Ganong's Review of Medical Physiology, 23rd Edition

arise centrally. For example, afferents from the vestibular
nuclei mediate the nausea and vomiting of motion sickness.
Other afferents presumably reach the vomiting control areas
from the diencephalon and limbic system, because emetic
responses to emotionally charged stimuli also occur. Thus, we
speak of “nauseating smells” and “sickening sights.”
Chemoreceptor cells in the medulla can also initiate vomiting
when they are stimulated by certain circulating chemical
agents. The chemoreceptor trigger zone in which these cells
are located (Figure 28–6) is in the area postrema, a V-shaped
band of tissue on the lateral walls of the fourth ventricle near
the obex. This structure is one of the circumventricular
organs (see Chapter34) and is not protected by the blood–
brain barrier. Lesions of the area postrema have little effect on
the vomiting response to gastrointestinal irritation or motion
sickness, but abolish the vomiting that follows injection of
apomorphine and a number of other emetic drugs. Such
lesions also decrease vomiting in uremia and radiation sickness,
both of which may be associated with endogenous production
of circulating emetic substances.
Serotonin (5-HT) released from enterochromaffin cells in
the small intestine appears to initiate impulses via 5-HT 3
receptors that trigger vomiting. In addition, there are dopamine
D2 receptors and 5-HT 3 receptors in the area postrema
and adjacent nucleus of the solitary tract. 5-HT 3 antagonists
such as ondansetron and D 2 antagonists such as chlorpromazine
and haloperidol are effective antiemetic agents. Corticosteroids,
cannabinoids, and benzodiazepines, alone or in
combination with 5-HT 3 and D 2 antagonists, are also useful
in treatment of the vomiting produced by chemotherapy. The
mechanisms of action of corticosteroids and cannabinoids are
unknown, whereas the benzodiazepines probably reduce the
anxiety associated with chemotherapy.
SMALL INTESTINE
In the small intestine, the intestinal contents are mixed with
the secretions of the mucosal cells and with pancreatic juice
and bile.
INTESTINAL MOTILITY
The MMCs that pass along the intestine at regular intervals in
the fasting state and their replacement by peristaltic and other
contractions controlled by the BER are described above. In the
small intestine, there are an average of 12 BER cycles/min in
the proximal jejunum, declining to 8/min in the distal ileum.
There are three types of smooth muscle contractions: peristaltic
waves, segmentation contractions, and tonic contractions.
Peristalsis is described above. It propels the intestinal contents
(chyme) toward the large intestines. Segmentation contractions
(Figure 28–1), also described above, move the
chyme to and fro and increase its exposure to the mucosal surface.
These contractions are initiated by focal increases in Ca 2+
influx with waves of increased Ca 2+ concentration spreading
CHAPTER 28 Gastrointestinal Motility 475
CLINICAL BOX 28–3
Ileus
When the intestines are traumatized, there is a direct inhibition
of smooth muscle, which causes a decrease in intestinal
motility. It is due in part to activation of opioid receptors
and is relieved by opioid-blocking drugs. When the peritoneum
is irritated, reflex inhibition occurs due to increased
discharge of noradrenergic fibers in the splanchnic nerves.
Both types of inhibition operate to cause paralytic (adynamic)
ileus after abdominal operations. Because of the
diffuse decrease in peristaltic activity in the small intestine,
its contents are not propelled into the colon, and it becomes
irregularly distended by pockets of gas and fluid.
Intestinal peristalsis returns in 6 to 8 h, followed by gastric
peristalsis, but colonic activity takes 2 to 3 d to return. Adynamic
ileus can be relieved by passing a tube through the
nose down to the small intestine and aspirating the fluid
and gas for a few days until peristalsis returns.
from each focus. Tonic contractions are relatively prolonged
contractions that in effect isolate one segment of the intestine
from another. Note that these last two types of contractions
slow transit in the small intestine to the point that the transit
time is actually longer in the fed than in the fasted state. This
permits longer contact of the chyme with the enterocytes and
fosters absorption (Clinical Box 28–3).
COLON
The colon serves as a reservoir for the residues of meals that cannot
be digested or absorbed (Figure 28–7). Motility in this segment
is likewise slowed to allow the colon to absorb water, Na + ,
and other minerals. By removal of about 90% of the fluid, it converts
the 1000 to 2000 mL of isotonic chyme that enters it each
day from the ileum to about 200 to 250 mL of semisolid feces.
MOTILITY OF THE COLON
The ileum is linked to the colon by a structure known as the
ileocecal valve, which restricts reflux of colonic contents, and
particularly the large numbers of commensal bacteria, into the
relatively sterile ileum. The portion of the ileum containing
the ileocecal valve projects slightly into the cecum, so that increases
in colonic pressure squeeze it shut, whereas increases
in ileal pressure open it. It is normally closed. Each time a peristaltic
wave reaches it, it opens briefly, permitting some of the
ileal chyme to squirt into the cecum. When food leaves the
stomach, the cecum relaxes and the passage of chyme through
the ileocecal valve increases (gastroileal reflex). This is presumably
a vagal reflex.