Digestive System Functions

The digestive system is the organ system that processes food, extracts
nutrients from it, and eliminates the residue. It does this in
five stages:
1. ingestion, the selective intake of food;
2. digestion, the mechanical and chemical breakdown of food
into a form usable by the body;
3. absorption, the uptake of nutrients into the blood and lymph;
4. compaction, absorbing water and consolidating the
indigestible residue into feces; and finally
5. defecation, the elimination of feces.
General Anatomy
The digestive system has two anatomical subdivisions, the digestive
tract and the accessory organs (fig. 24.1). The digestive tract is a
muscular tube extending from mouth to anus, measuring about 9 m
(30 ft) long in the cadaver. It is also known as the alimentary2 canal
or gut. It includes the mouth, pharynx, esophagus, stomach, small
intestine, and large intestine. Part of it, the stomach and intestines,
constitutes the gastrointestinal (GI) tract. The accessory organs are
the teeth, tongue, salivary glands, liver, gallbladder, and pancreas.
The digestive tract is open to the environment at both ends.
Most of the material in it has not entered any body tissues and is
considered to be external to the body until it is absorbed by epithelial
cells of the alimentary canal. In the strict sense, defecated food
residue was never in the body.
Most of the digestive tract follows the basic structural plan
shown in figure 24.2, with a wall composed of the following tissue
layers, in order from the inner to the outer surface:
Mucosa
Epithelium
Lamina propria
Muscularis mucosae
Submucosa
Muscularis externa
Circular layer
Longitudinal layer
Serosa
Areolar tissue
Mesothelium
Slight variations on this theme are found in different regions of
the tract.
The mucosa (mucous membrane), which lines the lumen, consists
of an inner epithelium, a loose connective tissue layer called
the lamina propria, and a thin layer of smooth muscle called the
muscularis mucosae (MUSS-cue-LAIR-is mew-CO-see). The epithelium
is simple columnar in most of the digestive tract, but the

mouth, pharynx, esophagus, and anal canal differ. These upper and
lower ends of the digestive tract are subject to more abrasion than
the stomach and intestines, and thus have a stratified squamous
epithelium. The muscularis mucosae tenses the mucosa, creating grooves and ridges that enhance
its surface area and contact with
food. This improves the efficiency of digestion and nutrient absorption.
The mucosa often exhibits an abundance of lymphocytes
and lymphatic nodulesthe mucosa-associated lymphatic tissue
(MALT) (see p. 646).
The submucosa is a thicker layer of loose connective tissue containing
blood vessels, lymphatic vessels, a nerve plexus, and in some
places, mucous glands. The MALT also extends into the submucosa
in some parts of the GI tract.
The muscularis externa consists of usually two layers of smooth
muscle near the outer surface. Cells of the inner layer encircle the
tract and those of the outer layer run longitudinally. In some places,
the circular layer is thickened to form valves (sphincters) that regulate
the passage of material through the digestive tract. The muscularis
externa is responsible for the motility that propels food and
residue through the tract.
The serosa is composed of a thin layer of areolar tissue topped
by a simple squamous mesothelium. It begins in the lower 3 to 4
cm of the esophagus and ends just before the rectum. The pharynx,
most of the esophagus, and the rectum are surrounded by a fibrous
connective tissue layer called the adventitia (AD-ven-TISH-ah),
which blends into the adjacent connective tissues of other organs.
Innervation
Tongue movements, mastication, and the initial actions of swallowing
employ skeletal muscles innervated by somatic motor fibers
from six of the cranial nerves (V, VII, and IXXII) and from the
ansa cervicalis; these muscles and their innervation are detailed in
table 11.3 (p. 304). The salivary glands are innervated by sympathetic
fibers from the superior cervical ganglion and parasympathetic
fibers from cranial nerves VII and IX (see figs. 15.31, p. 457;
15.33, p. 458; and 16.4, p. 472).
From the lower esophagus to the anal canal, most of the muscle
is smooth muscle (the external anal sphincter is the only exception),
and therefore receives only autonomic innervation. Parasympathetic
innervation dominates the digestive tract and comes
mainly from the vagus nerves, which supply all of the tract from
esophagus to transverse colon. The descending colon and rectum
receive their parasympathetic innervation from pelvic nerves arising
from the inferior hypogastric plexus (see fig. 16.7, p. 476). The
parasympathetic nervous system relaxes sphincter muscles and
stimulates gastrointestinal motility and secretion. Thus, in general,
it promotes digestion.

