PHS 351

GIT

Neural Control of Gastrointestinal Function

Enteric Nervous System

The gastrointestinal tract has a nervous system on its own called the enteric nervous system. It

lies entirely in the wall of the gut, beginning in the esophagus and extending all the way to the

anus. The number of neurons in this enteric system is about 100 million, almost exactly equal to

the number in the entire spinal cord. This highly developed enteric nervous system is especially

important in controlling gastrointestinal movements and secretion.

The enteric nervous system is composed mainly of two plexuses.

1. Outer plexus lying between the longitudinal and circular muscle layers, called the

myenteric plexus or Auerbach’s plexus. This controls mainly the gastrointestinal

movements.

2. Inner plexus, called the submucosal plexus or Meissner’s plexus that lies in the

submucosa. This controls mainly gastrointestinal secretion and local blood flow.

Although the enteric nervous system can function on its own, independently of these extrinsic

nerves, stimulation by the parasympathetic and sympathetic systems can greatly enhance or

inhibit gastrointestinal functions.

Differences between the Myenteric and Submucosal Plexuses

The myenteric plexus consists mostly of a linear chain of many interconnecting neurons that

extends the entire length of the gastrointestinal tract. Because the myenteric plexus extends all

the way along the intestinal wall and because it lies between the longitudinal and circular layers
of intestinal smooth muscle, it is concerned mainly with controlling muscle activity along the

length of the gut. When this plexus is stimulated, its principal effects are:

a. Increased tonic contraction, or “tone,” of the gut wall.

b. Increased intensity of the rhythmical contractions,

c. Slightly increased rate of the rhythm of contraction,

d. Increased velocity of conduction of excitatory waves along the gut wall, causing more

rapid movement of the gut peristaltic waves.

The myenteric plexus should not be considered entirely excitatory because some of its neurons

are inhibitory; their fiber endings secrete an inhibitory transmitter, possibly vasoactive intestinal

polypeptide or some other inhibitory peptide. The resulting inhibitory signals are especially

useful for inhibiting some of the intestinal sphincter muscles that impede movement of food

along successive segments of the gastrointestinal tract, such as the pyloric sphincter, which

controls emptying of the stomach into the duodenum, and the sphincter of the ileocecal valve,

which controls emptying from the small intestine into the cecum.

The submucosal plexus, in contrast to the myenteric plexus, is mainly concerned with controlling

function within the inner wall of each minute segment of the intestine. For instance, many

sensory signals originate from the gastrointestinal epithelium and are then integrated in the

submucosal plexus to help control local intestinal secretion, local absorption, and local

contraction of the submucosal muscle that causes various degrees of infolding of the

gastrointestinal mucosa.

Types of Neurotransmitters Secreted by Enteric Neurons

Neurotransmitter substances that are released by the nerve endings of different types of enteric

neurons includes: (1) acetylcholine and (2) norepinephrine (3) adenosine triphosphate
(4) serotonin (5) dopamine (6) cholecystokinin (7) substance P (8) vasoactive intestinal

polypeptide (9) somatostatin (10) leu-enkephalin (11) metenkephalin, and (12) bombesin.

Acetylcholine most often excites gastrointestinal activity. Norepinephrine almost always inhibits

gastrointestinal activity. The other aforementioned transmitter substances are a mixture of

excitatory and inhibitory agents.

GASTROINTESTINAL HORMONES

Gastrointestinal (GI) hormones are the hormones secreted in GI tract. These hormones are

polypeptides in nature. Major function of these hormones is to regulate the secretory activities

and motility of the GI tract.

CELLS SECRETING THE HORMONES

Enteroendocrine Cells

Enteroendocrine cells are the hormone-secreting cells in GI tract. These are the nerve cells and

glandular cells which are present in the gastric mucosa, intestinal mucosa and the pancreatic

cells.

Neuroendocrine Cells

Enteroendocrine cells which secrete hormones from amines are known as amine precursor

uptake and decarboxylation cells (APUD cells) or neuroendocrine cells. For the synthesis of a

GI hormone, first a precursor substance of an amine is taken up by these cells. Later, this

precursor substance is decarboxylated to form the amine. From this amine, the hormone is

synthesized. Because of the uptake of the amine precursor and decarboxylation of this precursor

substance, these cells are called APUD cells. This type of cells is also present in other parts of

the body, particularly the brain, lungs and the endocrine glands.

