Physiology

The Digestive
System
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Functions of the GI Tract

Motility:
Movement of of food through the GI tract.
Ingestion:
Taking food into the mouth.

Mastication:
Chewing the food and mixing it with saliva.

Deglutition:
Swallowing the food.

Peristalsis:
Rhythmic wave-like contractions that move food through
GI tract.

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Functions of the GI Tract

(continued)

Secretion:
Includes both exocrine and endocrine secretions.
Exocrine:
HCl, H20, HC03-, bile, lipase, pepsin, amylase, trypsin, elastase, and
histamine are secreted into the lumen of the GI tract.

Endocrine:
Stomach and small intestine secrete hormones to help regulate the GI
system.
Gastrin, secretin, CCK, GIP, GLP-1, guanylin, VIP, and
somatostatin.

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Functions of the GI Tract

(continued)

Digestion:
Breakdown of food particles into subunits (chemical
structure change).

Absorption:
Process of the passage of digestion (chemical subunits)
into the blood or lymph.

Storage and elimination:

Temporary storage and elimination of indigestible
food.

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Digestive System (GI)

GI tract divided
into:

Insert fig. 18.2

Alimentary
canal.
Accessory
digestive organs.

GI tract is 30 ft
long and extends
from mouth to
anus.

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Layers of GI Tract
Composed of 4 tunics:
Mucosa.
Submucosa.

Muscularis.
Serosa.

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Mucosa
Lines the lumen of GI tract.
Consists of simple columnar epithelium.

Lamina propria:
Thin layer of connective tissue containing lymph
nodules.

Muscularis mucosae:
Thin layer of smooth muscle responsible for the folds.
Folds increase surface area for absorption.

Goblet cells:
Secrete mucus.
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Submucosa
Thick, highly vascular layer of connective tissue.
Absorbed molecules enter the blood and
lymphatic vessels.
Submucosal plexus (Meissners plexus):
Provide autonomic nerve supply to the muscularis
mucosae.

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Muscularis
Responsible for segmental contractions and
peristaltic movement through the GI tract.
Inner circular layer of smooth muscle.
Outer longitudinal layer of smooth muscle.

Contractions of these layers move food through

the tract; pulverize and mix the food.
Myenteric plexus located between the 2 muscle
layers.
Major nerve supply to GI tract.
Fibers and ganglia from both sympathetic and
parasympathetic nervous systems.
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Serosa

Binding and protective outer layer.

Consists of areolar connective tissue covered with
simple squamous epithelium.
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Regulation of the GI Tract

Extrinsic innervation:
Parasympathetic nervous system:
Vagus and spinal nerves:

Stimulate motility and GI secretions.

Sympathetic nervous system:
Postganglionic sympathetic fibers that pass through
submucosal and myenteric plexuses and innervate GI tract:

Reduce peristalsis and secretory activity.

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Regulation of the GI Tract

(continued)

Enteric nervous system:

Sites where parasympathetic fibers synapse with
postganglionic neurons that innervate smooth muscle.

Submucosal and myenteric plexuses:

Local regulation of the GI tract.

Paracrine secretion:
Molecules acting locally.

Hormonal secretion:
Secreted by the mucosa.

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From Mouth to Stomach

Mastication (chewing):
Mixes food with saliva which contains salivary amylase.
Enzyme that can catalyze the partial digestion of starch.

Deglutition (swallowing):
Begins as a voluntary activity.
Involves 3 phases:
Oral phase is voluntary.
Pharyngeal and esophageal phases are involuntary.
Cannot be stopped.

Larynx is raised.
Epiglottis covers the entrance to respiratory tract.
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From Mouth to Stomach

(continued)

Involuntary muscular contractions and relaxations

in the mouth, pharynx, larynx, and esophagus are
coordinated by the swallowing center in the
medulla.
Esophagus:
Connects pharynx to the stomach.
Upper third contains skeletal muscle.
Middle third contains a mixture of skeletal and smooth
muscle.
Terminal portion contains only smooth muscle.

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Esophagus
Peristalsis:
Produced by a series of
localized reflexes in response
to distention of wall by bolus.