The sympathetic nervous system plays a lesser role, but in general

it inhibits motility and secretion and keeps the GI sphincters
contracted and closed. Thus, it inhibits digestion. Sympathetic efferent
pathways travel through the celiac ganglion to the stomach,
liver, and pancreas; through the superior mesenteric ganglion to
the small intestine and most of the large intestine; and through the
inferior mesenteric ganglion to the rectum (see fig. 16.4, p. 472).
Even though the digestive tract receives such extensive innervation
from the CNS, it can function independently even if
these nerves are severed. This is because the esophagus, stomach,
and intestines have their own extensive nervous network called million neuronsmore than the
spinal cord! These include sensory
neurons that monitor tension in the gut wall; interneurons;
and motor neurons that activate such effectors as smooth muscle
and gland cells of the gut. It also contains clusters of parasympathetic
postganglionic neurons that act essentially as autonomic
ganglia.
Neurons of the enteric nervous system are distributed in two
networks: the submucosal (Meissner4) plexus in the submucosa
and the myenteric (Auerbach5) plexus between the two layers of
the muscularis externa. Parasympathetic preganglionic fibers terminate
in the ganglia of the myenteric plexus. Postganglionic fibers
arising in this plexus not only innervate the muscularis externa,
but also pass through its inner circular layer and contribute to the
submucosal plexus. The myenteric plexus controls peristalsis and
other contractions of the muscularis externa, while the submucosal
plexus controls movements of the muscularis mucosae and glandular
secretion of the mucosa.
Sensory nerve fibers monitor stretching of the GI wall and
chemical conditions in the lumen. These fibers carry signals to adjacent
regions of the GI tract in short (myenteric) reflex arcs contained
in the myenteric plexus, and to the central nervous system
by way of long (vagovagal) reflex arcs, predominantly in the vagus
nerves. These visceral reflex arcs enable different regions of the GI
tract to regulate each other over both short and long distances.
Circulation
We will see near the end of this chapter that the embryonic digestive
tract forms in three segments: the foregut, midgut, and hindgut.
These segments are defined by their arterial blood supply (see table
21.6, pp. 616619).
The foregut includes the mouth, pharynx, esophagus,
stomach, and the beginning of the duodenum (to the point
where the bile duct empties into it). Above the diaphragm,
the thoracic aorta gives off a series of esophageal arteries to
the esophagus. Below the diaphragm, the foregut components
receive their blood from branches of the celiac trunk.

 The midgut begins at the opening of the bile duct and

includes the rest of the duodenum, the jejunum and ileum
(the second and third portions of the small intestine), and the
large intestine as far as the first two-thirds of the transverse
colon. It receives blood from the superior mesenteric artery
(fig. 21.24a, p. 619).
The hindgut includes the remainder of the large intestine,
from the end of the transverse colon through the anal canal. It
is supplied by branches of the inferior mesenteric artery (fig.
21.24b, p. 619).
The most noteworthy general point about the venous drainage of
the GI tract is that blood from the entire tract below the diaphragm
ultimately drains into the hepatic portal vein, which enters the liver.
The system of vessels connecting the lower digestive tract to the liver
is the hepatic portal system (table 21.7, p. 620). It routes all blood
from the stomach and intestines, as well as from some other abdominal
viscera, through the liver before returning it to the general circulation.
Like other portal systems, this one has two capillary networks
in series. Capillaries in the small intestine receive digested nutrients,
and capillaries in the liver (the hepatic sinusoids described later) deliver
these nutrients to the liver cells. This gives the liver a chance to
process most nutrients and cleanse the intestinal blood of bacteria.
Relationship to the Peritoneum
In processing food, the stomach and intestines undergo such strenuous
contractions that they need freedom to move in the abdominal
cavity. Thus, they are not tightly bound to the abdominal wall, but over
most of its length, the tract is loosely suspended from it by connective
tissue sheets called the mesenteries (see figs. A.9 and A.10, pp. 29,
30). The mesenteries also hold the abdominal viscera in their proper
relationship to each other and prevent the small intestine, especially,
from becoming twisted and tangled by changes in body position and
by its own contractions. Furthermore, the mesenteries provide passage
for the blood vessels and nerves that supply the digestive tract,
and contain many lymph nodes and lymphatic vessels.
The parietal peritoneum is a serous membrane that lines the
wall of the abdominal cavity (see p. 000). Along the posterior
(dorsal) midline of the body, it turns inward and forms the dorsal
mesentery, a translucent two-layered membrane extending to
the digestive tract. Upon reaching an organ such as the stomach or
small intestine, the two layers of the mesentery separate and pass
around opposite sides of the organ, forming the serosa. In some
places, the two layers come together again on the far side of that organ
and continue as another sheet of tissue, the ventral mesentery.
The ventral mesentery may hang freely in the abdominal cavity or
attach to the ventral abdominal wall or other organs. The relationship
between the dorsal and ventral mesenteries and the serosa are

shown in figure A.9.

