AnP 22

Chapter 22:
The
Digestive
System
Figure 22.1. Although the human diet varies widely with geography and cultural norms, it
commonly contains foods representing the three major nutrient classes: carbohydrates,
proteins, and fats. Korean temple cuisine appears in the photo above. [1]
22.1 Objectives
After completing this chapter, you should be able to:
22.2 Introduction
If food is the window to the soul, then reflecting on what you had for
breakfast this morning (you did eat breakfast, didn’t you?) can often be
an uncomfortable exercise for some of us. We voluntarily choose to eat
the foods we love because of their taste and texture, but caloric content
and health benefits must also play a role in our decision-making. Once a
meal enters our digestive system, however, the process of breaking it
down into nutrients and energy is entirely out of our control. In the video
below, take a closer look at what a brave journalist is having for
lunch—from inside his digestive system!
As the saying goes, “you are what you eat”—sort of. All food consists of
the same major nutrients (carbohydrates, proteins, and fats), but we
don’t often give credit to the complex mechanical and chemical
processes that allow them to be brought into the body. This chapter will
explore the many functions of the digestive system, and will underscore
the common themes emphasized throughout this text: communication
between cells, and homeostasis, and the idea that structure
complements function.
You will learn about the various components of the digestive system and
examine their macroscopic and microscopic structures. You will explore
how these structures are optimized to support their specific functions
and how those functions are regulated at the molecular level by the
central nervous system, the endocrine system, and the digestive system
itself.
Assigned as
Homework
Question 22.01
What is one thing that you would most like to learn about how the human digestive system
functions?
Responses
Reply
Showing
All Responses
ordered by
Newest Responses
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Aniyah White
an hour ago
I hate getting sick and I'm so sick that I'm throwing up so I want to know how the digestive system
and vomit correlate
Comments0
Alyssa Bell
an hour ago
How nutrients can absorb
Comments0
Caleb Murray
2 hours ago
How different molecules are digested
Comments0
Carl Borgstrom
17 hours ago
How our micronutrients turn helps tissues grow or atrophy.
Comments0
22.2.1 The Anatomy of the Gastrointestinal

Tract
Explore the components of the gastrointestinal (GI) tract in the diagram
below by interacting with the blue icons. Then, test your understanding in
the questions below!
Question 22.02
HomeworkUnansweredDue Oct 31st, 12:30 PM
Place the following digestive tract structures in the order they would be encountered by an ingested
food molecule.
Drag and drop options into correct order and submit. For keyboard navigation...SHOW MORE
Press space or enter to grab Ascending colon

Ascending colon
Press space or enter to grab Transverse colon

Transverse colon
Press space or enter to grab Ileum

Ileum
Press space or enter to grab Descending colon

Descending colon
Press space or enter to grab Stomach

Stomach
Press space or enter to grab Jejunum

Jejunum
Press space or enter to grab Pharynx

Pharynx
Press space or enter to grab Esophagus

Esophagus
Press space or enter to grab Cecum

Cecum
Press space or enter to grab Duodenum

Duodenum
Unanswered
Submit
Question 22.03
Match each of the following digestive tract structures with its general function.
Drag and drop options on the right-hand side and submit. For keyboard navigation...SHOW MORE
● Press space or enter to grab Major site of digestion

● Oral cavity
● Major site of digestion
● Press space or enter to grab Connects mouth to esophagus

● Duodenum
● Connects mouth to esophagus
● Press space or enter to grab Major site of water absorption

● Colon
● Major site of water absorption
● Press space or enter to grab Begins enzymatic digestion of proteins

● Esophagus
● Begins enzymatic digestion of proteins
● Press space or enter to grab Connects pharynx to stomach

● Stomach
● Connects pharynx to stomach
● Press space or enter to grab Terminal end large intestine

● Pharynx
● Terminal end large intestine
● Press space or enter to grab Vestigial immune structure

● Rectum
● Vestigial immune structure
● Press space or enter to grab Contains salivary glands

● Contains salivary glands
Unanswered
Submit
22.2.2 Accessory Organs and Structures of

the Gastrointestinal Tract
Repeat the process above, but for the organs and structures that play an
accessory role in the digestive process.
Question 22.04
Match the following accessory organs with their general function.

● Press space or enter to grab Begins digestion in the mouth

● Liver
● Begins digestion in the mouth
● Press space or enter to grab Bile storage

● Pancreas
● Bile storage
● Press space or enter to grab Carries bile into small intestine

● Salivary gland
● Carries bile into small intestine
● Press space or enter to grab Digestive enzyme production

● Gallbladder
● Digestive enzyme production
● Press space or enter to grab Carries pancreatic secretions into duodenum

● Common bile duct
● Carries pancreatic secretions into duodenum
● Press space or enter to grab Produces bile

● Produces bile
● Press space or enter to grab Aids in swallowing

● Aids in swallowing
Unanswered
Submit
22.3 Anatomical and
Functional Patterns of the
Digestive System
Imagine that you are shopping for a new car and making a comparison
between model A and model B. How would you approach this task? It
would be a waste of time to memorize every detail of every car
separately because both models have doors, a gasoline engine, a
stereo…etc. Instead, you will most likely focus on the
differences—model A has two doors, but model B is a hatchback. Model
A gets poor gas mileage, but model B is a hybrid. You can take a similar
approach when learning about the digestive system.
The complexity of the digestive system can be overwhelming to students

who use rote memorization to try to learn every detail about every region
independently. Instead, try to focus on seeing the bigger picture, on
identifying the patterns that are shared by many of the regions, and
finally on noticing the exceptions to the pattern that make each region
unique. To help get you started, this section will focus on some of the
major patterns to look out for.
22.3.1 The Digestive Tract Generally
Consists of Four Layers
There is a common pattern of overall tissue organization throughout the
gastrointestinal (GI) tract (Figure 22.2). The digestive tract is essentially
a large tube made up of four layers called tunics. The serosa is the
outermost layer, followed by the muscularis externa, then the
submucosa. The innermost layer, the mucosa, forms the lumen.
Figure 22.2. The structure of the wall of the digestive tract, including the mucosal,
submucosal, muscularis, and serosal layers, shown here in cross-section. The innermost
layer, the mucosa, consists of an epithelium, a lamina propria, and a muscularis mucosae. It
is surrounded by the submucosa, which contains the submucosal plexus (Meissner's
plexus). The mucosal and submucosal layers contain glands. The muscularis layer consists
of a circular smooth muscle layer and longitudinal smooth muscle layer. Finally, the serosal
layer is the outermost layer and is contiguous with the mesentery.
Did you know that recently, a whole new organ was discovered inside
the human body? This new organ, the mesentery, is the outermost
serosal layer and is contiguous with surrounding connective tissue. The
mesenteries anchor the digestive tract to the surrounding abdominal wall
(Figure 22.3). Blood vessels, sympathetic and parasympathetic nerves,
and lymphatic vessels branch through the mesentery to supply the
digestive tract. The mesentery also serves as a major site of visceral fat
deposition.
Figure 22.3. (a) The mesentery provides an anchor point for components of the digestive
tract, and also provides a path for nerves, lymphatic vessels, and blood vessels to service
the digestive tract. The mesentery is shown in the saggital cross-section schematic on the
left. (b) The picture at the bottom right shows one region of the mesenteries, known as the
omentum. [2]
Question 22.05
The innermost layer within the digestive tract is the ________.
Select an answer and submit. For keyboard navigation, use the up/down arrow keys to select an answer.
a
serosa
submucosa
mesenteries
mucosa
muscularis externa
Unanswered
Submit
Please select an answer to submit
The muscularis externa layer, as its name suggests, is composed largely

of muscle tissue. It consists of an outer layer of longitudinally arranged
smooth muscle (controlling length) and an inner layer of circular smooth
muscle (controlling radius). However, this layer also contains the enteric
nervous system and a structure known as the myenteric plexus. The
myenteric plexus is formed from postganglionic parasympathetic
neurons supplying cells in the esophagus, stomach, and intestines
(Figure 22.4); it regulates motility and secretion within the digestive
system. The submucosal layer, located deep inside the muscularis
externa, is composed of connective tissue that contains another layer of
nervous tissue (the submucosal plexus), as well as small circulatory
and lymphatic vessels. The submucosal plexus receives input from the
myenteric plexus, and together these two structures play a key role in
regulating digestive secretions.
Question 22.06
What is the result of damage to the myenteric plexus within the wall of the GI tract?
Gastric secretions may decrease
Pain information will be relayed through the enteric nervous system

c
Digestion will be impaired
Changes in gastric motility
Unanswered
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The innermost layer, the mucosa, is formed from three sublayers: a thin
smooth muscle layer known as the muscularis mucosae, a layer of
loose connective tissue known as the lamina propria, and an epithelial
tissue that forms the lumen. Regional specializations that allow each
region of the digestive system to have a unique function (discussed
later) are often found in this mucosal layer. For example, the lumen is
composed of stratified squamous epithelium in the mouth, oropharynx,
esophagus, and anal canal; elsewhere in the digestive tract, the lumen
consists of simple columnar epithelium.
Question 22.07
Place the tissue layers of the digestive tract in the order you would encounter them moving from the
lumen toward the abdominal cavity.

Press space or enter to grab Lamina propria

Lamina propria
Press space or enter to grab Circular layer of smooth muscle

Circular layer of smooth muscle
Press space or enter to grab Muscularis mucosa

Muscularis mucosa
Press space or enter to grab Mucosal epithelium

Mucosal epithelium
Press space or enter to grab Serosa

Serosa
Press space or enter to grab Submucosa

Submucosa
Press space or enter to grab Myenteric plexus

Myenteric plexus
Press space or enter to grab Longitudinal layer of smooth muscle

Longitudinal layer of smooth muscle
Unanswered
Submit
Why is the lumenal surface of the digestive
tract lined with epithelium?
Hover here to see the answer to Thought Question 22.01.
Figure 22.4 shows this pattern of organization throughout the GI

tract—you can see that each region has only a few specializations that
make it unique. This is a good diagram to refer back to later in the
chapter.
Figure 22.4. The wall structure along the length of the GI tract is specialized for different
functions. For example, the rugae and villi may be present or absent in different parts of the
GI tract. The diagram on the left depicts the different layers of the GI tract, beginning with
the mucosa on top in direct contact with the lumen, and the outermost layer, the serosa, at
the bottom. The diagram on the right shows a cross-section of different parts of digestive
tract, including the esophagus, stomach, duodenum, jejunum, ileum, and colon. Notable
features unique to the layers in specific areas of the tract are highlighted, such as Peyer's
patches in the ileum and gastric glands in the stomach.
22.3.2 Gastrointestinal Motility is a Major

Digestive Process
We are most aware of motility in the digestive system when we vomit.
Interestingly, powerful contractions of the stomach are capable of
projecting vomit several feet. Under normal circumstances, these
movements are much more subtle and often go unnoticed. Motility is due
to smooth muscle contraction. It occurs throughout the digestive tract
and carries ingested food bolus on a one-way journey from the
esophagus to the rectum (Figure 22.5).
Figure 22.5. Peristalsis drives the movement of food through the digestive tract, as pictured
on the left. The close-up image on the right illustrates the co-ordinated muscle contraction
and relaxation that enables the forward movement of a bolus of food.
The arrangement of smooth muscle in the digestive tract allows for two
different patterns of contraction—peristalsis and segmentation.
Peristalsis creates wave-like movements through the coordinated
contraction of circular smooth muscle behind a bolus and relaxation of
circular smooth muscle in front of the bolus. This is followed by the
contraction of longitudinal smooth muscle, which sends the bolus
forward and gives the process an overall appearance resembling an
inchworm. Peristaltic waves typically travel only short distances.
Figure 22.6. In this animation, peristalsis moves
the food bolus in the direction of the arrow along the digestive tract.
In contrast to peristalsis, segmentation is initiated by the simultaneous

contraction of circular smooth muscle in front of and behind the bolus,
creating a back and forth motion (much like kneading bread dough).
Segmentation contractions mix the bolus with digestive secretions.
Figure 22.7. Segmentation
causes the mixing of the contents of the GI tract with digestive secretions. This is illustrated
in the three images above, which show how the exclusively red or white bolus (left)
becomes mixed (middle and right). This movement is much like a baker kneading dough
with their hands.
Question 22.08
What type of contraction pushes food through the digestive tract?
a
Segmentation
Peristalsis
Sphincter
Circular
Unanswered
Submit
22.3.3 Digestion Can Be Mechanical or