Enteroendocrine cells which secrete serotonin are called enterochromaffin cells.

GI hormones include:

Hormone Source of secretion Actions

Gastrin G cells in stomach, TG cells in GI Stimulates gastric secretion and

tract motility
Islets in fetal pancreas. Promotes growth of gastric mucosa
Anterior pituitary Stimulates release of pancreatic
Brain hormones
Stimulates secretion of pancreatic
juice
Stimulates secretion of pancreatic

hormones

Secretin S cells of small intestine Stimulates secretion of watery and

alkaline pancreatic
secretion
Inhibits gastric secretion and
motility
Constricts pyloric sphincter
Increases potency of

cholecystokinin action

Cholecystokinin I cells of small intestine Contracts gallbladder

Stimulates pancreatic secretion with
enzymes
Accelerates secretin activity
Increases enterokinase secretion
Inhibits gastric motility
Increases intestinal motility
Augments contraction of pyloric
sphincter
Suppresses hunger
Induces drug tolerance to opioids

Gastric inhibitory peptide K cells in duodenum and jejunum Stimulates insulin secretion
(GIP) Antrum of stomach Inhibits gastric secretion and

motility

Vasoactive intestinal Stomach Dilates splanchnic (peripheral)

polypeptide (VIP) Small and large intestines blood vessels
Inhibits Hcl secretion in gastric
juice
Stimulates secretion of succus
entericus
Relaxes smooth muscles of
intestine
Augments acetylcholine action on
salivary glands
Stimulates insulin secretion
Glucagon α-cells in pancreas Increases blood sugar level
A cells in stomach
L cells in intestine

Glicentin L cells in duodenum and jejunum Increases blood sugar level

Glucagon-like polypeptide-1 α-cells in pancreas Stimulates insulin secretion

(GLP-1) Brain Inhibits gastric motility

GLP-2 L cells in ileum and colon Suppresses appetite

Somatostatin Hypothalamus Inhibits secretion of growth

D cells in pancreas hormone
D cells in stomach and small Inhibits gastric secretion and
Intestine motility
Inhibits secretion of pancreatic
juice
Inhibits secretion of GI hormones

Pancreatic polypeptide PP cells in pancreas Increases secretion of glucagons

Small intestine Decreases pancreatic secretion

Peptide YY L cells of ileum and colon Inhibits gastric secretion and

motility
Reduces secretion of pancreatic
juice
Inhibits intestinal motility and
bowel passage
Suppresses appetite and food intake
Neuropeptide Y Ileum and colon Increases blood flow in enteric
Brain and autonomic nervous blood vessels
system
(ANS)
Motilin Mo cells in stomach and intestine Accelerates gastric emptying
Enterochromoffin cells in intestine Increases movements of small
intestine
Increases peristalsis in colon
Substance P Brain Increases movements of small
Small intestine intestine
Ghrelin Stomach Promotes growth hormone (GH)
Hypothalamus release
Pituitary Induces appetite and food intake
Kidney Stimulates gastric emptying
Placenta
Digestion and Absorption of Carbohydrate

Enzymes involved in the digestion of carbohydrates are known as amylolytic enzymes. The only

amylolytic enzyme present in saliva is the salivary amylase or

Ptyalin.

In the Stomach

Gastric juice contains a weak amylase, which plays a minor role in digestion of carbohydrates.

In the Intestine

Amylolytic enzymes present in the small intestine are derived from pancreatic juice and succus

entericus.

Amylolytic Enzyme in Pancreatic Juice

Pancreatic juice contains pancreatic amylase

Amylolytic Enzymes in Succus Entericus

Amylolytic enzymes present in succus entericus are maltase, sucrase, lactase, dextrinase and

trehalase.

Final Products of Carbohydrate Digestion

Final products of carbohydrate digestion are monosaccharides, which are glucose, fructose and

galactose.

