Give
an account of gastrointestinal motility and its control.
Outline:
·
Role of motility in
digestion
·
Intrinsic innervation
·
Types of movement
·
Stomach:
- receptive relaxation
- gastric contractions
- gastric emptying and regulation
·
Small intestine:
- intestinal slow waves
- reflexes
- migrating motor complex
Essay:
Digestion involves both mechanical and chemical breakdown of ingested
material. For these processes to be carried out optimally, the contents must be
held long enough for breakdown and mixing to occur and gastrointestinal contents
must be moved analward to make room for more food and to remove wastes. The
motor activities of the gastrointestinal (GI) tract are regulated by myogenic
mechanisms arising from the intrinsic properties of smooth muscle.
Neurotransmitters and hormones act mainly to modulate this basic muscular
activity.
The smooth muscle cells of the GI tract are long and slender and are
electrically connected to one another through gap junctions, forming a
latticework of smooth muscle bundles enabling it to function as a syncytium.
They are subjected to a continual but slow electrical activity. Slow waves are
slow, undulating changes in the resting membrane potential. Its frequency varies
from 3 per minute in the stomach to 12 per minute in the duodenum. Slow waves
are generated by interstitial cells located in a thin layer between the
longitudinal and circular layers of the muscularis external. The amplitude of
the slow waves can be modulated by activity of the intrinsic and extrinsic
nerves and by hormones and paracrine substances. If the peak of the slow wave
exceeds the threshold potential, action potentials are triggered which enhances
the contractile force of the muscle.
The basic propulsive movement of the GI tract is peristalsis which cause
food to move forward along the tract at an appropriate rate for digestion and
absorption. The initiation of peristalsis begins with the appearance of a
contractile ring around the gut and then moves forward and spreads along the
tube, moving forward any material in front of it. At the same time, the gut
relaxes downstream (receptive relaxation) towards the anus, thus allowing the
food to be propelled easily analward.
The stomach serves as a reservoir for the large volume of food that can
be ingested at a single meal until the food can be processed in the duodenum.
The presence of food in the stomach increases its contractile activity, mixing
food with gastric secretions and grinding it into smaller particles to form
chyme. The gastric contents are delivered to the duodenum at a rate that affords
optimal mixing with the pancreatic-biliary secretions.
When food is passed from the esophagus to the stomach, the contractile
activity of the fundus is inhibited, enabling it to easily accomodate 1-2L of
food. Stretch receptors in the stomach detect the presence of food and initiate
a vagovagal reflex from the stomach to the brain stem and then back to the
stomach. The postganglionic vagal fibers release VIP or nitric oxide which
decreases the rate of vagal firing and hence the tone in the muscular wall of
stomach.
During the cephalic and gastric phases of digestion, the release of
acetylcholine from postganglionic parasympathetic nerve endings and gastrin from
the G cells stimulates gastric contractility. These contractions increase in
force and velocity as they approach the gastroduodenal junction and therefore
the major mixing activity occurs in the antrum of the stomach. As each
peristaltic wave reaches the pylorus, the pyloric sphincter snaps shut, so that
the stomach empties in small squirts. Hence, gastric emptying occurs when the
chyme is decomposed into small enough pieces to fit through the pyloric
sphincter. The rapid contraction of the antrum also propels the chyme back into
the antrum; this movement, called retropulsion, is effective at mixing and
breaking down gastric contents.
The rate at which the stomach empties is regulated by signals from both
the stomach and the duodenum. Increased food volume in the stomach promotes
increased emptying from the stomach. Local excitatory reflexes are initiated by
expansions of the antrum. The duodenal and jejunal mucosa contain receptors that
sense acidity, osmotic pressure, fat digestion products, peptides and amino
acids. Gastric emptying is slowed by hypertonic solutions in the duodenum, a
duodenal pH of below 3.5, and the presence of amino acids and fatty acids. As a
result of these mechanisms, the rate at which chyme enter the small intestine do
not exceed the rate at which the small intestine can process it. Acid is not
dumped into the duodenum more rapidly than it can be neutralized by pancreatic
and duodenal secretions.
The contractile activity of the smooth muscles lining the small intestine
serves to mix the chyme with the digestive juices and bile to facilitate
digestion and absorption and to propel chyme from the duodenum to the colon.
Segmentation is the most common type of intestinal contraction. During
segmentation, about 2cm of the intestinal wall contracts, forcing the chyme back
toward the stomach and toward the colon. When the muscle relaxes, the chyme
returns to the area from which it was displaced. This back-and-forth movement
enables the chyme to become thoroughly mixed with the digestive juices.
Segmentation occurs throughout the digestive period, being generated
continuously by the intestinal slow waves. The higher frequency of segmentation
in the proximal intestine than in the distal intestine propels the chyme slowly
toward the colon. The relative slow rate of net propulsion of chyme in the small
intestine allows time for digestion and absorption.
Peristalic activity of the small intestine increases after a meal and is
caused by the beginning entry of chyme into the duodenum. The gastroenteric
reflex is initiated by distention of the stomach and conducted through the
myenteric plexus from the stomach down along the wall of the small intestine.
Hormones such as CCK, gastrin, insulin and serotonin enhances intestinal
motility while secretin and glucagon inhibit it. The emptying of chyme from the
small intestine into the cecum is regulated by the ileocecal sphincter. When the
cecum is distended, the contraction of the ileocecal sphincter is intensified
and ileal peristalsis is inhibited delaying the emptying of additional chyme
from the ileum.
During the interdigestive period, any undigested food particles left in
the stomach is removed by the migrating motor complex (MMC). The MMC is a
peristaltic wave that repeats every 75 to 90 minutes from the esophagus and
travels through the entire GI tract to the terminal ileum. The MMC sweeps the
small bowel content clean and empty its contents in the colon.