Chemical
Did you know that the stain-fighting enzymes in laundry soap are very
similar to the ones produced inside your digestive system? Most
nutrients are ingested in an unusable form and need to be broken down
before being absorbed because they are too large to cross selectively
permeable membranes. The process of breaking down food molecules is
known as digestion. Mechanical digestion, which begins in the oral
cavity, does not break chemical bonds, but simply helps to increase
surface area. The increase in surface area increases the efficiency of
enzymatic or chemical digestion.
Although chemical digestion also begins in the oral cavity, it doesn’t

reach its peak until the stomach and small intestine. Chemical digestion
is carried out by many different enzymes (the names of which typically
end in the suffix -ase), each capable of working on a specific class of
nutrients. For example, complex carbohydrates are broken down into
disaccharides and simple sugars by carbohydrases; DNA and RNA a
broken down into nucleic acids by nucleases; proteins into peptides and
amino acids are broken down by proteases; and lipids are broken down
into free fatty acids and glycerol by lipases. A person’s ability to digest
food into nutrients is limited by the production of these enzymes. For
example, humans, unlike termites, cannot digest wood (or the paraffin
wax coating that makes our apples shiny in the grocery store).
22.3.4 Secretion and Absorption Are the

Last of the Major Digestive Processes
Secretion and absorption are very specific terms used to describe the
movement of substances into and out of the lumen of the digestive tract.
Secretion refers to the movement of a substance into the lumen. As you
will see, secretion is often tailored to the function of a particular region of
the digestive tract. For example, hydrochloric acid (HCI) is secreted in
the stomach to start the digestive process, whereas bicarbonate ions
(HCO3-) are secreted in the small intestine to protect the tissue from
stomach acid. Absorption refers to the movement of a substance out of
the lumen and into the cells. Like secretion, absorption is a highly
selective process that often allows specific molecules to be absorbed in
specific regions of the digestive tract. Molecules that are not absorbed
never leave the digestive tract and are removed from the body as waste.
The average person secretes up to 7 liters of water, electrolytes,
enzymes, and other molecules into the lumen each day, although the
total amount of water lost through feces is less than 500 ml—this means
that the vast majority of water is reabsorbed.
Question 22.09
Acid formation in the stomach is a result of ________.
absorption
b
secretion
peristaltic contractions
segmentation contractions
Unanswered
Submit
Lecture Video - Patterns of

the Digestive System
22.4 Regional
Specialization of the Oral
Cavity
22.4.1 Digestion Begins in the Oral Cavity
As humans, we are most familiar with the oral cavity because of our love
for food and speech. Generally speaking, the oral cavity serves the
following functions:
● Protects the mouth and digestive system against physical and

chemical abrasions, and food-borne pathogens (e.g., yeast,
bacteria)
● Increases the surface area of food
● Coats food with saliva
● Initiates the process of swallowing, which delivers food to the
stomach
The oral cavity (Figure 22.8) is the region where food is brought into the
body and the process of digestion begins. The tongue moves food within
the mouth and positions it for grinding by the teeth. The tongue's surface
is covered with 2,000-10,000 sensory or taste buds that provide
information about flavor (see Chapter 13: Specialized Senses).

Figure 22.8. The oral cavity ingests food, mixes it with saliva, and begins the digestion
process. The oral cavity is pictured on the left, with labels indicating the upper and lower
lips, the hard and soft palates, the uvula, tonsils, tongue, and gums. The picture on the right
depicts the lingual frenulum, which extends from the floor of the mouth to the midline of the
underside of the tongue.

There are 3 muscles used to close the jaw
but only 1 muscle used to open it, why?
Hover here to see the answer to Thought Question 22.02
Type caption for image (optional
22.4.2 Mastication Mechanically Digests

Food
Mastication, commonly known as chewing, breaks up large food
particles into smaller particles. This mechanical digestion increases the
surface area of food and mixes food with saliva. Mastication is
performed by the teeth. Chewing is a repetitive task carried out under
the control of the medulla oblongata. The presence of food within the
oral cavity stimulates sensory cells that send information to the medulla
and elicit relaxation of the mastication muscles. The muscles are
stretched as the mandible is lowered, which, in turn, activates sensory
cells that initiate a separate reflex to cause contraction of the mastication
muscles. Once the mouth is closed, the cycle is repeated. The
mastication reflex can be modified in rate and intensity, or stopped
altogether, by descending pathways from the cerebrum.
22.4.3 Saliva Secretion in the Oral Cavity

Saliva is secreted into the oral cavity by three glands: the parotid gland,
the submandibular gland, and the sublingual gland. Collectively, these
are known as the salivary glands. Saliva is a combination of thin serous
and thick mucus secretions composed of water, electrolytes, mucous,
leukocytes, epithelial cells, glycoproteins, enzymes, IgA, and lysozymes
(Figure 22.11). Its pH is between 6.5 and 7.5. Because it is mostly water,
saliva moistens epithelia, liquefies food, and begins the process of
enzymatic digestion of complex carbohydrates into disaccharides. This
chemical digestion of carbohydrates is carried out by the enzyme
salivary amylase, a component of serous saliva. The maltose that is
produced from this chemical reaction often causes food that lingers
inside the mouth to develop a “sweet” taste.
Figure 22.11. The mouth has several salivary glands that secrete saliva, aiding in the
ingestion and digestion of food. These glands include the parotid gland, the submandibular
gland, and the sublingual gland, as seen on the left. The cross-section on the right shows
the organization of a salivary gland consisting of acini and ducts, where primary secretions
(ptyalin, mucus, and extracellular fluid) from acinar cells, together with secondary secretion
modifications in ducts, produce saliva.
Because most of us eat our food in a hurry, there is only time for about
5% of the polysaccharides to be broken down by saliva. Additionally, not
all complex carbohydrates (e.g., cellulose in plants) are accessible to
salivary amylase unless they have been thoroughly cooked or chewed.
Saliva is also a component of the innate immune response (see Chapter
21: The Immune System). It helps to prevent oral bacterial infections by
removing bacteria from the oral mucosa. Lysozyme in saliva digests
bacterial cell walls, and antibodies to help protect the mouth against
bacterial infection.
A lack of saliva increases the risk of cavities and ulceration infections of

the oral mucosa. These anti-microbial properties are also the reason
why we often see injured pets licking their wounds! Both the
parasympathetic and sympathetic branches of the autonomic nervous
system stimulate salivary secretion, although the parasympathetic
system has a greater effect. Saliva secretion is stimulated through the
facial (VII) and glossopharyngeal (IX) cranial nerves in response to a
variety of stimuli, including tactile stimulation of the oral cavity, sour
tastes (low pH), and sensory input to higher brain areas (e.g., the sight
or smell of food).
22.4.4 Swallowing
Open your mouth and say “Ahhhhhhh.” When staring into the back of
your open mouth in the mirror, you can see behind the uvula (the
dangling soft palate structure at the back of your mouth) to the pharynx.
The pharynx is a tube that connects the inner ear, oral cavity, and larynx
(Figure 22.12). The upper portion is the nasopharynx, the middle portion
is the oropharynx, and the bottom portion is the laryngopharynx. The
pharynx provides a passageway for the movement of food, liquid, and air
to move into the esophagus and trachea, respectively. Swallowing, or
deglutition, is separated into three phases (Figure 22.13): voluntary,
pharyngeal, and esophageal.
During the initial voluntary phase, the mouth, tongue, teeth, and
secretions form a food bolus (Figure 22.13). The tongue pushes the food
bolus against the hard palate and toward the back of the oral cavity into
the oropharynx, where it is detected by receptors within the pharyngeal
wall. Parents who are trying to get small children to swallow pills will
often place the pill far back on the tongue, near the larynx (making this
voluntary phase a bit less voluntary).
Figure 22.12. An overview of the major structures of the oral cavity, used by both the
digestive and respiratory systems.The nasopharynx is seen in blue and contains the
opening to the auditory tube. The oropharynx is in green between the uvula and epiglottis.
The laryngopharynx is seen in pink.
In the pharyngeal phase, the contact of food components with sensory

receptors in the posterior wall of the oropharynx sends afferent
information to the swallowing center in the medulla. In response, motor
information travels to the soft palate and pharynx. This stimulates soft
palate elevation while closing the passageway between the nasopharynx
and oropharynx. Medial movement of the vestibular folds and anterior
movement of the epiglottis position the epiglottic cartilage to cover the
opening to the larynx, allowing food to safely travel into the esophagus
when the upper esophageal sphincter relaxes. Occasionally, food
travels “down the wrong pipe,” and water or food may enter the trachea.
There are sensory receptors in the wall of the upper trachea that
stimulate violent coughing when food enters. Despite being a skeletal
muscle sphincter, the relaxation of the upper esophageal sphincter is
involuntary (this is one of the rare cases where skeletal muscle is not
controlled voluntarily). Although it may sound very complex, the entire
pharyngeal phase lasts just over a second.
Figure 22.13. In the initial voluntary part of the swallowing reflex, the food bolus is pushed
by the tongue against the hard and soft palates and the oropharynx. During the pharyngeal
phase, the soft palate elevates, closing off the nasopharynx, and the pharynx is elevates
(left). Constriction of pharyngeal muscles forces the bolus from the pharynx into the
esophagus. The epiglottis is bent down over the larynx opening (middle). During the
esophageal phase, the upper esophageal sphincter relaxes, allowing the food bolus to enter
the esophagus and travel to the stomach (right). The food bolus is moved forward by
peristaltic contractions.
Swallowing terminates with the esophageal phase. The esophagus

provides a passageway for the movement of food and water from the
pharynx to the stomach (Figure 22.14). Recall Chapter 19: The
Circulation and Short-Term Bloom Pressure, where we discussed how
moving anything through a tube requires a pressure gradient? In the
digestive system, this pressure required to move the food bolus is
created by peristaltic muscle contractions.
Figure 22.14. The esophagus connects the oral cavity to the stomach, as shown above.
The following video shows an X-ray of a person swallowing some water.