Glucose represents 80% of the final product of carbohydrate digestion. Galactose and fructose

represent the remaining 20%.

Absorption of Carbohydrates

Carbohydrates are absorbed from the small intestine mainly as monosaccharides, viz. glucose,

galactose and fructose.

Absorption of Glucose

Glucose is transported from the lumen of small intestine into the epithelial cells in the mucus

membrane of small intestine, by means of sodium cotransport. Energy for this is obtained by the

binding process of sodium ion and glucose molecule to carrier protein.

From the epithelial cell, glucose is absorbed into the portal vein by facilitated diffusion.

However, sodium ion moves laterally into the intercellular space. From here, it is transported

into blood by active transport, utilizing the energy liberated by breakdown of ATP.

Absorption of Galactose

Galactose is also absorbed from the small intestine in the same mechanism as that of glucose.

Absorption of Fructose

Fructose is absorbed into blood by means of facilitated diffusion. Some molecules of fructose are

converted into glucose. Glucose is absorbed as described above.

DIGESTION OF PROTEINS

Enzymes responsible for the digestion of proteins are called proteolytic enzymes.

Digestion of proteins does not occur in mouth, since saliva does not contain any proteolytic

enzymes. So, the digestion of proteins starts only in stomach.

In the Stomach

Pepsin is the only proteolytic enzyme in gastric juice

In the Small Intestine

Most of the proteins are digested in the duodenum and jejunum by the proteolytic enzymes of the

pancreatic juice and succus entericus.

Proteolytic Enzymes in Pancreatic Juice

Pancreatic juice contains trypsin, chymotrypsin and carboxypeptidases. Trypsin and

chymotrypsin are called endopeptidases, as these two enzymes break the interior bonds of the

protein molecules.

Proteolytic Enzymes in Succus Entericus

Final digestion of proteins is by the proteolytic enzymes present in the succus entericus. It

contains dipeptidases, tripeptidases and aminopeptidases.

FINAL PRODUCTS OF PROTEIN DIGESTION

Final products of protein digestion are the amino acids, which are absorbed into blood from

intestine.

ABSORPTION OF PROTEINS

Proteins are absorbed in the form of amino acids from small intestine. The levo amino acids are

actively absorbed by means of sodium cotransport, whereas, the dextro amino acids are absorbed

by means of facilitated diffusion. Absorption of amino acids is faster in duodenum and jejunum

and slower in ileum.

DIGESTION OF LIPIDS

Lipids are digested by lipolytic enzymes.

IN THE INTESTINE

Almost all the lipids are digested in the small intestine because of the availability of bile salts,

pancreatic lipolytic enzymes and intestinal lipase.

Role of Bile Salts

Bile salts play an important role in the digestion of lipids

Lipolytic Enzymes in Pancreatic Juice

Pancreatic lipase is the most important enzyme for the digestion of fats. Other lipolytic

enzymes of pancreatic juice are cholesterol ester hydrolase, phospholipase A and phospholipase

B.

Lipolytic Enzyme in Succus Entericus

Intestinal lipase is the only lipolytic enzyme present in succus entericus.

Fatty acids, cholesterol and monoglycerides are the final products of lipid digestion.

ABSORPTION OF LIPIDS

Monoglycerides, cholesterol and fatty acids from the micelles enter the cells of intestinal mucosa

by simple diffusion. From here, further transport occurs as follows:

1. In the mucosal cells, most of the monoglycerides are converted into triglycerides.

Triglycerides are also formed by re-esterification of fatty acids with more than 10 to 12 carbon

atoms. Some of the cholesterol is also esterified.

Triglycerides and cholesterol esters are coated with a layer of protein, cholesterol and

phospholipids to form the particles called chylomicrons. Chylomicrons cannot pass through the

membrane of the blood capillaries because of the larger size. So, these lipid particles enter the

lymph vessels and then are transferred into blood from lymph.

2. Fatty acids containing less than 10 to 12 carbon atoms enter the portal blood from mucosal

cells and are transported as free fatty acids or unesterified fatty acids. Most of the fats are

absorbed in the upper part of small intestine. Presence of bile is essential for fat absorption.