Along the right superior margin (lesser curvature) of the stomach,
a ventral mesentery called the lesser omentum extends from
the stomach to the liver (fig. 24.3). Another membrane, a fatty
greater omentum, hangs from the left inferior margin (greater
curvature) of the stomach and loosely covers the small intestine
like an apron. At its inferior margin, the greater omentum turns
back on itself and passes upward, thus forming a deep pouch between
its deep and superficial layers. At its inner superior margin,
it continues as a serosa enclosing the spleen and transverse colon,
then continues still farther as the mesocolon, which anchors the
transverse colon to the dorsal abdominal wall. The omenta have a
loosely organized, lacy appearance due partly to many holes or gaps
in the membranes and partly to an irregular distribution of adipose
tissue. They adhere to perforations or inflamed areas of the stomach
or intestines, contribute immune cells to the site, and isolate
infections that might otherwise give rise to peritonitis.
When an organ is enclosed by mesentery (serosa) on both sides,
it is considered to be within the peritoneal cavity, or intraperitoneal.6 When an organ lies
against the dorsal body wall and is covered
by peritoneum on the ventral side only, it is said to be outside
the peritoneal cavity, or retroperitoneal.7 The duodenum, most of
the pancreas, and parts of the large intestine are retroperitoneal. The
stomach, liver, and other parts of the small and large intestines are
intraperitoneal.
The Mouth
The mouth is also known as the oral (buccal8) cavity. Its functions
include ingestion (food intake), taste and other sensory responses to
food, chewing, chemical digestion, swallowing, speech, and respiration.
The mouth is enclosed by the cheeks, lips, palate, and tongue
(fig. 24.4). Its anterior opening between the lips is the oral fissure,
and its posterior opening into the throat is the fauces9 (FAW-seez).
The mouth is lined with stratified squamous epithelium. It is keratinized
in areas subject to the greatest abrasion, such as the gums
and hard palate, and nonkeratinized in other areas such as the floor
of the mouth, the soft palate, and the inside of the cheeks and lips.
The Cheeks and Lips
The cheeks and lips retain food and push it between the teeth for
mastication, and are essential for articulate speech and for sucking
and blowing actions, including suckling by infants. Their fleshiness
is due mainly to subcutaneous fat, the buccinator muscles of the
cheeks, and the orbicularis oris muscle of the lips. A median fold
called the labial frenulum10 attaches each lip to the gum, between
the anterior incisors. The vestibule is the space between the cheeks
or lips and the teeththe space where you insert your toothbrush

when brushing the outer surfaces of the teeth.

The lips are divided into three areas: (1) The cutaneous area is
colored like the rest of the face and has hair follicles and sebaceous
glands; on the upper lip, this is where a mustache grows. (2) The red
area (vermilion), is the hairless region where the lips meet (where
some people apply lipstick). It has unusually tall dermal papillae,
which allow blood capillaries and nerve endings to come closer to
the epidermal surface. Thus, this area is redder and more sensitive
than the cutaneous area. (3) The labial mucosa is the inner surface
of the lip, facing the gums and teeth.
The Tongue
The tongue (fig. 24.5), although muscular and bulky, is an agile
and sensitive organ with several functions: it aids in food intake;
it has sensory receptors for taste, texture, and temperature that are
important in the acceptance or rejection of food; it compresses and
breaks up food; it maneuvers food between the teeth for mastication;
it secretes mucus and enzymes; it compresses the chewed
food into a soft mass, or bolus, that is easier to swallow; it initiates
swallowing; and it is necessary for articulate speech. Its surface is
covered with nonkeratinized stratified squamous epithelium and
exhibits bumps and projections called lingual papillae, the site of
the taste buds. The types of papillae and sense of taste are discussed
in chapter 17.
The anterior two-thirds of the tongue, called the body, occupies
the oral cavity; the posterior one-third, the root, occupies the
oropharynx. The boundary between the body and root is marked
by a V-shaped row of vallate papillae and, behind these, a groove
called the terminal sulcus. A ventral median fold called the lingual
frenulum attaches the body of the tongue to the floor of the
mouth. The root of the tongue contains the lingual tonsils. Amid
the tongue muscles are serous and mucous lingual glands, which
secrete a portion of the saliva.
The muscles of the tongue, which compose most of its mass, are
described in chapter 11. The intrinsic muscles, contained entirely
within the tongue, produce the relatively subtle tongue movements
of speech. The extrinsic muscles, with origins elsewhere and insertions
in the tongue, produce the stronger movements of food
manipulation. The extrinsic muscles include the genioglossus, hyoglossus,
styloglossus, and palatoglossus (see table 11.3, p. 304).
The Palate
The palate, separating the oral cavity from the nasal cavity, makes
it possible to breathe while chewing food. Its anterior portion, the