As mentioned above, peristaltic muscle contractions facilitate the
movement of the water down the esophagus.
In contrast to other areas of the digestive tract, the upper esophagus has
an outer longitudinal and inner circular skeletal muscle layer (Figure
22.14). The lower esophagus is composed entirely of smooth muscle.
Therefore, the upper skeletal muscle is consciously controlled to initiate
swallowing, whereas the lower smooth muscle is unconsciously
(involuntarily) controlled during the later stages of swallowing.
Question 22.14
During which phase of swallowing does food enter the stomach?
a
Voluntary phase
Pharyngeal phase
Esophageal phase
Unanswered
Submit
As food travels through the esophagus, it stimulates stretch receptors

that signal through the enteric nerve plexus to smooth muscle, which
stimulates contraction and forms peristaltic waves. In turn, this activates
efferent neurons that stimulate the contraction of esophageal smooth
muscle to reinforce peristaltic contractions. If you’ve ever had a peanut
butter sandwich, then you know that this is particularly useful when
something gets “stuck” in your throat and a secondary peristaltic wave is
necessary.
Though not recommended, you can eat and drink upside down. While
swallowing is normally aided by gravity, peristaltic waves are strong
enough to work against it. In fact, a key part of the training of future
astronauts is to help them become comfortable eating without the help of
gravity! However, many people have a difficult time swallowing while
lying on their backs and prefer to elevate their head when swallowing. As
food approaches the stomach, the lower esophageal sphincter (smooth
muscle) relaxes, allowing the food bolus into the stomach.
When the lower esophageal sphincter doesn't close properly, acidic

secretions from the stomach can reflux (leak back) into the esophagus
and cause inflammation. This can lead to a feeling of heartburn in the
patient. This heartburn, in turn, can lead to gastroesophageal reflux
disease (GERD) (Figure 22.15). If left untreated, GERD can lead to very
serious complications, including ulceration of the esophagus wall,
leading to massive blood loss and esophageal cancer. Treatment
includes changing the diet (food type, amount, and frequency),
medication, and in serious cases, surgery.
Figure 22.15. (a) GERD refers to the backflow (reflux) of acidic stomach contents into the
esophagus, as seen in the enlarged schematic (top right). (b) An x-ray of the chest and
abdomen of a patient with severe GERD, which shows barium contrast moving upward from
the stomach into the esophagus. (c) An endoscopic image of peptic stricture, or narrowing
of the esophagus near the junction with the stomach, which is a common complication of
chronic GERD and can cause difficulty swallowing by creating a high pressure environment
in the lower esophagus. [4]

Lecture Video - Oral Cavity
22.5 The Stomach
The next stop for ingested food is the stomach, which is an essential part
of the human digestive system that has several unique functions:
● To act as a short-term storage reservoir, allowing a large meal to

be ingested quickly but digested over an extended period of time.
This is especially true for protein-rich meals.
● To mix and grind stomach contents with HCl and the enzyme
pepsin.
● To continue chemical and enzymatic digestion, particularly of
proteins.
● To move stomach contents (chyme) into the small intestine for
further processing.
As a continuation of the digestive tract, the stomach is a hollow organ

that connects to the esophagus in the cardiac region (because of its
closeness to the heart) (Figure 22.16). Inferior and medial to the cardiac
region is the body, the largest part of the stomach. The body and
fundus form the greater and lesser curvatures. The body narrows to
form the pylorus, which joins with the duodenum of the small intestine.
The opening between the pylorus and the small intestine is the pyloric
canal, which is surrounded by a thick ring of smooth muscle known as
the pyloric sphincter. The stomach’s specialized walls contain large
folds known as rugae that disappear with distension (stretching). This
allows the stomach to increase in volume to accommodate a large meal.
Figure 22.16. The stomach acts as a food reservoir and begins the chemical and enzymatic
digestion of food. The cardia is the region of the stomach that connects to the esophagus,
the body, and fundus from the greater and lesser curvatures; the pylorus joins the stomach
with the duodenum. As seen in the cross-sections above, the stomach consists of three
muscle layers: a longitudinal smooth muscle layer, a circular smooth muscle layer, and an
oblique muscle layer. The stomach wall also contains rugae, which are large folds that
accommodate the increase in volume that accompanies large meals.
The stomach wall also contains an additional smooth muscle layer

known as the inner oblique layer, which increases the strength of its
contractions (which are felt when we vomit). The mucosal layer is also
specialized to form invaginations known as gastric pits. Gastric pits are
made of simple squamous epithelium and are the major secretory
structures of the stomach (Figure 22.17).
Why is the stomach one of the few places

within the digestive tract that has 3 layers of
smooth muscle within its wall?
22.5.1 Secretion in the Stomach: The Gastric

Pit
The enzymatic digestion of food begins in the stomach, converting the
food bolus into chyme, an acidic fluid containing partially digested food.
Pepsins, the major proteases active in the stomach, are synthesized and
secreted predominantly by chief cells in the gastric pit (Figure 22.17).
Although there are at least 8 different forms of pepsin produced in the
human stomach, all are synthesized as inactive proenzymes (see
Chapter 16: The Endocrine System) consisting of a signal peptide, an
activation peptide, and an enzyme.
Figure 22.17. A cross-sectional of the stomach wall is shown on the left and consists of the
surface epithelium, mucosa, submucosa muscularis externa (oblique, circular, and
longitudinal smooth muscle layers), and serosa. Gastric pits within the lining of the stomach
(shown on the right) are openings for cells that secrete mucus, acid, enzymes, and
hormones. Parietal cells, chief cells, G cells that secrete gastrin, and D cells that secrete
somatostatin, can be found in gastric pits.
The signal peptide is cleaved into a proenzyme, pepsinogen, that is

transported to the Golgi and condensed into secretory granules, which
are then released into the lumen. Pepsinogens are activated when they
become exposed to the acidic conditions of the stomach lumen. The
altered pepsinogen is digested, and the resulting protein fragment is an
active peptidase, pepsin. Pepsins are endopeptidases, which recognize
amino acid sequences embedded within proteins. Exopeptidases, which
will be covered in Section 22.9.1, cleave peptides from the ends of
proteins. Chief cells contain receptors for several hormones that
stimulate pepsin release, including secretin, vasoactive intestinal
peptide, and epinephrine.
Question 22.16
Which of the following are produced in the gastric pit?
Mucus
b
Chyme
Alkaline buffer
Glucose
Unanswered
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Because pepsinogen requires an acidic pH for activation, protein

digestion is ultimately dependent on hydrochloric acid (HCl) secretion.
HCl is secreted by parietal cells (Figure 22.18) which gives the lumen a
pH of approximately 0.8. HCl at this concentration is capable of etching
cement. The acid is secreted into large canaliculi, which are deep
invaginations of the apical parietal cell membrane that are continuous
with the lumen of the stomach. Following stimulation, there is a dramatic
change in the parietal cell membrane, characterized by a large increase
in canaliculi.
Figure 22.18. (a) A microscopic view of parietal cells. (b) A schematic representation of
parietal cell morphology, with intracellular canaliculi on the apical membrane continuous
with the stomach lumen. [5]
Remember that important chemical reaction that takes place inside red
blood cells and creates the buffer system in the blood? It happens inside
parietal cells as well! Parietal cells use the enzyme carbonic anhydrase
to combine CO2 and H2O—which are abundant inside the cell—into
carbonic acid (H2CO3). Dissociation of carbonic acid yields H+ and HCO3-
. The H+ concentration in the lumen of the stomach is roughly 3 million
times greater than in the blood. Given this tremendous concentration
gradient, parietal cells must secrete H+ via active transport using the
H+/K+-ATPase (“proton pump”) located on their apical (luminal)
membrane. H+ is pumped out of the parietal cells into the lumen in
exchange for K+, which is recycled back into the parietal cells.
Bicarbonate (HCO3-) is transported out of the basolateral (blood side)
membrane in exchange for Cl-. The Cl- and K+ ions are transported into
the lumen of the canaliculus by conductance channels on the apical
membrane. This system of acid secretion allows for the formation of HCl
in the stomach lumen and prevents damage to the parietal cells that
would occur if HCl were formed in the cytoplasm. Interestingly, as the
lumen of the stomach becomes more acidic, the outflow of HCO3– into
the blood results in a slight increase of blood pH, which is known as the
alkaline tide.
The regulation of acid secretion is complex but critically important. To aid

in your understanding, review earlier portions of this textbook that
discuss hormones, neurotransmitters. and transport processes across
the membrane. Parietal cells are regulated locally by paracrine
mechanisms, and globally by hormones and the central nervous system.
The local release of histamine from enterochromaffin (ECL) cells
within the stomach stimulates acid secretion, but the magnitude of the
stimulus results from a complex interaction with other signals. Parietal
cells may also be stimulated by gastrin, a hormone released from G
cells in response to distension of the stomach by food. Gastrin binds to
receptors on the basolateral membrane of parietal cells, leading to
elevations in intracellular Ca2+, activation of Ca2+-dependent kinases, and
increased transport of the H+/K+-ATPase to the cell membrane.
Interestingly, gastrin is also capable of stimulating enterochromaffin cells
to release histamine, and it can also stimulate chief cells to release
pepsinogen. Gastrin is, therefore, a potent regulator of acid formation in
the stomach.
Figure 22.19. Hydrochloric acid (HCl) is produced within parietal cells in the gastric pits of
the stomach and is transported across the cell membrane by a H+/K+-ATPase, as pictured
above. Parietal cell secretion is regulated by surrounding cells in the gastric mucosa that
release factors such as gastrin, histamine and acetylcholine.
Gastric secretions are also regulated by the parasympathetic nervous

system through the vagus nerve. The binding of acetylcholine to its
receptors results in the secretion of acid from parietal cells (via
Ca2+-dependent kinases), pepsin from chief cells, and gastrin and
histamine from endocrine cells. This results in additional gastrin being
released into the circulation and traveling to parietal and chief cells, thus
stimulating additional HCl and pepsinogen release. The synergistic effect
of acetylcholine, histamine, and gastrin on parietal cells results in greater
HCl secretion than any agent is capable of producing on its own. A
variety of substances are capable of reducing gastric acid secretion,
including hormones like secretin and gastric inhibitory peptide (from the
small intestine), glucagon (from the pancreas), and somatostatin (from
D cells in the gastric pit). Somatostatin inhibits gastrin and histamine
secretion and also appears to have a direct inhibitory effect on parietal
cells.
The stomach epithelium is intrinsically resistant to the damaging effects

of gastric acid and pepsin because of a mucous layer that is formed by
the mucus-secreting epithelium. Excessive secretion of gastric acid can
eventually damage the stomach wall, leading to gastritis, gastric ulcers,
and peptic acid disease. If excessive acid secretion occurs where there
is an infection of the stomach wall by H. pylori, the damage to the wall is
increased.
How does excessive coffee consumption

(who's to say what's excessive?!) affect the
stomach?
Gastric pits also secrete a critical intrinsic factor that is unrelated to acid
secretion. Intrinsic factor is a glycoprotein secreted by parietal cells that
plays an important role in the absorption of vitamin B12 in the intestine.
Failure to produce or utilize intrinsic factor results in pernicious anemia
(see Chapter 17: Blood). Dietary vitamin B12 is released from ingested
proteins in the stomach through the actions of pepsin and acid. It is
rapidly bound by one of two vitamin B12-binding proteins that are present
in gastric juice. At an acidic pH, these binding proteins have a greater
affinity for vitamin B12 than intrinsic factor does. In the small intestine,
pancreatic proteases digest the binding proteins, releasing vitamin B12,
which then becomes bound to intrinsic factor. There are receptors for
intrinsic factor on mucosal cells that bind the complex, allowing vitamin
B12 to be absorbed into the portal blood. Because the efficient absorption
of vitamin B12 in humans depends on intrinsic factor, diseases that
decrease the secretion of intrinsic factor, interfere with cleavage of the
binding proteins, or decrease binding and absorption of the intrinsic
factor-vitamin B12 complex, can result in anemia.
Because of all the crosstalk, secretions from the gastric pit can be a bit
daunting to learn about for the first time. Grab a blank sheet of paper,
and let’s review them in the Lightboard video below.
Question 22.18
Match each cell type with the molecule it releases.
● Press space or enter to grab Somatostain

● G cell
● Somatostain
● Press space or enter to grab Histamine

● Parietal cell
● Histamine
● Press space or enter to grab Acetylcholine

● Enterochomaffin (ECL) cell
● Acetylcholine
● Press space or enter to grab Gastrin