Insert 18.4a

Wave-like muscular
contractions:
Circular smooth muscle
contract behind, relaxes in
front of the bolus.
Followed by longitudinal
contraction (shortening) of
smooth muscle.
Rate of 2-4 cm/sec.

After food passes into

stomach, LES constricts.
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Stomach
Most distensible part of GI tract.
Empties into the duodenum.

Functions of the stomach:

Stores food.
Initiates digestion of proteins.
Kills bacteria.
Moves food (chyme) into intestine.

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Stomach
Contractions of
the stomach
churn chyme.
Mix chyme
with gastric
secretions.
Push food
into intestine.

(continued)

Insert fig. 18.5

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Stomach

(continued)

Gastric mucosa
has gastric pits in
the folds.
Cells that line the
folds deeper in
the mucosa, are
gastric glands.

Insert fig. 18.7

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Gastric Glands
Secrete gastric juice:

Goblet cells: mucus.

Parietal cells: HCl and intrinsic factor.
Chief cells: pepsinogen.
Enterochromaffin-like cells (ECL): histamine and
serotonin.
G cells: gastrin.
D cells: somatostatin.
Stomach: ghrelin.

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HCl Production
Parietal cells secrete
H+ into gastric lumen
by primary active
transport, through
H+/ K+ ATPase
pump.

Insert fig. 18.8

Parietal cells
basolateral
membrane takes in
Cl- against its
electrochemical
gradient, by
coupling its
transport with
HC03-.
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HCl Production

(continued)

HCl production is stimulated:

Indirectly by gastrin.
Indirectly by ACh.

ACh and gastrin stimulate release of histamine.

Histamine:
Stimulates parietal cells to secrete HCl.

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HCl Functions
Makes gastric juice
very acidic.

Insert fig. 18.9

Denatures ingested
proteins (alter
tertiary structure) so
become more
digestible.

Activates
pepsinogen to
pepsin.
Pepsin is more
active at pH of 2.0.

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Digestion and Absorption in the

Stomach
Proteins partially digested by pepsin.
Carbohydrate digestion by salivary amylase is
soon inactivated by acidity.
Alcohol and aspirin are the only commonly
ingested substances absorbed.

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Gastric and Peptic Ulcers

Peptic ulcers:
Erosions of the mucous membranes of the stomach or duodenum
produced by action of HCl.

Zollinger-Ellison syndrome:
Ulcers of the duodenum are produced by excessive gastric acid
secretions.

Helicobacter pylori:
Bacterium that resides in GI tract that may produce ulcers.

Acute gastritis:
Histamine released by tissue damage and inflammation stimulate
further acid secretion.
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Protective Mechanisms of Stomach

Parietal and chief cells impermeable to HCl.
Alkaline mucus contains HC03-.
Tight junctions between adjacent epithelial cells.
Rapid rate of cell division (entire epithelium
replaced in 3 days).
Prostaglandins inhibit gastric secretions.

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Small Intestine
Each villus is a fold in the
mucosa.
Covered with columnar
epithelial cells interspersed
with goblet cells.
Epithelial cells at the tips of
villi are exfoliated and
replaced by mitosis in crypt
of Lieberkuhn.
Lamina propria contain
lymphocytes, capillaries,
and central lacteal.

Insert fig. 18.12

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Absorption in Small Intestine

Duodenum and jejunum:
Carbohydrates, amino acids, lipids, iron, and Ca2+.

Ileum:
Bile salts, vitamin B12, electrolytes, and H20.

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Intestinal Enzymes
Microvilli contain brush border enzymes that are
not secreted into the lumen.
Brush border enzymes remain attached to the cell
membrane with their active sites exposed to the chyme.

Absorption requires both brush border enzymes

and pancreatic enzymes.

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Intestinal Contractions and Motility

2 major types of contractions
occur in the small intestine:
Peristalsis:
Slow movement.
Pressure at the pyloric end of
small intestine is greater than
at the distal end.

Segmentation:
Major contractile activity of
the small intestine.
Contraction of circular
smooth muscle.
Mix chyme.
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Insert fig. 18.14

Contractions of Intestinal Smooth

Muscles
Occur automatically in
response to endogenous
pacemaker activity.
Rhythm of contractions is
paced by graded
depolarizations called
slow waves.