hard (bony) palate, is supported by the palatine processes of the

maxillae and by the smaller palatine bones. It has transverse ridges
called palatine rugae that aid the tongue in holding and manipulating
food. Posterior to this is the soft palate, which has a more
spongy texture and is composed mainly of skeletal muscle and
glandular tissue, but no bone. It has a conical medial projection,
the uvula,11 visible at the rear of the mouth. The uvula helps to
retain food in the mouth until one is ready to swallow.
A pair of muscular arches on each side of the oral cavity begin
dorsally near the uvula and follow the wall of the cavity to its
floor. The anterior one is the palatoglossal arch and the posterior
one is the palatopharyngeal arch. The latter arch marks the
beginning of the pharynx. The palatine tonsils are located on the
wall between the arches.
The Teeth
The teeth are collectively called the dentition. They serve to masticate
the food, breaking it into smaller pieces. This not only makes
the food easier to swallow, but also exposes more surface area to the
action of digestive enzymes and thus speeds up chemical digestion.
Adults normally have 16 teeth in the mandible and 16 in the maxilla.
From the midline to the rear of each jaw, there are two incisors, a canine, two premolars, and up
to three molars (fig. 24.6a). The
incisors are chisel-like cutting teeth used to bite off a piece of food.
The canines are more pointed and act to puncture and shred it.
They serve as weapons in many mammals but became reduced in
the course of human evolution until they now project barely beyond
the other teeth. The premolars and molars have relatively
broad surfaces adapted for crushing and grinding.
Each tooth is embedded in a socket called an alveolus, forming a

joint called a gomphosis between the tooth and bone (fig. 24.7). The
alveolus is lined by a periodontal (PERR-ee-oh-DON-tul) ligament,
a modified periosteum whose collagen fibers penetrate into the bone
on one side and into the tooth on the other. This anchors the tooth
firmly in the alveolus, but allows for a slight amount of movement
under the pressure of chewing. The gum, or gingiva (JIN-jih-vuh),
covers the alveolar bone. Regions of a tooth are defined by their relationship
to the gingiva: the crown is the portion above the gum, the
root is the portion inserted into the alveolus below the gum, and the
neck is the line where the crown, root, and gum meet. The space between
the tooth and gum is the gingival sulcus. The hygiene of this
sulcus is especially important to dental health (see Insight 24.1).
Most of a tooth consists of hard yellowish tissue called dentine,
covered with enamel in the crown and cementum in the root.
Dentine and cementum are living connective tissues with cells or
cell processes embedded in a calcified matrix. Cells of the cementum
(cementocytes) are scattered more or less randomly and occupy
tiny cavities similar to the lacunae of bone. Cells of the dentine
(odontoblasts) line the pulp cavity and have slender processes that
travel through tiny parallel tunnels in the dentine. Enamel is not a
tissue but a cell-free secretion produced before the tooth emerges
above the gum. Damaged dentine and cementum can regenerate,
but damaged enamel cannotit must be artificially repaired.
Internally, a tooth has a dilated pulp cavity in the crown and
upper root, and a narrow root canal in the lower root. These spaces
are occupied by pulpa mass of loose connective tissue, blood and
lymphatic vessels, and nerves. These nerves and vessels enter the
tooth through a pore, the apical foramen, at the inferior end of
each root canal.
Tooth and Gum Disease
Food leaves a sticky residue on the teeth called plaque, composed
mainly of bacteria and sugars. If plaque is not thoroughly removed by
brushing and flossing, bacteria accumulate, metabolize the sugars, and
release lactic acid and other acids. These acids dissolve the minerals of
enamel and dentine, and the bacteria enzymatically digest the collagen
and other organic components. The eroded cavities of the tooth are
known as dental caries.12 If not repaired, caries may fully penetrate the
dentine and spread to the pulp cavity. This requires either extraction
of the tooth or root canal therapy, in which the pulp is removed and
replaced with inert material.
When plaque calcifies on the tooth surface, it is called calculus
(tartar). Calculus in the gingival sulcus wedges the tooth and gum
apart and allows bacterial invasion of the sulcus. This leads to gingivitis,
or gum inflammation. Nearly everyone has gingivitis at some time. In
some cases, bacteria spread from the sulcus into the alveolar bone and
begin to dissolve it, producing periodontal disease. About 86% of people