● D cell
● Gastrin
● Press space or enter to grab Hydrochloric acid

● Vagal neuron
● Hydrochloric acid
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A gastric, or duodenal ulcer, is a break in the normal tissue lining the

stomach or small intestine (Figure 22.20). Benign gastric ulcers are
caused by an imbalance between the defences provided by the stomach
mucosal lining and the secretion of acid and pepsin. Gastric ulcers are
caused by inhibition of mucous secretion, resulting tissue damage. A
number of risk factors are associated with gastric ulcers, including
chronic inflammation, excessive use of aspirin and nonsteroidal
anti-inflammatory medications, and smoking. However, nearly 90% of all
cases are attributed to bacterial infection with H. pylori. Contrary to
popular belief, stress and spicy foods do not cause, or worsen, gastric
ulcers.
Symptoms associated with ulcers include abdominal pain, nausea,

indigestion, vomiting blood, blood in stools, and weight loss. Some
symptoms are alleviated by antacid medications (which react with HCl to
form salts in a neutralization reaction). These medications are often
taken in combination with antibiotics and an H2 receptor antagonist or a
proton pump inhibitor to suppress acid production. These medications
are commonly used in conjunction with dietary changes, such as
avoiding alcohol, caffeine-rich drinks, fried food, lemonade or grapefruit
juice, and spicy or pickled foods. Patients are encouraged to drink water,
low-caffeine teas, or low acid fruit juice, and to eat complex starches,
fruit, fresh and cooked vegetables, low-fat meat substitutes, and nuts.
Figure 22.20. Gastric ulcers are erosions in the mucosal layer of the stomach (left). On the
right is an endoscopic photo of a large gastric ulcer. [6]
Understanding the mechanisms involved in stomach acid secretion has

led to the development of several drugs capable of inhibiting acid
secretion. The most effective inhibitors fall into two classes: H2 receptor
agonists and proton pump inhibitors. The parietal cell receptor for
histamine is of the H2 type, and antihistamines that bind to the H1
receptor have no effect on acid secretion. Evidence of histamine's role in
acid secretion is strongly supported by the finding that H2 receptor
antagonists are quite effective in inhibiting acid secretion. Four primary
H2 antagonists have been developed and are used clinically: cimetidine
(Tagamet™), ranitidine (Zantac™), famotidine, and nizatidine. These
drugs, particularly cimetidine, are among the most widely prescribed
drugs in the United States. Acid secretion is dependent on the activity of
H+/K+-ATPase located in the parietal canalicular membrane. Several
drugs have been developed that bind and inactivate this ATPase,
resulting in strong inhibition of acid secretion. Omeprazole (Prilosec™) is
an acid-activated drug that binds covalently to two cysteines on the
H+/K+-ATPase, resulting in its irreversible inactivation. Two other
inhibitors, lansoprazole and pantoprazole, have similar modes of action.
22.5.2 Stomach Motility

Segmentation contractions in the stomach mix the food bolus with
stomach secretions to convert the bolus into chyme. Peristaltic
contractions move the chyme toward the pyloric sphincter of the
stomach and small intestine, where the chyme is mixed with pancreatic
and small intestinal secretions. As food moves into the stomach, the
rapid expansion of the stomach walls prevent increases in stomach
pressure until the volume nears maximum capacity. This allows the
upper stomach to act as a storage chamber during the digestion of
protein-rich meals. Mixing waves, which begin at the stomach body and
move toward the antrum, push chyme against the pyloric sphincter.
Roughly 80% of gastric contractions are mixing waves; the rest are
stronger peristaltic waves. The volume and composition of chyme
regulates how long it is stored within the stomach. A typical meal
containing fats, proteins, and carbohydrates leaves the stomach within
four hours. A fatty, protein-rich meal is cleared more slowly. While the
pyloric sphincter is usually closed, stomach distension stimulates the
release of gastrin, which increases the formation of peristaltic waves and
relaxes the sphincter to create a path to the duodenum (small
intestines). Peristaltic contractions are sufficiently strong to push chyme
out of the stomach into the small intestine. The presence of food within
the duodenum, in turn, will slow the process of gastric emptying. We will
talk more about this in Section 22.6.
Question 22.19
Which of the following might be a consequence of bariatric surgery, where a “band” is placed around
the stomach in order to reduce its volume?

The stomach should behave in the same way.
Meals must be consumed more frequently because digestion occurs more rapidly.
c
Digestion would be decreased and fewer nutrients would be absorbed.
The stomach will no longer be capable of peristaltic contractions.
Unanswered
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22.5.3 Vomiting
All humans have experienced vomiting—the involuntary, forceful
expulsion of stomach contents through the mouth, which acts to remove
potentially harmful materials from the stomach and small intestine.
Vomiting is initiated by receptors in an area of the fourth ventricle of the

brain known as the chemoreceptor trigger zone. Vomiting is caused by
the coordinated contraction of abdominal muscles, the diaphragm, and
the stomach. This coordinated contraction results in tremendous
pressure built up inside the stomach, which provides the force to expel
stomach contents. The contents of the small intestine can also be part of
vomit with the relaxation of the pyloric sphincter. Vomiting can be
induced by emetics, and can be blocked by antiemetics that target the
parasympathetic nervous system, as well as by antihistamines and
cannabinoids. The antiemetic properties of cannabinoids are one of the
strongest arguments in favor of medicinal marijuana.
Question 22.20
Which of the following is most rapidly cleared from the stomach after eating?
Fats
Proteins
Carbohydrates
Unanswered
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A 43-year-old man presents to the emergency room one clear Tuesday
morning complaining of severe pain in his epigastric and umbilical
regions. He describes the pain as a burning sensation that began as he
was sitting in rush hour traffic. He was concerned that he may be having
a heart attack, so he got to the hospital as fast as he could. Upon further
investigation, his physician discovers that this is not the first time he has
experienced this pain. He has frequent nausea and occasional vomiting,
but notices that the pain is worse between meals. He has a high-stress
job and takes antacids regularly for heartburn. He notes that during the
last month, he has been very tired and has had dark-colored stools.
Diagnosis: Peptic Ulcer
Hover here to find out how this condition would be treated.
Lecture Video - Stomach

22.6 The Small Intestine
The small intestine is a continuation of the stomach and is one of the
longest structures in the human body—stretched out it would likely be at
least three times as tall as you!
The major functions of the small intestine are:
● To mix liver and pancreatic secretions with chyme.

● To continue digesting carbohydrates and proteins, and to initiate
fat digestion.
● To absorb nutrients.
● To move chyme toward the large intestine.
● To produce hormones that regulate the digestive system.
● To produce large numbers of immune cells that protect against
pathogens associated with food and the microbiome.
Until now, we have discussed how digestion begins in the oral cavity and
accelerates in the stomach. However, 90% of all digestion occurs in the
small intestine. In fact, the small intestine is often described as the
major site of digestion. Very few compounds are absorbed in the
stomach, so it is also a major site of absorption. Each day, approximately
9 L of liquid enters the small intestine from ingested food and secretions,
but less than a liter enters the colon.
Explore the three parts of the small intestine by interacting with the 3D
model below.
Activate
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The small intestine consists of three parts: the duodenum, the jejunum,
and the ileum. The duodenum is the shortest segment (4%), followed by
the jejunum (40%), and the ileum (56%). Although the small intestine
produces its own secretions, it also receives secretions from accessory
organs in the abdominal cavity. The duodenal papillae are located
about two-thirds of the way down the duodenum. It is here that the
common bile duct and pancreatic duct join with the
hepatopancreatic ampulla. This allows the small intestine to receive
secretions from the liver, pancreas, and gallbladder.
Figure 22.21. A hematoxylin
and eosin (H&E)-stained section showing plicae within the wall of the small intestine that
increase its surface area by 600-fold, accelerating absorption. [7]
The intestinal wall has several modifications that increase its surface
area approximately 600-fold (Figure 22.21). The mucosal and
submucosal layers form a series of large projections called plicae
(Figure 22.21). The mucosal layer forms numerous villi (finger-like
projections), which are 0.5 to 1.5 mm in length (Figure 22.22). Inferior to
each villus is a blood and lymphatic capillary network known as a
lacteal. The luminal surface of the columnar epithelium is covered with
cytoplasmic extensions called microvilli, which further increase the
surface area. The microvilli form the brush border (so-named because
it resembles a fine brush), which is a major site of nutrient digestion and
absorption. Villi formation is increased by contraction of the mucosal
layer.
Figure 22.22. Villi and microvilli on plicae dramatically increase the surface area of the small
intestine, accelerating absorption, as seen in the series of cross-sections above of the small
intestine (far left), plicae, villi, and microvilli (far right). Each lacteal is surrounded by a
capillary bed and lymphatic vessel, providing a rapid means to absorb nutrients.
Question 22.22
Rank the following structures in order from smallest to largest.
Press space or enter to grab Microvilli
Microvilli
Press space or enter to grab Villi

Villi
Press space or enter to grab Plicae

Plicae
Unanswered
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Why is the small intestine so long?

Intestinal epithelia lining the lumen of the small intestine are one of four
different cell types: absorptive cells that have extensive microvilli and
produce digestive enzymes; goblet cells that produce a protective
mucous; granular cells that may participate in the immune protection of
the small intestine; and endocrine cells that produce several different
hormones. These epithelial cells are continuously replaced as stem cells
divide within tubular invaginations of the mucosal layer known as
intestinal crypts, which are located at the base of each villus.
Absorptive and goblet cells migrate from the intestinal glands to cover
the villi surface and are eventually shed at the tip. The granular and
endocrine cells remain at the bottom of the crypts.
Question 22.23
Villi formation is due to contraction of cells in the mucosal layer of the digestive tract.
True
False
Unanswered
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The jejunum and ileum share structural similarities with the duodenum,
except that there is a gradual decrease in intestinal wall diameter and
thickness, and a decrease in the number of plicae and villi, as the
intestine progresses. The duodenum and jejunum, which have a large
surface area, are the major sites of nutrient reabsorption. This is
because reabsorption is driven by diffusion, which is proportional to
surface area. Lymph nodes, known as Peyer’s patches (Figure 22.23),
are numerous in the ileum mucosal and submucosal layers.
Figure 22.23. Peyer's patches, circled in this hematoxylin and eosin (H&E)-stained section,
are part of the immune surveillance within the small intestine. [8]
The ileum and the large intestine join at the ileocecal junction. The
ileocecal junction has a ring of smooth muscle, a sphincter (the ileocecal
sphincter), and the one-way ileocecal valve. The ileocecal valve ensures
that the contents of the large intestine do not enter the small intestine.
22.6.1 Motility in the Small Intestine

Both segmentation and peristaltic contractions occur in the small
intestine. Segmentation contractions mix intestinal contents, and
peristaltic contractions propel the chyme through the small intestine.
However, very few peristaltic contractions travel the entire length of the
small intestine. Under normal circumstances, it takes chyme 3 to 5 hours
to travel from the pyloric sphincter to the ileocecal valve. The ileocecal
sphincter is mildly contracted most of the time to slow the rate of chyme
movement and allow time for digestion and absorption to occur. When
peristaltic waves reach the ileocecal sphincter, the sphincter relaxes to
allow chyme to enter the large intestine. The distension of the cecum,
the first part of the large intestine, activates a local reflex that contracts
the ileocecal sphincter and prevents movement of chyme into the
cecum.
22.6.2 Secretion from the Small Intestine

into the Lumen
In contrast to the stomach, secretions from the small intestine do not
contain HCl or pepsin. Instead, small intestine secretions contain mucus,
electrolytes, and water. The duodenal glands, intestinal glands, and
goblet cells secrete large amounts of mucus that protect the intestinal
wall against the gastric acid entering the duodenum.
Question 22.24
Match each intestinal feature with its major function.
● Press space or enter to grab Produce the hormone secretin

● Peyer’s patch
● Produce the hormone secretin
● Press space or enter to grab Supports the immune system

● Villi
● Supports the immune system
● Press space or enter to grab Increase surface area for digestion and absorption
● Goblet cell
● Increase surface area for digestion and absorption
● Press space or enter to grab Protects the small intestine from stomach acid and
abrasion
● Endocrine cell
● Protects the small intestine from stomach acid and abrasion
● Press space or enter to grab Contains stem cells