Insert fig. 18.15

Slow waves produced by

interstitial cells of Cajal.
Slow waves spread from 1
smooth muscle cell to
another through nexuses.
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Contractions of Intestinal Smooth Muscles

When slow waves above threshold, it triggers APs
by opening of VG Ca2+ channels.
Inward flow of Ca2+:
Produces the upward depolarization phase.
Stimulates contraction of smooth muscle.

Repolarization:
VG K+ channels open.
Slow waves decrease in amplitude as they are conducted.

May stimulate contraction in proportion to the

magnitude of depolarization.
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Cells and Electrical Events in the

Muscularis
Insert fig. 18.16

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Large Intestine
Outer surface bulges outward to form haustra.
Little absorptive function.
Absorbs H20, electrolytes, several vitamin B complexes, vitamin K,
and folic acid.
Intestinal microbiota produce significant amounts of folic acid and
vitamin K.
Bacteria ferment indigestible molecules to produce short-chain fatty
acids.
Does not contain villi.

Secretes H20, via active transport of NaCl into intestinal

lumen.
Guanylin stimulates secretion of Cl- and H20, and inhibits
absorption of Na+ (minor pathway).
Membrane contains Na+/K+ pumps.
Minor pathway.
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Fluid and Electrolyte Absorption in the Intestine

Small intestine:
Most of the fluid and electrolytes are absorbed by small
intestine.
Absorbs about 90% of the remaining volume.

Absorption of H20 occurs passively as a result of the

osmotic gradient created by active transport.
Aldosterone stimulates NaCl and H20 absorption in the ileum.

Large intestine:
Absorbs about 90% of the remaining volume.
Absorption of H20 occurs passively as a result of the osmotic gradient
created by active transport of Na+ and Cl-.

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Defecation
Waste material passes to the rectum.
Occurs when rectal pressure rises and external anal
sphincter relaxes.
Defecation reflex:
Longitudinal rectal muscles contract to increase rectal
pressure.
Relaxation of internal anal sphincter.

Excretion is aided by contractions of abdominal and

pelvic skeletal muscles.
Push feces from the rectum.

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Structure of Liver

Liver largest internal organ.

Hepatocytes form hepatic plates that are 12 cells thick.
Arranged into functional units called lobules.

Plates separated by sinusoids.

More permeable than other capillaries.

Contains phagocytic Kupffer cells.

Secretes bile into bile canaliculi, which are drained by
bile ducts.

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Structure of Liver

(continued)

Insert fig. 18.20

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Hepatic Portal System

Products of digestion that are absorbed are
delivered to the liver.
Capillaries drain into the hepatic portal vein,
which carries blood to liver.
blood is deoxygenated.
Hepatic vein drains liver.

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Enterohepatic Circulation
Compounds that
recirculate between liver
and intestine.
Many compounds can be
absorbed through small
intestine and enter
hepatic portal blood.
Variety of exogenous
compounds are secreted
by the liver into the bile
ducts.

Insert fig. 18.22

Can excrete these

compounds into the
intestine with the bile.
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Major Categories of Liver Function

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Bile Production and Secretion

The liver produces and secretes 2501500 ml of bile/day.
Bile pigment (bilirubin) is produced in spleen, bone
marrow, and liver.
Derivative of the heme groups (without iron) from hemoglobin.

Free bilirubin combines with glucuronic acid and forms

conjugated bilirubin.
Secreted into bile.

Converted by bacteria in intestine to urobilinogen.

Urobilogen is absorbed by intestine and enters the hepatic vein.
Recycled, or filtered by kidneys and excreted in urine.

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Metabolism of Heme and Bilirubin

Insert fig. 18.23

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Bile Production and Secretion

Bile acids are derivatives of
cholesterol.
Major pathway of cholesterol
breakdown in the body.

(continued)

Insert fig. 18.25

Principal bile acids are:

Cholic acid.
Chenodeoxycholic acid.
Combine with glycine or
taurine to form bile salts.
Bile salts aggregate as
micelles.

95% of bile acids are

absorbed by ileum.
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Detoxification of the Blood

Liver can remove hormones, drugs, and other
biologically active molecules from the blood by:
Excretion into the bile.
Phagocytosis by Kupffer cells.
Chemical alteration of the molecules.
Ammonia is produced by deamination of amino acids in the
liver.
Liver converts it into urea.
Excreted in urine.