over age 70 have periodontal disease, and many suffer tooth loss as a
result. This accounts for 80% to 90% of adult tooth loss.
The meeting of the teeth when the mouth closes is called occlusion
(ah-CLUE-zhun), and the surfaces where they meet are called
the occlusal surfaces. The occlusal surface of a premolar has two
rounded bumps called cusps; thus the premolars are also known as
bicuspids. The molars have four to five cusps. Cusps of the upper
and lower premolars and molars mesh when the jaws are closed and
slide over each other as the jaw makes lateral chewing motions. This
grinds and tears food more effectively than if the occlusal surfaces
were flat.
Teeth develop beneath the gums and erupt (emerge) in predictable
order. Twenty deciduous teeth (milk teeth or baby teeth)
erupt from the ages of 6 to 30 months, beginning with the incisors
(fig. 24.6a). Between 6 and 25 years of age, these are replaced by 32
permanent teeth. As a permanent tooth grows deep to a deciduous
tooth (fig. 24.8), the root of the deciduous tooth dissolves and
leaves little more than the crown by the time it falls out. The third
molars (wisdom teeth) erupt around ages 17 to 25, if at all. Over
the course of human evolution, the face became flatter and the jaws
shorter, leaving little room for the third molars. Thus, they often
remain below the gum and become impactedso crowded against
neighboring teeth and bone that they cannot erupt.
The Salivary Glands
Saliva moistens the mouth, digests a small amount of starch and
fat, cleanses the teeth, inhibits bacterial growth, dissolves molecules
so they can stimulate the taste buds, and moistens food and binds
particles together to aid in swallowing. It is a solution of 97.0% to
99.5% water and the following solutes:
salivary amylase, an enzyme that begins starch digestion in
the mouth;
lingual lipase, an enzyme that is activated by stomach acid
and digests fat after the food is swallowed;
mucus, which binds and lubricates the food mass and aids in
swallowing;
lysozyme, an enzyme that kills bacteria;
immunoglobulin A (IgA), an antibody that inhibits bacterial
growth; and
electrolytes, including sodium, potassium, chloride,
phosphate, and bicarbonate salts.
There are two kinds of salivary glands, intrinsic and extrinsic. The

intrinsic salivary glands are an indefinite number of small glands

dispersed amid the other oral tissues. They include lingual glands in
the tongue, labial glands on the inside of the lips, and buccal glands
on the inside of the cheeks. They secrete saliva at a fairly constant rate
whether we are eating or not, but in relatively small amounts. This
saliva keeps the mouth moist and inhibits bacterial growth.
The extrinsic salivary glands are three pairs of larger, more
discrete organs located outside of the oral mucosa. They communicate
with the oral cavity by way of ducts (fig. 24.9). They are:
1. The parotid13 gland, located just beneath the skin anterior
to the earlobe. Its duct passes superficially over the masseter,
pierces the buccinator, and opens into the mouth opposite the
second upper molar tooth. Mumps is an inflammation and
swelling of the parotid gland caused by a virus.
2. The submandibular gland, located halfway along the body of
the mandible, medial to its margin, just deep to the mylohyoid
muscle. Its duct empties into the mouth at a papilla on the
side of the lingual frenulum, near the lower central incisors.
3. The sublingual gland, located in the floor of the mouth. It
has multiple ducts that empty into the mouth posterior to the
papilla of the submandibular duct.
These are all compound tubuloacinar glands with a treelike arrangement
of branching ducts ending in acini (see p. 96). Some
acini have only mucous cells, some have only serous cells, and some
have a mixture of both (fig. 24.10). Mucous cells secrete salivary
mucus, and serous cells secrete a thinner fluid rich in amylase and
electrolytes.
Salivation is controlled by groups of neurons called salivatory
nuclei in the medulla oblongata and pons. They receive signals
from sensory receptors in the mouth as well as from higher brain
centers that respond to the odor, sight, or thought of food. Efferent
nerve pathways to the salivary glands were described earlier
(p. 686). Salivation is mostly under the control of parasympathetic
fibers in cranial nerves VII and IX, which stimulate the secretion of
watery, enzyme-rich saliva. Sympathetic fibers from cervical ganglia
stimulate the secretion of thicker, mucus-rich saliva.
The Pharynx
The pharynx, as described in chapter 23, consists of three regions
called the nasopharynx, oropharynx, and laryngopharynx
(fig. 23.2c, p. 665). The first is exclusively respiratory and is lined
with pseudostratified columnar epithelium; the last two are shared
by the respiratory and digestive tracts and are lined with nonkeratinized
stratified squamous epithelium, an adaptation to withstanding

abrasion by passing food.