● Intestinal crypts
● Contains stem cells
Unanswered
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Intestinal mucosal enzymes are attached to the microvilli that form the
brush border membrane. These membrane-bound enzymes include
disaccharidases, which cleave disaccharides to monosaccharides;
peptidases, which hydrolyze peptide bonds in chains of amino acids;
and nucleases, which digest nucleic acids. These membrane-bound
enzymes play an important role in digestion by taking advantage of the
increased surface area arising from the villi, which maximizes the
contact of chyme with these enzymes. The small intestine releases a
variety of hormones into the bloodstream, including cholecystokinin
(CCK), gastric inhibitory peptide (GIP), and secretin. These
hormones, known as enterogastrones, are important for regulating
other regions of the GI tract, the pancreas, and the gallbladder.
22.6.3 Secretions into the Small Intestine

from Accessory Organs
Although the intestinal mucosa produces its own secretions, liver and
pancreatic secretions also enter the digestive tract in the duodenum. The
pancreas supplies digestive enzymes and an alkaline (HCO3-) broth to
neutralize the acidic chyme delivered by the stomach. The liver and
gallbladder provide bile salts that aid in lipid digestion. We will talk more
about these accessory organs later in the chapter.
Lecture Video - Small
Intestines
22.7 The Large Intestine
The large intestine, which is the last portion of the digestive system, has
several critical roles, including:
● Absorption of water from the remaining chyme, converting the

chyme to semi-solid feces.
● Secretion of a protective mucous and facilitation of the movement
of the remaining undigested food.
● Providing a hospitable area for the growth of extensive bacterial
colonies that metabolize nutrients and steroids, produce vitamin
K, and prevent the growth of pathogenic bacteria.
● Production of large numbers of immune cells, organized into
Peyer’s patches and primary follicles, that provide protection
against pathogens.
The large intestine extends from the ileocecal sphincter to the anus
and consists of the cecum, the colon, and the rectum (Figure 22.24).
Figure 22.24. The large intestine consists of the cecum, ascending colon, transverse colon,
descending colon, and sigmoid colon, as shown above.
The cecum forms the proximal end of the large intestine and includes the
vermiform appendix. The appendix is a small, closed tube that is
studded with lymph nodules that provides immune surveillance against
foreign molecules and pathogens. The large intestine beyond the cecum
is known as the colon. The colon is two meters in length and is made up
of four parts that move up, across, and down through the peritoneal
cavity—the ascending, transverse, descending, and sigmoid colon,
respectively. The colon wall is made up of mucosal, submucosal,
muscularis, and serosal layers (like the rest of the digestive tract). The
mucosal lining of the colon consists of simple columnar epithelium;
notably, it does not form folds or villi like the small intestine. It does,
however, have tubular crypts that are similar to the crypts of the small
intestine.
Question 22.25
What can you deduce from the fact that the large intestine wall is composed of simple columnar
epithelia?
The large intestine is the major site of digestion of small peptides into single amino acids.
The large intestine is the major site of carbohydrate absorption.
Water absorption by the large intestine does not require plicae.

d
The large intestine is the major site of acid and enzyme secretion.
Unanswered
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Question 22.26
Assigned as
Homework
Question 22.26
Why does the large intestine not contain villi and microvilli?
Responses
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ordered by
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Click here to see the answer to Question 22.26.
Appendicitis is the inflammation of the appendix, which is found at the

base of the ascending colon. It is thought that the appendix is a vestigial
organ that played some role in immune surveillance of the large
intestine. Patients with appendicitis present with right lower abdominal
pain, nausea, vomiting, and decreased appetite. This condition arises
when the appendix becomes blocked, leading to infection. The appendix
can be blocked by a calcified fragment of feces, inflamed lymphoid
tissue, parasites, gallstones, or tumors. If left untreated, the appendix
can burst and release bacteria into the abdomen, leading to sepsis and
death. The following video shows the surgical removal of the appendix.
Not all of the colon smooth muscle layers are complete; the circular
muscle layer is complete, whereas the longitudinal muscle layer is not.
The longitudinal smooth muscle layer forms three bands (like adhesive
tape) known as the teniae coli, which run the length of the colon, giving
it a puckered, segmented appearance at the level of its gross anatomy.
Each of these segments is known as a haustrum (Figure 22.25). It
normally takes the better part of a day for chyme to pass through the
large intestine. As chyme moves through the large intestine, it is
transformed into feces through water and salt absorption, mucous
secretion, and by the actions of normal gut flora. Feces are stored in the
colon until they are eliminated by defecation. About 2 L of chyme enters
the cecum each day and more than 90% of the volume is reabsorbed,
leaving approximately 200 mL of waste to be eliminated.
Figure 22.25. The teniae coli, which is formed by three bands of longitudinal smooth
muscle, create haustra and give the large intestine a puckered appearance.
The rectum is a straight, muscular tube that extends from the sigmoid
colon to the anal canal. It is lined with simple columnar epithelium and
has a relatively thick smooth muscle layer that is required to expel fecal
material. The last few centimeters of the digestive tract, known as the
anal canal, extends from the rectum to the anus, which is the external
digestive tract opening. A thick, smooth muscle layer forms the internal
anal sphincter at the superior end of the anal canal, but skeletal muscle
forms the external anal sphincter at the inferior end. The external anal
sphincter is under voluntary motor control (via skeletal muscle), but the
internal anal sphincter is under involuntary motor control (via smooth
muscle). The presence of stool inside the rectum is sensed by the
nervous system, which results in parasympathetic relaxation of the
internal anal sphincter (a process known as the defecation reflex);
“bearing down” is usually sufficient to expel fecal material through the
external anal sphincter (Figure 22.26).
Figure 22.26. The defecation reflex involves the activation of stretch receptors, which result
in the involuntary relaxation of the internal anal sphincter. In order to expel feces from the
body, the external anal sphincter must also be relaxed through voluntary and conscious
signals from the cerebral cortex.
22.7.1 Motility in the Large Intestine

Because digestion is essentially completed by the time the chyme enters
the colon, there are few colon segmentation contractions. Peristalsis
moves the chyme through the ascending colon. Approximately 3-4 times
per day, large peristaltic contractions, known as mass movements,
move through the transverse and descending colon. Mass movements
extend over a larger part of the colon (over more than 20 cm) compared
to normal peristaltic contractions and propel chyme toward the anus.
Mass movements are commonly initiated after meals when the stomach
and duodenum increase in volume, especially during the first meal of the
day. These changes in volume are known as the enterocolic reflex and
the duodenocolic reflex, respectively. The mass movements usually
persist for 10 to 30 minutes and then stop for 10 to 12 hours. Mass
movements in the colon stimulate the defecation reflex. A person is
usually unaware of mass movements through the large intestine.
Question 22.27
Match the following with their function.
● Press space or enter to grab Activates large intestine motility in response to small
intestine stretching
● Enterocolic reflex
● Activates large intestine motility in response to small intestine stretching
● Press space or enter to grab Activates large intestine motility in response to stomach
stretching
● Duodenocolic reflex
● Activates large intestine motility in response to stomach stretching
● Press space or enter to grab Carry chyme over large distances in the large intestine
● Segmentation contraction
● Carry chyme over large distances in the large intestine
● Press space or enter to grab Occur infrequently in the large intestine

● Peristaltic contractions
● Occur infrequently in the large intestine
Unanswered
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Feces-induced rectal wall distension is a stimulus for the defecation

reflex. Locally, the defecation reflex stimulates weak wall contractions
and relaxation of the internal anal sphincter. Central nervous system
reflexes (mediated by parasympathetic fibers) cause strong wall
contractions and are normally responsible for most of the defecation
reflex. Rectal distension triggers sensory neurons that send information
to the sacral spinal cord and trigger fibers to stimulate strong colonic and
rectal contractions. The defecation reflex causes the internal anal
sphincter to relax, whereas the external anal sphincter is under
conscious control and prevents feces from moving through the anal
opening. When the external anal sphincter is consciously relaxed, feces
are expelled. Fortunately, the defecation reflex is short-lived and quickly
declines.
22.7.2 Secretions of the Large Intestine
The colon’s mucosal layer is lined with goblet cells scattered along its
length and numerous crypts that are lined almost exclusively with goblet
cells. There is little digestive activity associated with colon secretions,
which are predominantly mucous. Mucous coats the colon wall and
provides a matrix that helps fecal matter stick together. Mucous
secretion is increased by the presence of fecal material within the large
intestine that activates wall mechanoreceptors. Parasympathetic
stimulation of the large intestine can also increase mucous secretion.
The large mass of gut flora produces acid in the colon. A molecular
pump in the colon epithelial cells exchanges bicarbonate ions for Cl-, and
another pump exchanges Na+ for H+. The increased NaCl in the colon
pulls water through the colon wall by osmosis. The feces that leave the
digestive tract consist of water, mucus, undigested food,
microorganisms, and sloughed-off epithelial cells. Microorganisms
account for approximately 30% of feces’ dry weight. Irritation or
inflammation of the large intestine due to bacterial infection (bacterial
enteritis) causes the colon to secrete large amounts of mucus,
electrolytes, and water. The increased colon secretion causes distension
of the colon and an abnormally frequent discharge of fluid feces known
as diarrhea. Diarrhea can cause significant losses of fluid and
electrolytes and is a potentially serious problem in young children.
However, diarrhea also helps to remove pathogenic bacteria from the
intestine more rapidly.
Question 22.28
Which of the following are secreted into the large intestine?
HCl
Pepsinogen
Pepsin
CO
e
Mucus
Unanswered
Submit
A 22-year-old girl presents to the emergency room with nausea,

vomiting, fever, and abdominal pain. On physical examination, her
abdominal pain is located to the right iliac region. The pain she is
experiencing is classified as rebound tenderness, meaning that when
pressure is applied there is no pain, but when pressure is released there
is extreme pain.
Diagnosis: Appendicitis
Hover here to find out how this condition would be treated.
22.7.3 The Microbiome of the Large Intestine

If you have seen a yogurt advertisement recently, then you have
probably heard about the strong link between the normal gut microbiome
and human health. The normal human gut microbiome is composed of
bacteria from two major phyla: Bacteroidetes and Firmicutes. Most
humans have between 500 to 1000 species within their microbiome,
which are most commonly anaerobic bacteria. The infant gut microbiome
is more variable than the adult, and the adult microbiome is commonly
established by the 3rd year of life. In the adult microbiome, there are
temporal and spatial variations from the esophagus to the rectum during
a person's life. The normal microbiome plays key roles in nutrient and
drug metabolism, maintenance of the mucosa, and the immune system.
The composition of the microbiome varies with birthing method (e.g.,
vaginal vs. cesarean), diet as an infant (e.g., breast milk vs. formula),
diet as an adult (e.g., vegan vs. animal proteins), and pharmaceutical
use (e.g., antibiotics).
In addition to fermenting carbohydrates that escape digestion in the

small intestine, the gut microbiome aids in the synthesis of vitamin K and
several forms of vitamin B (linoleic acid, deoxycholic, and lithocholic
acids). It also modulates host immunity by preventing the growth of
pathogenic bacteria, helping to maintain the mucosal layer, influencing
the gut-associated lymphoid tissues (GALT), and helping to ensure
normal structure and function in the digestive tract. This is one reason
why many people choose to take probiotic supplements. Bacteria in the
large intestine also metabolize a variety of sterols and steroids, which
are excreted with bile into the intestines. This often changes biological
half-life (see Chapter 16: The Endocrine System). For example,
antibiotic therapy can sometimes lead to unplanned pregnancy by
interfering with the reabsorption of the steroids found in birth control pills.
Question 22.30
Which of the following is not a normal function of the large intestine?
Microbiome fermentation of starches to small organic molecules
Inhibition of the growth of pathogenic bacteria
Absorption of water
d
Microbiome metabolism of sterols and steroids
Absorption of nutrients
Unanswered
Submit
Where do the bacteria that colonize the

large intestine come from in a newborn
baby?
Lecture Video - Large

Intestines
22.8 Accessory Organs
Unless a person’s life has been touched by a disease of the digestive
system, such as hepatitis C or gallstones, most of us know very little
about what our accessory organs actually do. These organs are critical
for digestive function and for life. For example, we cannot survive without
our liver and pancreas. These organs provide essential secretions
(enzymes, detergents, electrolytes) required for proper digestion. The 3D
model below shows how the organs of the GI tract, and the accessory
organs, fit together to form the digestive system.
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22.8.1 The Liver and Gallbladder

