border enzymes that degrade
proteins into peptides that are one, two, or three amino acids
long. Similar to glucose, many amino acids are transported
Intestinal across the mucosal membrane in a sodium-linked process that
H2O
lumen uses the Na+/K+- ATPase pump as an energy source.
Body Glucose
fluids Glucose Glucose In summary, the digestion and absorption of foodstuffs
Carrier
+
take place in the small intestine. Digestion is the process of
Glucose Na dismantling foods into their constituent parts. Digestion re-
+
Carrier Na+ Na quires hydrolysis, enzyme cleavage, and fat emulsication.
Glucose
Na+ Proteins, fats, carbohydrates, and other components of the
K+
Glucose diet are broken down into molecules that can be transported
ATP from the intestinal lumen into the body uids. Absorption is
Na+ the process of moving nutrients and other materials from the
external environment of the gastrointestinal tract into the in-
ternal environment. Brush border enzymes break carbohy-
FIGURE 26-11 The hypothetical sodium-dependent transport
drates into monosaccharides that can be transported across
system for glucose. Both sodium and glucose must attach to the
the intestine into the bloodstream. The digestion of proteins
transport carrier before either can be transported into the cell.
The concentration of glucose builds up in the intestinal cell until begins in the stomach with the action of pepsin and is further
a diffusion gradient develops, causing glucose to move into the facilitated in the intestine by the pancreatic enzymes, such as
body uids. Sodium is transported out of the cell by the energy- trypsin, chymotrypsin, carboxypeptidase, and elastase. The
dependent Na+/K+-ATPase pump. This creates the gradient needed absorption of glucose and amino acids is facilitated by a
to operate the transport system. Water is passively absorbed sodium-dependent transport system. Fat in the diet is broken
along a concentration gradient generated by the absorption of down by pancreatic lipase into triglycerides containing
solutes.
Chapter 26: Structure and Function of the Gastrointestinal System 471
Vomiting, or emesis, is the sudden and forceful oral expulsion In summary, the signs and symptoms of many gastro-
of the contents of the stomach. It usually is preceded by nau- intestinal tract disorders are manifested by anorexia, nausea,
sea. The contents that are vomited are called vomitus. Vomit- and vomiting. Anorexia, or loss of appetite, may occur alone
ing, as a basic physiologic protective mechanism, limits the or may accompany nausea and vomiting. Nausea, which is an
possibility of damage from ingested noxious agents by empty- ill-dened, unpleasant sensation, signals the stimulation of the
ing the contents of the stomach and portions of the small in- medullary vomiting center. It often precedes vomiting and
testine. Nausea and vomiting may represent a total-body re- frequently is accompanied by autonomic responses, such as
sponse to drug therapy, including overdosage, cumulative salivation and vasoconstriction with pallor, sweating, and
effects, toxicity, and side effects. tachycardia. The act of vomiting, which is integrated by the
Vomiting involves two functionally distinct medullary cen- vomiting center, involves the forceful oral expulsion of the
ters: the vomiting center and the chemoreceptor trigger zone. gastric contents. It is a basic physiologic mechanism that rids
The act of vomiting is integrated by the vomiting center, which the gastrointestinal tract of noxious agents.
is located in the dorsal portion of the reticular formation of the
472 Unit Seven: Alterations in the Gastrointestinal System
REVIEW QUESTIONS
Describe the physiologic function of the digestive system in Visit the Connection site at connection.lww.com/go/porth
terms of transport of food, digestion, absorption of nutrients, for links to chapter-related resources on the Internet.
and storage of waste products.
Approximately 7000 mL of uid is secreted into the gastro-
intestinal tract, with only about 200 mL being eliminated in the BIBLIOGRAPHY
stool. Explain the source, composition, function, and removal
Berne R.M., Levy M.N. (1997). Principles of physiology (3rd ed., pp. 354
of this uid from the gastrointestinal tract. 400). St. Louis: C.V. Mosby.
Describe the site of gastric acid and pepsin production and Ganong W.F. (1999). Review of medical physiology (19th ed., 459491).
secretion in the stomach. Stanford: Appleton & Lange.
Gershon M.D. (1999). The enteric nervous system: A second brain.
Describe the function of the gastric mucosal barrier. Hospital Practice 34 (7), 3152.
Relate the characteristics of the small intestine to its absorp- Guyton A.C., Hall J.E. (2000). Textbook of medical physiology (10th ed.,
tive function and explain the function of intestinal brush border pp. 718770). Philadelphia: W.B. Saunders.
enzymes in the digestion and absorption of carbohydrates, fats, Johnson L.R. (2001). Gastrointestinal physiology (6th ed.). St. Louis: C.V.
Mosby.
and proteins.
Explain the relationship between the enteric nervous system
and the autonomic nervous system in the regulation of gastro-
intestinal function.
Describe the innervation of the reexive and voluntary
components of defecation.