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Detoxification of the Blood

(continued)

Inactivation of steroid hormones and drugs.

Conjugation of steroid hormones and xenobiotics
make them anionic.
Can be transported into bile by multispecific organic anion
transport carriers.

Steroid and xenobiotic receptors stimulate production

of cytochrome P450 enzymes.

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Secretion of Glucose, Triglycerides and

Ketones
Liver helps regulate blood glucose concentration
by:
Glycogenesis and lipogenesis.
Glycogenolysis and gluconeogenesis.

Contains enzymes required to convert free fatty

acids into ketone bodies.

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Production of Plasma Proteins

Albumin and most of the plasma globulins
(except immunoglobulins) are produced by
the liver.
Albumin:
Constitutes 70% of the total plasma protein.
Contributes most to the colloid osmotic pressure in the
blood.

Globulins:
Transport cholesterol and hormones.
Inhibit trypsin.
Produce blood clotting factors I, II, III, V, VII, IX, XI.
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Gallbladder
Sac-like organ attached to the inferior surface of
the liver.
Stores and concentrates bile.
When gallbladder fills with bile, it expands.
Contraction of the muscularis layer of the gallbladder,
ejects bile into the common bile duct into duodenum.

When small intestine is empty, sphincter of Oddi

closes.
Bile is forced up to the cystic duct to gallbladder.

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Pancreas
Exocrine:
Acini:

Insert fig. 18.26

Secrete
pancreatic
juice.

Endocrine:
Islets of
Langerhans:
Secrete
insulin and
glucagon.
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Pancreatic Juice
Contains H20, HC03- and digestive enzymes.

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Pancreatic Juice
Complete digestion of food
requires action of both
pancreatic and brush
border enzymes.
Most pancreatic enzymes
are produced as
zymogens.
Trypsin (when activated
by enterokinase) triggers
the activation of other
pancreatic enzymes.

Fig. 18.29

Pancreatic trypsin inhibitor

attaches to trypsin.
Inhibits its activity in the
pancreas.
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Neural and Endocrine Regulation

Neural and endocrine mechanisms modify the
activity of the GI system.
GI tract is both an endocrine gland, and a target
for the action of hormones.

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Regulation of Gastric Function

Gastric motility and secretion are automatic.
Waves of contraction are initiated spontaneously
by pacesetter cells.
Extrinsic control of gastric function is divided into 3
phases:
Cephalic phase.
Gastric phase.
Intestinal phase.

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Cephalic Phase
Stimulated by sight, smell, and taste of food.
Activation of vagus:

Stimulates chief cells to secrete pepsinogen.

Directly stimulates G cells to secrete gastrin.
Directly stimulates ECL cells to secrete histamine.
Indirectly stimulates parietal cells to secrete HCl.

Continues into the 1st 30 min. of a meal.

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Gastric Phase
Arrival of food in stomach stimulates the gastric phase.
Gastric secretion stimulated by:
Distension.
Chemical nature of chyme (amino acids and short polypeptides).
Stimulates G cells to secrete gastrin.
Stimulates chief cells to secrete pepsinogen.
Stimulates ECL cells to secrete histamine.
Histamine stimulates secretin of HCl.

Positive feedback effect.

As more HCl and pepsinogen are secreted, more polypeptides and
amino acids are released.

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Gastric Phase

(continued)

Secretion of HCl is also

regulated by a negative
feedback effect:

Insert. Fig. 18.30

HCl secretion decreases

if pH < 2.5.
At pH of 1.0, gastrin
secretion ceases.
D cells stimulate
secretion of
somatostatin.
Paracrine
regulator to
inhibit secretion
of gastrin.

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Intestinal Phase
Inhibits gastric activity when chyme enters the
small intestine.
Arrival of chyme increases osmolality and
distension.
Activates sensory neurons of vagus and produces an
inhibitory neural reflex:
Inhibits gastric motility and secretion.
In the presence of fat, enterogasterone inhibits gastric motility
and secretion.