Found in the right upper quadrant of the abdomen and against the
inferior surface of the diaphragm, the liver is the body’s largest internal
organ (Figure 22.27). At the level of gross anatomy, students often
confuse the liver with the stomach (or sometimes the heart) because of
its large size. The major physiological functions of the liver and
gallbladder in the digestive system are:
● To detoxify a wide range of chemicals brought into the body

through the ingestion of food (liver).
● To metabolize and interconvert nutrients into a wide range of
forms for storage or use by other tissues (liver).
● To store bile until it is needed for fat digestion (gallbladder).
Figure 22.27. The liver processes absorbed nutrients, adjusts blood levels of nutrients, and
produces bile which is stored in the gallbladder (green). As seen in the magnified image
above, the gallbladder (green) consists of a fundus, body, and neck that is connected by the
cystic duct to the common hepatic duct, which then goes on to become the bile duct.
Like our other large organs (e.g., the lungs), the liver is divided into
lobes. The major (left and right) lobes are larger than the two minor
(caudate and quadrate) lobes.
Question 22.31
Bile is produced by the ________ and stored in the ________.
gallbladder; liver
liver; small intestine
small intestine; liver
d
liver; gallbladder
Unanswered
Submit
The organization of the liver is unique and central to the organ’s ability to
carry out its physiological functions (remember that structure dictates
function) (Figure 22.28). Connective tissue forms a branching network of
septa (walls) that make up the three-dimensional structure of the liver.
Blood, lymphatic vessels, nerves, and ducts follow the connective tissue
branches throughout the liver. Inside the liver, the sheets of connective
tissue divide the liver into thousands of lobules. Each lobule is roughly
hexagonal (six-sided) in shape, with portal triads at the vertices and a
central vein in the middle. The portal triads are formed by three
structures: a hepatic portal vein, a hepatic artery, and a bile duct. Nerves
and lymphatic vessels are also located in these areas. Blood from the
hepatic portal vein and hepatic artery move through the lobules to the
central vein in the center of each lobule. Central veins unite to form the
hepatic veins, which exit the liver on the posterior and inferior surfaces
and empty into the inferior vena cava.
Hepatic cords radiate out from each lobule central vein like wheel
spokes. The hepatic cords are filled with hepatocytes. Hepatocytes are
responsible for bile production, nutrient metabolism and interconversion,
detoxification of harmful substances, and synthesis of blood
components. The spaces between hepatic cords form hepatic sinusoids
that act as blood channels. The bile canaliculus, a cleft-like lumen, lies
between each cord.
Figure 22.28. Liver hepatocytes are organized into lobules (top right) and have direct
access to venous blood that is collected from the digestive tract, as seen in the enlarged
image (bottom left).Portal triads, consisting of a bile duct, hepatic artery, and hepatic portal
vein, are found at the vertices of lobules and the central vein is found in the middle.
Activate
22.8.2 The Hepatic Portal System

The circulatory system of the liver is unlike that of any other organ. The
majority of the liver's blood supply is venous blood rather than arterial
blood. Roughly 75% of the blood entering the liver is from the portal vein,
which carries all of the venous blood returning from the small intestine,
stomach, pancreas, and spleen. The portal vein empties into the hepatic
sinusoids. Sinusoids are channels that are lined with fenestrated
endothelial cells and are surrounded by hepatocytes. The remaining
25% of the blood supply to the liver is arterial blood from the hepatic
artery, which also empties into the sinusoids. Hepatocytes remove O2
and nutrients from the blood to support their metabolic needs. The
nutrients are consumed, interconverted, stored, or converted to new
molecules. As blood flows through the sinusoids, a considerable amount
of plasma is filtered into the space between the endothelium and
hepatocytes, providing a major fraction of the body's lymphatic fluid.
Blood drains from the sinusoids into the central vein of each lobule.
Central veins drain into the larger hepatic vein, which exits the liver and
ultimately empties into the inferior vena cava.
Figure 22.29. The hepatic portal vein collects blood from the intestinal tract (intestines,
pancreas, spleen, and stomach) and makes it available to the liver, which can then adjust
the concentration of critical nutrients in the blood.Blood in the hepatic vein returns to the
heart via the inferior vena cava, where it then oxygenates and returns to the body via the
aorta. It then returns to the intestinal tract or liver via the hepatic artery.
This unique arrangement of blood flow is known as the hepatic portal
system. It ensures that the liver is the first organ to be exposed to
absorbed nutrients and dietary toxins. This positions the liver to act on
potentially harmful substances before they can damage other body
organ, and it also allows it to make important metabolic decisions to
regulate homeostasis.
22.8.3 Liver Secretions: Bile

Bile, which is produced by hepatocytes, is a complex fluid that contains
water, electrolytes, and organic molecules including bile acids,
cholesterol, lecithin, and bilirubin. Bile emulsifies fat particles (makes
them into small droplets) within the small intestine, making them easier
to digest and absorb. It also aids in the absorption of fat soluble vitamins
and neutralizes acidic chyme. Many waste products are removed from
the body by secretion into bile and elimination in feces.
Figure 22.30. Emulsification involves coating large fat droplets with amphipathic bile salts,
thus making them accessible to lipases and allowing for micelle formation.
Bile flows through the biliary tract into the small intestine. The biliary
system is a series of channels and ducts that carry bile from the liver into
the lumen of the small intestine. Hepatocytes secrete bile into
canaliculi. At the ends of the canaliculi, bile flows into bile ducts that join
into progressively larger ducts and eventually form the common bile
duct, which carries bile into the duodenum. Bile can also be diverted
through the cystic duct into the gallbladder, where it can be stored until
it is needed. As bile flows through the bile ducts, it is modified by the
addition of a watery, bicarbonate-rich secretions from ductal epithelial
cells. The gallbladder stores and concentrates bile (up to five-fold)
between meals.
The flow of bile is lowest during fasting when the majority of the bile is
diverted into the gallbladder for storage. When chyme enters the small
intestine, acid and partially digested fats and proteins stimulate the
secretion of cholecystokinin (CCK) and secretin. CCK is released in
response to the presence of fat in the duodenum. Once released, CCK
stimulates contractions of the gallbladder and common bile duct,
delivering bile to the small intestine. Secretin is secreted in response to
acid in the duodenum. Secretin stimulates biliary duct cells to secrete
bicarbonate and water, which expands the volume of bile and increases
its flow out into the intestine. Bile salts also increase bile secretion
through a positive feedback mechanism. Most of the bile salts are
reabsorbed in the ileum and transported back to the liver where they are
reused. The loss of bile salts in the feces is greatly reduced by this
recycling process. Bile secretion continues into the duodenum until it
empties.
Why are dietary triglycerides digested to

free fatty acids and glycerol, if they are then
recombined to form triglycerides within the
cell?
Hover here to see the answer to TQ 22.07.
22.9 The Pancreas

The pancreas is an essential part of the digestive system with very
important functions:
● To produce a watery secretion that is rich in HCO3–.

● To produce enzymes that aid in chemical digestion in the small
intestine.
Figure 22.31. The pancreas, pictured above, lies next to the duodenum and contains a
ductal network.
The pancreas is an elongated organ that lies next to the duodenum

(Figure 22.31). Most of the pancreas is composed of pancreatic exocrine
glands, which contain ducts. Embedded within the exocrine tissue,
however, are roughly one million small clusters of cells called Islets of
Langerhans, which are the endocrine cells of the pancreas that secrete
insulin, glucagon, and several other hormones. Pancreatic exocrine cells
are arranged in grapelike clusters called acini (singular, ancinus) (Figure
22.32).
Figure 22.32. Pancreatic exocrine glands contain acini and ducts, as seen in the
cross-section above. Secretions from the acini travel through the ducts where they can be
modified, and then flow into the pancreatic duct. Islets of Langerhans can also be found in
clusters in the pancreas (seen above on the right). Islets of Langerhans secrete insulin,
glucagon, and other hormones.
The exocrine cells themselves are packed with membrane-bound

granules that contain digestive enzymes, which are released by
exocytosis into the lumen of the acinus. The lumen of each acinus is
connected by small intercalated ducts to larger intralobular ducts. These
intralobular ducts fuse and ultimately become the pancreatic duct, which
drains directly into the duodenum. A smooth muscle sphincter surrounds
the pancreatic duct where it enters the hepatopancreatic ampulla.
22.9.1 Secretions of the Pancreas

The pancreatic exocrine secretions, known as pancreatic juice, are
made up of two products critical to proper digestion: digestive enzymes
and HCO3– (Figure 22.33). The enzymes are synthesized and secreted
from the exocrine acinar cells, whereas HCO3– is secreted from the
epithelial cells lining small pancreatic ducts. The pancreas secretes a
wide range of enzymes that can reduce many macromolecules into
monomeric subunits, which can be absorbed. Several major groups of
enzymes are critical for efficient digestion: proteases, lipases, and
amylases.
Figure 22.33. Diagram showing the steps of bicarbonate secretion by the pancreatic ductal
cells.
The pancreatic ductal cells play an important role in the neutralization of

chyme (Figure 22.33). The mechanism underlying HCO3– secretion is
essentially the same as that for acid secretion by parietal cells and is
dependent on carbonic anhydrase. In ductal cells, however, HCO3– is
secreted into the lumen of the duct (vs. the extracellular fluid in parietal
cells) and hence into pancreatic juice. The increased intestinal pH that
results from pancreatic secretions neutralizes the acidic chyme and
adjusts the pH for pancreatic enzymes. This is important because the
intestinal wall, unlike the stomach, does not resist acid, and most
digestive organs are not optimized for low pH.
The digestion of proteins is initiated by pepsin in the stomach, but the

majority of proteins are digested by pancreatic proteases. Several
proteases are synthesized in the pancreas and secreted into the lumen
of the small intestine. The two major pancreatic proteases are trypsin
and chymotrypsin, which are synthesized and packaged into zymogen
vesicles as the inactive proenzymes, trypsinogen and
chymotrypsinogen. The secretory vesicles also contain a trypsin
inhibitor that serves as an additional safeguard in case the trypsinogen is
activated to trypsin before being secreted into the duodenum. This
inhibitor is diluted in the duodenum and becomes ineffective. Once
trypsinogen and chymotrypsinogen are released into the small intestine
lumen, they must be converted into their active forms to digest proteins.
Trypsinogen, which is embedded in the intestinal mucosa, is activated by
the enzyme enterokinase by cleavage of a specific peptide sequence,
changing its structure and function. Once trypsin is formed, it activates
chymotrypsinogen (as well as additional molecules of trypsinogen) by
cleavage of another specific peptide sequence. This positive feedback
loop rapidly activates small intestine proteases. Trypsin and
chymotrypsin digest proteins and peptides into smaller peptides, but,
as endopeptidases, they cannot digest proteins and peptides into single
amino acids. Other pancreatic proteases, including carboxypeptidase
(an exopeptidase), can break peptides into individual amino acids.
However, the final digestion of peptides into amino acids is largely
carried out by peptidases located on the brush border.
Question 22.33
Match the following molecules with its function.
● Press space or enter to grab An exopeptidase

● Trypsin
● An exopeptidase
● Press space or enter to grab An inactive endopeptidase released from pancreatic

vesicles
● Chymotrypsinogen
● An inactive endopeptidase released from pancreatic vesicles
● Press space or enter to grab A peptidase that activates other peptidases

● Enterokinase
● A peptidase that activates other peptidases
● Press space or enter to grab An enzyme of the intestinal mucosa