The pharynx has a deep layer of longitudinally oriented skeletal
muscle and a superficial layer of circular skeletal muscle. The circular
muscle is divided into superior, middle, and inferior pharyngeal
constrictors, which force food downward during swallowing. When one is not swallowing, the
inferior constrictor remains contracted to
exclude air from the esophagus. This constriction is regarded as the
upper esophageal sphincter, although it is not an anatomical feature
of the esophagus. It disappears at the time of death when the muscle
relaxes. Thus it is regarded as a physiological sphincter rather than a
constant anatomical structure.
The Esophagus
The esophagus is a straight muscular tube 25 to 30 cm long, posterior
to the trachea (see figs. 24.1 and 24.2). Its superior opening lies
between vertebra C6 and the cricoid cartilage of the larynx. After
passing downward through the mediastinum, the esophagus penetrates
the diaphragm at an opening called the esophageal hiatus,
continues another 3 to 4 cm, and meets the stomach at the level of
vertebra T7. Its opening into the stomach is called the cardiac orifice
(for its proximity to the heart). Food pauses briefly at this point before
entering the stomach because of a constriction called the lower
esophageal sphincter (LES). The LES is also a physiological rather
than an anatomical sphincter, and thus is not found in the cadaver. It
is thought to be either a constriction of the diaphragm surrounding
the esophageal hiatus, or muscle tone in the smooth muscle of the
esophagus. The LES prevents stomach contents from regurgitating
into the esophagus, thus protecting the esophageal mucosa from the
corrosive effect of the stomach acid (see Insight 24.2).
The wall of the esophagus is organized into the tissue layers described
earlier, with some regional specializations. The mucosa has
a nonkeratinized stratified squamous epithelium. The submucosa
contains esophageal glands, which secrete lubricating mucus into
the lumen. When the esophagus is empty, the mucosa and submucosa
are deeply folded into longitudinal ridges, giving the lumen a
starlike shape in cross section.
The muscularis externa is composed of skeletal muscle in the
upper one-third of the esophagus, a mixture of skeletal and smooth
muscle in the middle one-third, and only smooth muscle in the lower
one-third. This transition corresponds to a shift from voluntary to involuntary
phases of swallowing as food passes down the esophagus.
Most of the esophagus is in the mediastinum. Here, it is covered

with a connective tissue adventitia that merges into the adventitias

of the trachea and thoracic aorta. The short segment below the
diaphragm is covered by a serosa.
Swallowing, or deglutition (DEE-glu-TISH-un), is a complex
action involving over 22 muscles in the mouth, pharynx, and
esophagus, coordinated by the swallowing center, a pair of nuclei
in the medulla oblongata. This center communicates with muscles
of the pharynx and esophagus by way of the trigeminal, facial, glossopharyngeal,
and hypoglossal nerves (cranial nerves V, VII, IX, and
XII), and coordinates a complex series of muscle contractions to
produce swallowing without choking.
The stomach is a muscular sac in the upper left abdominal cavity
immediately inferior to the diaphragm (see fig. 24.1). It functions
primarily as a food storage organ, with an internal volume of about 50 mL when empty and 1.0
to 1.5 L after a typical meal.
When extremely full, it may hold up to 4 L and extend nearly as far
as the pelvis. The stomach mechanically breaks up food particles,
liquefies the food, and begins the chemical digestion of proteins
and a small amount of fat. This produces a soupy or pasty mixture
of semidigested food called chyme15 (pronounced kime). Most
digestion occurs after the chyme passes on to the small intestine.
Gross Anatomy
The stomach is somewhat J-shaped (fig. 24.11) and vertical in tall
people, whereas in short people it is more nearly horizontal. The
lesser curvature of the stomach extends the short distance (about
10 cm) from esophagus to duodenum along the medial to superior
aspect, facing the liver, and the greater curvature extends the
longer distance (about 40 cm) from esophagus to duodenum on
the lateral to inferior aspect. As described earlier, the lesser omentum
extends from the lesser curvature to the liver, and the greater
omentum is suspended from the greater curvature and overhangs
the intestines below.
The stomach is divided into four regions: (1) The cardiac region
(cardia) is the small area within about 3 cm of the cardiac
orifice. (2) The fundic region (fundus) is the dome-shaped portion
superior to the esophageal attachment. (3) The body (corpus)
makes up the greatest part of the stomach distal to the cardiac orifice.
(4) The pyloric region is a slightly narrower pouch at the distal
end; it is subdivided into a funnel-like antrum16 and a narrower
pyloric canal. The latter terminates at the pylorus,17 a narrow passage
into the duodenum. The pylorus is surrounded by a thick ring
of smooth muscle, the pyloric (gastroduodenal) sphincter, which
regulates the passage of chyme into the duodenum.
Microscopic Anatomy
The stomach wall has tissue layers similar to those of the esophagus,

