CHAPTER 26
Overview of Gastrointestinal Function & Regulation 431
protects the cells to some extent from the effects of digestive
enzymes. Some digestive enzymes are also actually part of the
brush border, being membrane-bound proteins. These so-
called “brush border hydrolases” perform the final steps of
digestion for specific nutrients.
GASTROINTESTINAL SECRETIONS
SALIVARY SECRETION
The first secretion encountered when food is ingested is saliva.
Saliva is produced by three pairs of salivary glands that drain into
the oral cavity. It has a number of organic constituents that serve
to initiate digestion (particularly of starch, mediated by amylase)
and which also protect the oral cavity from bacteria (such as
immunoglobulin A and lysozyme). Saliva also serves to lubricate
the food bolus (aided by mucins). Saliva is also hypotonic com-
pared with plasma and alkaline; the latter feature is important to
neutralize any gastric secretions that reflux into the esophagus.
The salivary glands consist of blind end pieces (acini) that
produce the primary secretion containing the organic constit-
uents dissolved in a fluid that is essentially identical in its
composition to plasma. The salivary glands are actually
extremely active when maximally stimulated, secreting their
own weight in saliva every minute. To accomplish this, they
are richly endowed with surrounding blood vessels that dilate
when salivary secretion is initiated. The composition of the
saliva is then modified as it flows from the acini out into ducts
that eventually coalesce and deliver the saliva into the mouth.
Na
+
and Cl
- are extracted and K
- and bicarbonate are added.
Because the ducts are relatively impermeable to water, the loss
of NaCl renders the saliva hypotonic, particularly at low
secretion rates. As the rate of secretion increases, there is less
time for NaCl to be extracted and the tonicity of the saliva
rises, but it always stays somewhat hypotonic with respect to
plasma. Overall, the three pairs of salivary glands that drain
into the mouth supply 1000 to 1500 mL of saliva per day.
Salivary secretion is almost entirely controlled by neural
influences, with the parasympathetic branch of the autonomic
nervous system playing the most prominent role (Figure 26–3).
Sympathetic input slightly modifies the composition of saliva
(particularly by increasing proteinaceous content), but has little
influence on volume. Secretion is triggered by reflexes that are
stimulated by the physical act of chewing, but is actually initi-
ated even before the meal is taken into the mouth as a result of
central triggers that are prompted by thinking about, seeing, or
smelling food. Indeed, salivary secretion can readily be condi-
tioned, as in the classical experiments of Pavlov where dogs
were conditioned to salivate in response to a ringing bell by
associating this stimulus with a meal. Salivary secretion is also
prompted by nausea, but inhibited by fear or during sleep.
Saliva performs a number of important functions: it facili-
tates swallowing, keeps the mouth moist, serves as a solvent
for the molecules that stimulate the taste buds, aids speech by
facilitating movements of the lips and tongue, and keeps the
mouth and teeth clean. The saliva also has some antibacterial
action, and patients with deficient salivation
(xerostomia)
have a higher than normal incidence of dental caries. The
buffers in saliva help maintain the oral pH at about 7.0. They
also help neutralize gastric acid and relieve heartburn when
gastric juice is regurgitated into the esophagus.
- and bicarbonate are added.
GASTRIC SECRETION
Food is stored in the stomach; mixed with acid, mucus, and
pepsin; and released at a controlled, steady rate into the
duodenum (see Clinical Box 26–1).
ANATOMIC CONSIDERATIONS
The gross anatomy of the stomach is shown in Figure 26–4.
The gastric mucosa contains many deep glands. In the cardia
and the pyloric region, the glands secrete mucus. In the body
of the stomach, including the fundus, the glands also contain
parietal (oxyntic) cells,
which secrete hydrochloric acid and
intrinsic factor, and
chief (zymogen, peptic) cells,
which se-
crete pepsinogens (Figure 26–5) These secretions mix with
FIGURE 26–2
The structure of intestinal villi and crypts.
(Reproduced with permission, from Fox SI:
Human Physiology
, 10th ed. McGraw-Hill,
2008.)
Simple columnar
epithelium
Lacteal
Capillary network
Goblet cells
Intestinal crypt
Lymph vessel
Arteriole
Venule
Villus