● Carboxypeptidase
● An enzyme of the intestinal mucosa
Unanswered
Submit
The major form of dietary fat is triglyceride, which cannot be directly
absorbed across the intestinal mucosa. Rather, it must first be digested
into 2-monoglyceride and two free fatty acids by pancreatic lipase, which
is delivered into the lumen of the gut as a constituent of pancreatic juice.
Liver bile salts must also be present in the lumen of the intestine for
lipase to efficiently digest dietary triglyceride and for the resulting fatty
acids and monoglycerides to be absorbed. As a result, normal digestion
and absorption of dietary fat are critically dependent on secretions from
both the pancreas and the liver. The major dietary carbohydrate is
starch, a storage form of plant glucose. Salivary amylase initiates plant
starch digestion in the mouth, but this enzyme is inactivated by stomach
acid. In the stomach, pancreatic amylase continues to hydrolyze starch
into maltose (a glucose-glucose disaccharide), trisaccharide maltotriose,
and small branch-point fragments called dextrins. In addition to the
proteases, lipase, and amylase, the pancreas produces a host of other
digestive enzymes, including ribonuclease (which digests RNA),
deoxyribonuclease (which digests DNA), gelatinase, and elastase (which
digests connective tissue).
Pancreatic exocrine secretions are regulated by both neural and

endocrine controls (Table 22.1). Click here to view the text version of
Table 22.1. Between meals, very little secretion takes place, but as food
enters the stomach and chyme flows into the small intestine, pancreatic
secretions are strongly stimulated. Like the stomach, the pancreas is
innervated by the vagus nerve, which applies a low-level stimulus to
secretion in response to anticipation of a meal. However, the most
important stimuli for pancreatic secretion come from three hormones
secreted by the digestive tract (Figure 22.34).
Table 22.1. Key hormones involved in the control of digestive tract secretions.
Cholecystokinin (CCK) is a hormone synthesized and secreted by

duodenal endocrine cells (Figure 22.34). The presence of partially
digested proteins and fats in the small intestine stimulates CCK
secretion into the blood. CCK binds to receptors on pancreatic acinar
cells, stimulating them to secrete large quantities of digestive enzymes.
Figure 22.34. Cholecystokinin (CCK), which is secreted primarily from duodenal cells,
inhibits gastric motility, stimulates enzyme production in the pancreas, and stimulates bile
release from the gallbladder, as illustrated in the schematic above.
Secretin, which is also produced by duodenal endocrine cells, is

secreted in response to acid in the duodenum when chyme enters from
the stomach. Secretin stimulates pancreatic duct cells to secrete water
and HCO3–. As soon as this occurs, digestive enzymes secreted by the
acinar cells are flushed out of the pancreas through the pancreatic duct
into the duodenum.
Gastrin, which is very similar to CCK, is secreted in large amounts by

the stomach in response to gastric distention and irritation of the
stomach wall. In addition to stimulating acid secretion by parietal cells,
gastrin stimulates pancreatic acinar cells to secrete digestive enzymes.
The pancreas is also an important endocrine organ. It secretes insulin

and glucagon (glucagon is also secreted by the gastrointestinal tract) in
response to changes in blood glucose levels. Insulin decreases blood
glucose levels after a meal is eaten, whereas glucagon is one of several
hormones that increases blood glucose levels between meals. The role
of insulin and glucagon in energy metabolism is discussed in detail in
Chapter 23: Nutrition and Metabolism. There are at least 30 hormones
and paracrine factors released within the digestive tract, which should
not be surprising given that the digestive tract is a disassembly line,
where coordination over its entire length is essential.
22.10 Regulation of the

Digestive System
Given its many components, the digestive system requires an equally
complex control system to keep it functioning properly and to maintain
homeostasis. As we discuss this complex regulation of the digestive
system below, always try to keep in mind the major functional patterns of
this chapter—movement, digestion, absorption, and secretion.
The digestive system is regulated at many levels, including:
● At higher brain centers (responding to taste and smell)

● Locally in response to changing sensory information (responding
to chemicals and pH)
● Via reflex loops (both short and long)
● Via hormones (released from the gastrointestinal and other
endocrine tissues)
22.10.1 Regulation of the Digestive System

by the Nervous System
There are two levels of nervous system control in the digestive system.
Local control comes from the enteric nervous system in the submucosal
and myenteric plexuses. Higher-level control comes from the
parasympathetic branch of the autonomic nervous system. There are
three major types of enteric neurons: sensory neurons that detect
chemical changes in the composition of the digestive tract contents and
mechanical changes in the digestive tract wall (stretch); motor neurons
that stimulate or inhibit smooth muscle contraction or glandular
secretions; and interneurons that integrate signals between afferent
and efferent neurons.
The enteric nervous system is capable of controlling the digestive tract

independently of the central nervous system. One example of this is the
basal electrical rhythm induced by the enteric nervous system. This
electrical depolarization is usually not sufficient to trigger action
potentials in smooth muscle on its own, but it can be modified and
brought to threshold with the help of other signals. The enteric nervous
system regulates local reflexes that control activities within specific
regions of the digestive tract. For example, vomiting requires the
coordinated action of the stomach and the esophagus. The enteric
nervous system also relays signals sent from the central nervous system
that protect the digestive tract against external threats. When the brain
signals danger, postganglionic parasympathetic fibers stimulate small
intestine mast cells to release histamine and other inflammatory
chemicals, which attract immune cells from the bloodstream to the area.
Have you ever noticed your stomach growling loudly during an exam
because you skipped your breakfast? Digestive tract reflexes initiated by
the central nervous system can also be activated by the sight, smell, and
taste of food, which stimulates the sensation of hunger and causes the
stomach to growl. The stereotypical growl is created by air movement
within an empty stomach that occurs when the stomach walls contract.
The vast majority of innervation to the digestive tract is parasympathetic
via the vagus nerve, although there is a small amount of sympathetic
innervation that inhibits contraction and secretion and decreases blood
flow to the digestive tract during a fight or flight response.
Communication between the central nervous system and the enteric

nervous system is bi-directional, which means that the GI tract can also
relay information to the central nervous system. As an example, consider
the regulation of stomach secretions by the central nervous system,
which occurs through several reflexes that are integrated by the enteric
nervous system. The regulation of stomach secretion is divided into
three phases: cephalic, gastric, and intestinal. The following video
demonstrates the components of a reflex arc, similar to those found in
the GI tract.
In the cephalic phase of regulation of stomach secretion (Figure 22.35),
neurons within the medulla oblongata are stimulated by olfactory and
gustatory afferent neurons, information from tactile receptors during
chewing and swallowing, and memories of food. Information, in the form
of preganglionic parasympathetic fibers, is sent through the vagus (X)
nerve to the submucosal enteric plexus. Preganglionic fibers synapse on
postganglionic parasympathetic cells within the submucosal plexus,
which, in turn, stimulate mucosal glands.
Figure 22.35. The cephalic phase of regulation of stomach secretion occurs in response to
the sight, smell, taste, or thought of food. Stimulation from the cerebral cortex,
hypothalamus, and medulla is relayed via the vagus nerve and parasympathetic neurons to
trigger stomach acid secretion, whereas loss of appetite or depression can trigger the
cerebral cortex to activate sympathetic neurons that inhibit stomach acid secretion.
The greatest volume of stomach secretion is produced during the

gastric phase, which is initiated by the presence of food within the
stomach (Figure 22.36). The primary stimuli are stomach distension and
the presence of amino acids and peptides within the chyme. Distension
of the stomach wall stimulates mechanoreceptors. Mechanoreceptors
initiate reflexes that involve both the central nervous system and the
enteric nervous system and results various secretions from specialized
cells. These secretions include mucus, HCl and intrinsic factor from
parietal cells, pepsinogen from chief cells, and the hormone gastrin from
G cells. Amino acids and peptides released by the digestive action of
pepsin on proteins also stimulate gastrin and HCl release. These
secretions are limited, however, by the low pH that they create. When
stomach pH declines to less than 2, somatostatin inhibits acid secretion.
This negative feedback loop protects the stomach wall from the
damaging effects of acid and pepsin (and ensures that acid and pepsin
are secreted into the stomach only when food is present). The exact
mechanism by which this response is mediated is not clearly
understood.
Figure 22.36. The gastric phase of regulation of stomach secretion is initiated by the
presence of food within the stomach, triggering local reflexes or stomach distension
(stretch) that initiates the vagovagal reflex and stimulates the vagus nerve via the medulla.
Meanwhile, excessive stomach acid can inhibit gastrin, and emotional upset can trigger
sympathetic neurons to inhibit stomach acid secretion.
Question 22.34
Which of the following does not occur during the cephalic phase?
a
Saliva secretion in the mouth
Stomach growling
Secretion of mucus in the stomach
Release of secretin from the small intestine
Unanswered
Submit
Gastric emptying brings acidic chyme into the small intestine and
triggers the intestinal phase of the regulation of stomach secretions
(Figure 22.37). During this phase, both neural and hormonal
mechanisms stimulate intestinal secretions while inhibiting stomach
secretions. The chyme effect is mediated by the duodenum’s secretion
of the hormone secretin into the general circulation. Secretin inhibits
both parietal and chief cells. Acidic chyme can also initiate a local enteric
reflex, which also inhibits stomach secretions. Chyme that contains fat
digestion products, fatty acids, and certain other lipids causes the
release of gastric inhibitory peptide (GIP) and CCK from the duodenum.
GIP strongly inhibits gastric secretions, and CCK inhibits gastric
secretions to a lesser extent. The inhibition of gastric secretions is also
under nervous control. Distension of the duodenal wall and reduced
duodenal pH activate the central nervous system-mediated enterogastric
reflex and reduce stomach secretions.
Figure 22.37. The intestinal phase of regulation of stomach secretions occurs as food
moves beyond the stomach. Peptides and amino acids in the duodenum activate gastrin,
whereas distension, fats, and irritants in the duodenum can trigger the enterogastric reflex
and inhibit stomach acid secretions via the medulla, vagus nerve, and parasympathetic
neurons, or through stomach neuroendocrine cells and factors such as secretin, gastric
inhibit inhibitory polypeptide (GIP), cholecystokinin (CCK), and vasoactive intestinal peptide
(VIP).
Question 22.35
The intestinal phase is triggered by ________________.
the sight, smell, and taste of food
the presence of food in the stomach
the secretion of hormones by the pancreas
the presence of chyme in the small intestine
Unanswered
Submit
22.10.2 Endocrine Regulation of the

Digestive System
The digestive system is a major endocrine tissue (see Chapter 16: The
Endocrine System) that produces a wide range of hormones that act on
cells within the digestive system and also regulate cells and tissues
outside the digestive system. These hormones are a major part of the
cell-to-cell communication pathways that regulate the activity of the
digestive system. Hormones regulate smooth muscle contraction and
glandular secretion. There are also a number of paracrine compounds
(including histamine) that are released locally within the digestive system
and influence the activity of nearby cells.
22.10.3 Physical Regulation of the Digestive

System
The digestive system is also influenced by the physical presence of food
inside the digestive tract. The distension of the digestive tract wall
induces smooth muscle contraction, altering the tone of the digestive
tract. The presence of specific food components (sugars, amino acids,
and fats) within the digestive tract also affects smooth muscle
contraction and glandular secretion, as does the osmolarity of the
digestive tract contents.
22.11 Digestion and
Absorption at the Molecular
Level
Absorption is the process by which molecules are moved out of the
digestive tract and into the circulatory system for distribution throughout
the body. Absorption can occur throughout the digestive tract, although
the vast majority of digestion takes place in the duodenum and jejunum
of the small intestine. However, some small fat-soluble components can
be absorbed through the thin mucosa of the oral cavity, aspirin and
alcohol are absorbed in the stomach, and water is absorbed in the large
intestine. Absorption and digestion are closely linked processes because
most (but not all) molecules must be broken down before they are
absorbed. Exceptions include vitamins, minerals, and water.
22.11.1 Carbohydrates Are Absorbed as

Simple Sugars
Most ingested carbohydrates are from plant sources and consist
primarily of polysaccharides, starches, and glycogen. The human diet
also has disaccharides, including sucrose (white sugar) and lactose (milk
sugar); and monosaccharides, including glucose and fructose. During
the digestion process, polysaccharides are broken down by amylases
into smaller fragments and disaccharides. Thus, carbohydrate digestion
begins in the oral cavity with salivary amylase and is completed in the
small intestine with the help of pancreatic amylases and brush border
enzymes. Brush border enzymes, which are found in intestinal epithelial
microvilli membranes, are disaccharidases that digest disaccharides into
monosaccharides for absorption.
22.11.2 Proteins Are Absorbed in Many