with some variations. The surface of the mucosa is a simple

columnar glandular epithelium (fig. 24.12). The apical regions of
its surface cells are filled with mucin. After it is secreted, mucin
swells with water and becomes mucus. When the stomach is full,
the mucosa and submucosa are flat and smooth, but as it empties,
these layers fold into longitudinal wrinkles called gastric rugae18
(ROO-gee). The lamina propria is almost entirely occupied by tubular
glands, to be described shortly. The muscularis externa has
three layers, rather than twoan outer longitudinal, middle circular,
and inner oblique layer (see fig. 24.11a).
The gastric mucosa is pocked with depressions called gastric
pits lined with the same columnar epithelium as the mucosal surface.
Two or three tubular glands open into the bottom of each
gastric pit and span the rest of the lamina propria. The glands are
simple wavy or coiled tubes of more or less uniform diameter, except
for a constriction called the neck at the point where the gland
opens into the pit. In the cardiac and pyloric regions, they are called
cardiac glands and pyloric glands, respectively. In the rest of the
stomach, they are called gastric glands (fig. 24.12b, c). Collectively,
the glands have the following cell types:
Mucous cells, which secrete mucus, predominate in the cardiac
and pyloric glands. In gastric glands, they are called mucous
neck cells and are concentrated in the neck of the gland.
Regenerative (stem) cells, found in the base of the pit and neck
of the gland, divide rapidly and produce a continual supply of
new cells. Newly generated cells migrate upward to the gastric
surface as well as downward into the glands to replace cells that
die and fall off into the lumen of the stomach.
Parietal cells, found mostly in the upper half of the gland,
secrete hydrochloric acid (HCl) and intrinsic factor. They
are found mostly in the gastric glands, but a few occur in the
pyloric glands.
Chief cells, so named because they are the most numerous,
secrete chymosin (formerly called rennin) and lipase in
infancy and pepsinogen throughout life. They dominate the
lower half of the gastric glands but are absent from cardiac
and pyloric glands.
Enteroendocrine cells, concentrated especially in the lower
end of a gland, secrete hormones and paracrine messengers
that regulate digestion. They are found in all regions of the
stomach, but are most numerous in the gastric and pyloric
glands. There are at least eight different kinds in the stomach,
each of which produces a different chemical messenger. G
cells, for example, secrete a hormone called gastrin, which
stimulates the exocrine cells of the gastric glands to secrete
acid and enzymes.

In general, the cardiac and pyloric glands secrete mainly mucus;

acid and enzyme secretion occur predominantly in the gastric
glands; and hormones are secreted throughout the stomach. Table
24.1 describes the functions of the gastric gland secretions.
It may seem that the stomach would digest itself; we can, after all,
digest tripe (animal stomachs) as readily as any other meat. The living
stomach, however, is protected from self-digestion in three ways:
1. Mucous coat. A thick, highly alkaline mucus resists the action
of acid and enzymes.
2. Tight junctions. The epithelial cells are joined by tight
junctions that prevent gastric juice from seeping between
them and digesting the connective tissue of the lamina propria
or beyond.
3. Epithelial cell replacement. In spite of these other
protections, the stomachs epithelial cells live only 3 to 6 days
and are then sloughed off into the chyme and digested with
the food. They are replaced just as rapidly, however, by the
division of stem cells in the gastric pits.
The stomach spits about 3 mL of chyme at a time into the small
intestine. Nearly all chemical digestion and nutrient absorption occur
here. To perform these roles efficiently, the small intestine must
have a large surface area exposed to the chyme. This surface area is
imparted to it by extensive folding of the mucosa, and by the great
length of the small intestine. It measures about 2.7 to 4.5 m long in
a living person, but in the cadaver, where there is no muscle tone,
it is 4 to 8 m long. The expression small intestine refers not to its
length but to its diameterabout 2.5 cm (1 in.).

Gross Anatomy
The small intestine is a coiled mass filling most of the abdominal
cavity inferior to the stomach and liver. It is divided into three regions:
the duodenum, jejunum, and ileum (fig. 24.14).
The duodenum (dew-ODD-eh-num or DEW-oh-DEE-num)
constitutes the first 25 cm (10 in.). It begins at the pyloric sphincter,
arcs around the head of the pancreas and passes to the left, and ends
at a sharp bend called the duodenojejunal flexure. Its name refers
to its length, about equal to the width of 12 fingers.19 The first 2 cm
of the duodenum is intraperitoneal, but the rest is retroperitoneal,
along with the pancreas.
Internally, the duodenum exhibits transverse to spiral ridges,
up to 10 mm high, called circular folds (plicae circulares) (see fig.
24.20). They cause the chyme to flow on a spiral path along the mucosa,
slowing its progress, causing more contact with the mucosa, and
promoting thorough mixing, digestion, and nutrient absorption.
Adjacent to the head of the pancreas, the duodenal wall has
a prominent wrinkle called the major duodenal papilla where
the bile and pancreatic ducts open into the intestine. This papilla
marks the boundary between the foregut and midgut. In most people,
there is a smaller minor duodenal papilla a little proximal to
this, which receives an accessory pancreatic duct.
The duodenum receives and mixes the stomach contents, pancreatic
juice, and bile. Stomach acid is neutralized here by bicarbonate