Hormone secretion:
Inhibit gastric activity:
Somatostatin, CCK, and GLP-1.
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Enteric Nervous System

Submucosal and myenteric plexuses contain 100
million neurons.
Include preganglionic parasympathetic axons,
ganglion cell bodies, postganglionic sympathetic
axons; and afferent intrinsic and extrinsic sensory
neurons.

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Enteric Nervous System

Peristalsis:
ACh and substance
P stimulate smooth
muscle contraction
above the bolus.
NO, VIP, and ATP
stimulate smooth
muscle relaxation
below the bolus.

(continued)

Insert fig. 18.31

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Paracrine Regulators of the Intestine

Serotonin (5-HT):
Stimulates intrinsic afferents, which send impulses into intrinsic
nervous system; and activates motor neurons.

Motilin:
Stimulates contraction of the duodenum and stomach antrum.

Guanylin:
Activates guanylate cyclase, stimulating the production of cGMP.
cGMP stimulates the intestinal cells to secrete Cl- and H20.
Inhibits the absorption of Na+.

Uroguanylin:
May stimulate kidneys to secrete salt in urine.
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Intestinal Reflexes
Intrinsic and extrinsic regulation controlled by
intrinsic and paracrine regulators.
Gastroileal reflex:
Increased gastric activity causes increased motility of
ileum and movement of chyme through ileocecal
sphincter.

Ileogastric reflex:
Distension of ileum, decreases gastric motility.

Intestino-intestinal reflex:
Overdistension in 1 segment, causes relaxation
throughout the rest of intestine.
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Secretion of Pancreatic Juice

Secretion of pancreatic juice and bile is stimulated by:
Secretin:
Occurs in response to duodenal pH < 4.5.
Stimulates production of HC03- by pancreas.
Stimulates the liver to secrete HC03- into the bile.
CCK:
Occurs in response to fat and protein content of chyme in
duodenum.
Stimulates the production of pancreatic enzymes.
Enhances secretin.
Stimulates contraction of the sphincter of Oddi.
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Digestion and Absorption of

Carbohydrates
Salivary amylase:
Begins starch digestion.

Insert fig. 18.32

Pancreatic amylase:
Digests starch to
oligosaccharides.
Oligosaccharides
hydrolyzed by brush
border enzymes.

Glucose is transported by
secondary active
transport with Na+ into
the capillaries.
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Digestion and Absorption of Protein

Digestion begins in the stomach when pepsin
digests proteins to form polypeptides.
In the duodenum and jejunum:
Endopeptidases cleave peptide bonds in the interior of
the polypeptide:
Trypsin.
Chymotrypsin.
Elastase.

Exopeptidases cleave peptide bonds from the ends of

the polypeptide:
Carboxypeptidase.
Aminopeptidase.
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Digestion and Absorption of Protein

(continued)

Free amino acids

absorbed by
cotransport with Na+.
Dipeptides and
tripeptides
transported by
secondary active
transport using a H+
gradient to transport
them into the
cytoplasm.

Insert fig. 18.33

Hydrolyzed into free

amino acids and then
secreted into the
blood.
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Digestion and Absorption of Lipids

Arrival of lipids in the duodenum serves as a
stimulus for secretion of bile.
Emulsification:
Bile salt micelles are secreted into duodenum to break
up fat droplets.

Pancreatic lipase and colipase hydrolyze

triglycerides to free fatty acids and monglycerides.
Colipase coats the emulsification droplets and anchors
the lipase enzyme to them.
Form micelles and move to brush border.
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Digestion and Absorption of Lipids

(continued)

Free fatty acids, monoglycerides, and lysolecithin

leave micelles and enter into epithelial cells.
Resynthesize triglycerides and phospholipids within cell.
Combine with a protein to form chylomicrons.

Secreted into central lacteals.

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Transport of Lipids
In blood, lipoprotein lipase hydrolyzes triglycerides
to free fatty acids and glycerol for use in cells.
Remnants containing cholesterol are taken to the
liver.
Form VLDLs which take triglycerides to cells.
Once triglycerides are removed, VLDLs are converted
to LDLs.
LDLs transport cholesterol to organs and blood vessels.

HDLs transport excess cholesterol back to liver.

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Absorption of Fat

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