Forms
Protein digestion begins in the stomach and continues in the small
intestine. Once protein and polypeptide chains leave the stomach, the
pancreatic proteases (trypsin, chymotrypsin, and carboxypeptidases)
digest proteins into smaller polypeptide chains. These are then digested
to tripeptides, dipeptides, and single amino acids by peptidase at the
brush border.
The mechanism by which amino acids are absorbed into intestinal

epithelial cells is similar to that of monosaccharides. The microvilli
membrane has at least four Na+-dependent amino acid transporters, one
each for acidic, basic, neutral, and hydrophobic amino acids. These
active transporters bind amino acids only after binding Na+. Absorption of
amino acids, similar to monosaccharides, generates an osmotic gradient
that drives water absorption. The basolateral membrane of intestinal
epithelial cells contains additional transporters that export amino acids
from the cells into the blood. These are not dependent on Na+ gradients.
There is virtually no absorption of peptides longer than three amino acids

because larger peptides are too large to fit through transporters, and
proteins cannot permeate tight junctions. However, dipeptides and
tripeptides are readily absorbed in the small intestine independent of
Na+. Once inside the intestinal epithelial cells, the bulk of the dipeptides
and tripeptides are digested into amino acids by cytoplasmic peptidases,
then exported into blood.
Both digestion and absorption are also linked to solubility. Whereas

carbohydrates and most proteins are soluble in water, lipids are not. This
complicates the process of lipid digestion by requiring that the first step
in lipid digestion be emulsification, which breaks large fat droplets up
into smaller lipid droplets. This is sort of like shaking up a bottle of salad
dressing to mix the oil in with the water and seasonings. Emulsification
aids digestion by increasing the surface area for digestive lipases to
work. Without emulsification, many fats would go undigested because
lipases, which are water-soluble themselves, would only have access to
fats in the outermost layer (Figure 22.39).
Figure 22.39. A schematic of micelle fat absorption from dietary fats.
Lipases are primarily secreted by the pancreas (although there is a small

amount of salivary and gastric lipase). Lipases produce free fatty acids,
diglycerides, monoglycerides, and glycerol. After digestion, these
components form micelles, which combine with bile salts. Bile salts are
amphipathic: one end of the molecule is water-soluble (hydrophilic) and
the other end is fat-soluble (hydrophobic). Lecithins, which give bile this
property, are commonly found in non-stick cooking spray and processed
foods. The hydrophobic ends point toward the center of the micelles,
which contains free fatty acids and monoglycerides, while the hydrophilic
ends interact with the aqueous environment. When a micelle comes into
contact with an intestinal epithelial cell, free fatty acids and
monoglycerides enter intestinal epithelial cells by simple diffusion across
the cell membrane.
Inside the aqueous environment of the cell, free fatty acids are combined
with glycerol to form triglycerides in the endoplasmic reticulum. The
resulting molecules, known as chylomicrons, are made up of
triglycerides (90%), cholesterol (5%), phospholipids (4%), and protein
(1%). Chylomicrons are extruded from the Golgi into vesicles, which are
transported to the basolateral membrane of the intestinal epithelial cells.
The vesicles fuse with the cell membrane and undergo exocytosis,
releasing the chylomicrons into the space outside the cells. Unlike all
other absorbed nutrients which enter the hepatic portal system via the
portal vein, chylomicrons are absorbed into the lymphatic system
through vessels known as lacteals. This is necessary because
lymphatic capillaries, unlike blood vessels, have no basement
membrane and are permeable to larger particles. Chylomicrons move
through the lymphatic system to the venous blood, and then to the liver
or adipose tissue. Details of this process are covered in Chapter 23:
Nutrition and Metabolism.
Question 22.37
Place the following events related to lipid digestion in order.
Press space or enter to grab Movement across basolateral membrane

Movement across basolateral membrane
Press space or enter to grab Diffusion across luminal membrane

Diffusion across luminal membrane
Press space or enter to grab Micelles form

Micelles form
Press space or enter to grab Chylomicron forms

Chylomicron forms
Press space or enter to grab Chylomicron enters lacteal

Chylomicron enters lacteal
Press space or enter to grab CCK stimulates gallbladder

CCK stimulates gallbladder
Press space or enter to grab Free fatty acids enter endoplasmic reticulum
Free fatty acids enter endoplasmic reticulum
Press space or enter to grab Emulsification of fats

Emulsification of fats
Press space or enter to grab Pancreatic lipases begin to digest triglycerides

Pancreatic lipases begin to digest triglycerides
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Lecture Video - Accessory

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Question 22.38
What topic did you understand the least from this chapter? Explain. (Remember that we are looking
for what you understood the least and not necessarily something that you didn’t understand.)
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Function of the organs
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22.13 Summary
This chapter reviews the structures and functions of the digestive
system. One of the most fundamentally important things we do each day
is eating. We eat when we are hungry, when we are happy, and when we
are sad. Eating food makes us feel full and satisfied or sometimes
nauseous, but most of us have a limited understanding of what happens
to that food when it enters our body. We all know that food enters
through our mouths, where we chew (masticate) it, breaking it into
smaller pieces that we mix with saliva and swallow.
Anatomy of the Digestive System

● Food enters a very long tube (up to 22 feet in length)—the
digestive tract—that is functionally differentiated, with specific
segments responsible for different activities.
● Food moves from the mouth into the esophagus by the process of
swallowing, a coordinated effort of the tongue and pharyngeal
muscles. From there, the food bolus travels to the stomach,
where chemical and enzymatic digestion of the food occurs.
● After digestion, food, now known as chyme, enters the small
intestine, where the majority of enzymatic digestion occurs. The
small intestine is subdivided into the duodenum, jejunum, and
ileum.
● The small intestine is the site of the majority of absorption
(bringing nutrients into the body).
● What remains after passing through the small intestine travels to
the large intestine, where water and ions are reabsorbed and
where the microbiome, a collection of bacteria, aids in digestion,
absorption, and immune surveillance.
● The large intestine is composed of the ascending, transverse,
descending, and sigmoidcolon, terminating in the anus where
waste material—feces—is excreted from the body.
● The digestion of food and absorption of nutrients is critically
dependent on the properties of the wall of the digestive tract. The
digestive wall is uniquely composed of four layers or tunics, the
outer serosal layer, then a layer of smooth muscle, then a
submucosal layer, and then the innermost mucosal layer.
● Within the muscle and submucosal layers is the enteric nervous
system that controls smooth muscle activity and the secretion of
various buffers, enzymes, and other factors into the lumen.
● The enteric nervous system is composed of the myenteric
plexus and the submucosal plexus.
● Given that the digestive tract is functionally differentiated, it is not
surprising that the digestive wall varies in different regions of the
digestive tract. For example, the stomach wall has gastric pits
that secrete acid, buffers, and enzymes. On the small intestine,
lacteals in the intestinal wall aid in food absorption.
Movement
● Movement along the length of the digestive tract occurs via
peristalsis—a coordinated contraction of smooth muscle layers
that moves food (or chyme) directionally along the length of the
tube.
● Segmentation is a different pattern of smooth muscle
contractions that mixes food with secretions.
● Movements within the digestive tract are under the control of the
enteric nervous system and are modifiable by the actions of the
CNS and hormones.
Secretions and Absorption
● Secretions and movement aid in the breakdown of food into its
smallest possible components, which are absorbed into the
hepatic portal blood supply, which delivers nutrients to the liver
for processing.
● Secretion begins with saliva in the mouth, mucus all along the
digestive tract, acid and enzymes in the stomach, and enzymes
and buffers in the small intestine.
● Absorption of nutrients and water into the body requires energy
and the presence of specific transporters and channels in the
intestinal wall.
Accessory Organs
● The digestion and absorption of food require the activities of other
organs. The liver is responsible for converting nutrients absorbed
from the digestive tract into compounds cells can use and storing
them until needed. It also produces bile salts for theemulsification
of fats, filters toxins from the blood, and assesses the body's
energy status.
● The pancreas is responsible for the secretion of alkaline buffers
into the small intestine to neutralize stomach acid along with a
wide range of proteases and enzymes that digest lipids, sugars,
and nucleic acids. It also plays an important role in the regulation
of blood sugar through the secretion of insulin and glucagon
(discussed in Chapter 16).
● The gallbladder stores bile for secretion into the small intestine.
Regulation
● The control of the digestive tract is as complex as the tract itself.
The tract is primarily controlled by the autonomic nervous system
(ANS) and the enteric nervous system, both of which generate
complex and local reflexes respectively. These reflexes regulate
all aspects of movement, secretion, and absorption along the
gastrointestinal tract.
● Secretions are regulated in three phases:
○ Cephalic phase – responds to olfactory and gustatory
stimuli, chewing and swallowing and food memories
○ Gastric phase – most secretions are produced in this
phase; stimulated by the presence of food in the stomach
○ Intestinal phase – stimulated by the presence of acidic
chyme in the small intestine
Health
● The functioning of the digestive tract has a significant impact on
our health by affecting our weight, and by the ingestion of
nutrients, minerals, and water. We ignore the functioning of the
digestive tract until we have a problem with it—like constipation,
diarrhea, or irritable bowel. However, the normal functioning of the
digestive tract is essential to our existence.

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Chapter 22:

How nutrients can absorb

How different molecules are digested

How our micronutrients turn helps tissues grow or atrophy.

22.2.1 The Anatomy of the Gastrointestinal

​ Press space or enter to grab Ascending colon

​ Press space or enter to grab Transverse colon

​ Press space or enter to grab Ileum

​ Press space or enter to grab Descending colon

​ Press space or enter to grab Stomach

​ Press space or enter to grab Jejunum

​ Press space or enter to grab Pharynx

​ Press space or enter to grab Esophagus

​ Press space or enter to grab Cecum

​ Press space or enter to grab Duodenum

HomeworkUnansweredDue Oct 31st, 12:30 PM

● Press space or enter to grab Major site of digestion

● Major site of digestion

● Press space or enter to grab Connects mouth to esophagus

● Connects mouth to esophagus

● Press space or enter to grab Major site of water absorption

● Major site of water absorption

● Press space or enter to grab Begins enzymatic digestion of proteins

● Begins enzymatic digestion of proteins

● Press space or enter to grab Connects pharynx to stomach

● Connects pharynx to stomach

● Press space or enter to grab Terminal end large intestine

● Terminal end large intestine

● Press space or enter to grab Vestigial immune structure

● Vestigial immune structure

● Press space or enter to grab Contains salivary glands

22.2.2 Accessory Organs and Structures of

HomeworkUnansweredDue Oct 31st, 12:30 PM

Match the following accessory organs with their general function.

● Press space or enter to grab Begins digestion in the mouth

● Begins digestion in the mouth

● Press space or enter to grab Bile storage

● Press space or enter to grab Carries bile into small intestine

● Carries bile into small intestine

● Press space or enter to grab Digestive enzyme production

● Digestive enzyme production

● Press space or enter to grab Carries pancreatic secretions into duodenum

● Carries pancreatic secretions into duodenum

● Press space or enter to grab Produces bile

● Press space or enter to grab Aids in swallowing

The complexity of the digestive system can be overwhelming to students

is surrounded by the submucosa, which contains the submucosal plexus (Meissner's

layer is the outermost layer and is contiguous with the mesentery.

HomeworkUnansweredDue Oct 31st, 12:30 PM

The innermost layer within the digestive tract is the ________.

Please select an answer to submit

The muscularis externa layer, as its name suggests, is composed largely

HomeworkUnansweredDue Oct 31st, 12:30 PM

Gastric secretions may decrease

Pain information will be relayed through the enteric nervous system

Digestion will be impaired

Changes in gastric motility

Please select an answer to submit

Press space or enter to grab Ascending colon

Press space or enter to grab Transverse colon

Press space or enter to grab Ileum

Press space or enter to grab Descending colon

Press space or enter to grab Stomach

Press space or enter to grab Jejunum

Press space or enter to grab Pharynx

Press space or enter to grab Esophagus

Press space or enter to grab Cecum

Press space or enter to grab Duodenum

Press space or enter to grab Lamina propria

Press space or enter to grab Circular layer of smooth muscle

Press space or enter to grab Muscularis mucosa

Press space or enter to grab Mucosal epithelium

Press space or enter to grab Serosa

Press space or enter to grab Submucosa

Press space or enter to grab Myenteric plexus

Press space or enter to grab Longitudinal layer of smooth muscle

with their hands.

with the stomach lumen. [5]

release factors such as gastrin, histamine and acetylcholine.