in the pancreatic juice, fats are physically broken up (emulsified)

by the bile, pepsin is inactivated by the rise in pH, and pancreatic
enzymes take over the job of chemical digestion.
The jejunum (jeh-JOO-num) is the next 2.5 m (8 ft), or by definition,
the first 40% of the small intestine beyond the duodenum.
Its name refers to the fact that early anatomists typically found it to be empty.20 The jejunum
begins in the upper left quadrant of the
abdomen but lies mostly within the umbilical region (see fig. A.6,
p. 27). It has large, tall, closely spaced circular folds. Most digestion
and nutrient absorption occur here. Its wall is relatively thick and
muscular, and it has an especially rich blood supply that gives it a
relatively red color.
The ileum21 forms the last 3.6 m (12 ft), or 60% of the postduodenal
small intestine. (The lengths given here are for the cadaver.)
The ileum occupies mainly the hypogastric region and part
of the pelvic cavity. Compared to the jejunum, its wall is thinner,
less muscular, and less vascular, and it has a paler pink color. Its
circular folds are smaller and more sparse, and are lacking from the
distal end. On the side opposite from its mesenteric attachment, the
ileum has prominent lymphatic nodules in clusters called Peyer22
patches, which are readily visible to the naked eye and become progressively
larger approaching the large intestine.
The end of the small intestine is the ileocecal (ILL-ee-oh-SEEcul)
junction, where the ileum joins the cecum of the large intestine.
The muscularis of the ileum is thickened at this point to form a sphincter, the ileocecal (ILLee-oh-SEE-cul) valve, which protrudes
into the cecum and regulates the passage of food residue into
the large intestine. Both the jejunum and ileum are intraperitoneal
and thus covered with a serosa, which is continuous with the complex,
folded mesentery that suspends the small intestine from the
dorsal abdominal wall.
Microscopic Anatomy
The tissue layers of the small intestine are reminiscent of those in
the esophagus and stomach with modifications appropriate for nutrient
digestion and absorption. The lumen is lined with simple
columnar epithelium. The muscularis externa is notable for a thick
inner circular layer and a thinner outer longitudinal layer.
Effective digestion and nutrient absorption require that the
small intestine have a large internal surface area. This is provided
by its relatively great length and by three kinds of internal folds or
projections: the circular folds, villi, and microvilli. If the mucosa of
the small intestine were smooth, like the inside of a hose, it would
have a surface area of about 0.3 to 0.5 m2, but with these surface
elaborations, its actual surface area is about 200 m2clearly a great
advantage for nutrient absorption. The circular folds increase the
surface area by a factor of 2 to 3; the villi by a factor of 10; and the
microvilli by a factor of 20.
The largest of these elaborations, the circular folds, were described
earlier. They occur from the duodenum to the middle of
the ileum. They involve only the mucosa and submucosa; they are
not visible on the external surface, which is smooth.

If the mucosa is examined more closely, it appears fuzzy, like

a terrycloth towel. This is due to the villi (VIL-eye; singular, villus),
tongue- to finger-shaped projections about 0.5 to 1.0 mm
high (fig. 24.15). The villi are largest in the duodenum and become
progressively smaller in more distal regions of the small intestine.
A villus is covered with two kinds of epithelial cellscolumnar absorptive
cells and mucus-secreting goblet cells. Like epithelial cells
of the stomach, those of the small intestine are joined by tight junctions
that prevent digestive enzymes from seeping between them.
The core of a villus is filled with areolar tissue of the lamina
propria. Embedded in this tissue are an arteriole, a bed of blood
capillaries, a venule, and a lymphatic capillary called a lacteal
(LAC-tee-ul) (fig. 24.15c). Blood capillaries of the villus absorb
most nutrients, but the lacteal absorbs most dietary lipid. The reason
for this difference is that when lipids pass through the intestinal
absorptive cells, the Golgi complex packages them into proteinand
phospholipid-coated droplets called chylomicrons, then releases
them from the base of the epithelium into the core of the
villus. Chylomicrons are too large (60 to 750 nm) to pass into the
bloodstream through the blood capillary walls, but they can pass
through the larger gaps between the cells of lymphatic capillaries
and thus enter the lymph. The lymphatic system, of course, eventually
delivers the chylomicrons to the bloodstream. The fatty lymph
in the lacteal is called chyle. It has a milky appearance for which
the lacteal is named.23 The core of the villus also has a few smooth
muscle cells that contract periodically. This enhances mixing of the
chyme in the intestinal lumen and milks lymph down the lacteal to
the larger lymphatic vessels of the submucosa.