SECRETORY FUNCTIONS OF THE ALIMENTARY TRACT

Guyton & Hall, Chapter 65

I. GENERAL PRINCIPLES OF ALIMENTARY TRACT SECRETION

A. anatomical types of glands
1. mucous cells = goblet cells
a. single-cell mucous glands
b. surface epithelium
c. respond to epithelial stimulation
d. extrude mucus for lubrication to protect surfaces
2. pits - invaginations of the epithelium
a. crypts of Lieberkühn - small intestine
b. deep
c. contain specialized secretory cells
3. tubular glands - stomach and upper duodenum
 A typical tubular gland can be seen in Figure 65-4, which shows an acid
and pepsinogen-secreting gland of the stomach (oxyntic gland).
4. complex glands: (alimentary tract)
a. compound acinous glands
1. salivary glands
2. pancreas
b. liver
c. provide secretions for digestion or emulsification of food
d. lie outside the walls of alimentary tract
e. contain millions of acini lined with secreting glandular cells
f. acini feed into system of ducts that empty into alimentary tract

B. stimulation of alimentary tract glands

 1. mechanical presence of food
2. enteric nervous system
a. tactile stimulation
b. chemical irritation
c. distention of gut wall
3. nervous reflexes stimulate both mucous cells on the epithelial surface
and deeper glands in the mucosa

C. Autonomic Nervous System (ANS) stimulation of secretion

1. parasympathetic stimulation
a. increase rate of secretion
b. upper portion of tract (innervated by glossopharyngeal & Vagus
……………………………Parasympathetic nerves)
1. salivary glands
2. esophageal glands
3. gastric glands
4. pancreas
5. Brunner's glands in the duodenum
c. innervated by vagus & other cranial parasympathetic nerves
d. distal portion of large intestine
e. innervated by pelvic parasympathetic nerves
f. (secretion in the remainder of the small intestine and in the 1st
2/3 of the large intestine occurs mainly in response to
local neural and hormonal stimuli)
2. sympathetic stimulation
a. slight to moderate increase in secretion in some parts
b. constriction of blood vessels - leads to reduced secretion from
glands

D. regulation of glandular secretion by hormones

 1. stomach and intestines
2. from gastrointestinal mucosa
3. stimulated by the presence of food in gut lumen
4. carried by blood to glands
5. increases output of gastric juice and pancreatic juice
6. hormonal stimulation causes constriction of gallbladder to empty bile
7. gastrointestinal hormones are polypeptides or polypeptide derivatives

E. basic mechanism of secretion by glandular cells

1. all are not known
2. nutrient material needed for formation of the secretion must diffuse or
be actively transported from the capillary into the base of the glandular
cell
3. many mitochondria located inside the cell near its base use oxidative
energy for formation of adenosine triphosphate (ATP)
4. energy from the ATP, along with appropriate substrates provided by
the nutrients, is them used for synthesis of the organic substances; this
synthesis occurs almost entirely in the endoplasmic reticulum and
Golgi complex. The ribosomes adherent to the reticulum are
specifically responsible for formation of the proteins that are to be
secreted.
5. secretory materials are transported through the tubules of the
endoplasmic reticulum, passing in about 20 min. all the way to the
vesicles of the Golgi complex, which lies near the secretory ends of the
cells
6. in the Golgi complex, materials are modified, added to, concentrated,
and discharged into the cytoplasm in the form of secretory vesicles,
which are stored in the apical ends of the secretory cells
7. vesicles remain stored until nervous or hormonal control signals cause
the cells to extrude the vesicular contents through the cell's surface -
exocytosis

F. water and electrolyte secretion

 1. nerve stimulation has a specific effect on the basal portion of the cell
membrane to cause active transport of chloride ions to the
interior
2. resulting increase in electronegativity induced inside the cell by excess
chloride ions then causes positive ions also to move to the
interior of the cell
3. excess of both negative and positive ions inside the cell creates an
osmotic force that pulls water to the interior - increasing the
hydrostatic pressure inside the cell and causing the cell itself to swell
4. pressure in the cell results in minute ruptures of the secretory border of
the cell and causes flushing of the water, electrolytes, and
organic materials out of the secretory end of the glandular cell
and into the lumen of the gland

G. lubricating & protective properties of mucus and its importance in the GI tract
1. mucus
a. thick secretion composed mainly of water, electrolytes, and
glycoproteins
b. adherent qualities
c. prevents contact of food particles with the mucosa
d. low resistance for slippage - particles slide
e. causes adherence to fecal matter
f. strongly resistant to digestion by the GI enzymes
g. amphoteric properties - pH buffer
h. contain bicarbonate ions - neutralizes acids
2. mucus allows easy slippage of food along GI tract and prevents
excoriative or chemical damage to epithelium
II. SECRETION OF SALIVA
A. principal glands (tubuloalveolar)
1. parotid secretory end-pieces serous cells

2. submandibular
3. sublingual intercalated duct

4. buccal striated duct mucous cells

5. lingual glands excretory duct

6. other glands
B. daily rate of secretion = 1000 - 1500 ml
C. types of saliva secretion
1. serous secretion
a. ptyalin
1. an α-amylase
2. salivary amylase
b. for digestion of starches
c. produced by parotid, submandibular and sublingual glands
2. mucous
a. mucin (glycoprotein)
b. for lubrication
c. produced by buccal, submandibular and sublingual glands

Mucous Cells
3. muramidase
a. lysozyme
b. acts on muramic acid of the cell wall
c. bacteria can not maintain osmotic pressure
 Serous Cells
** 4. lingual lipase
a. fat digesting enzyme
b. digest milk fat
c. important in infants

Na

HCO3-
Cl-
K+

5. lactoferrin
a. protein
b. binds with iron
6. epidermal growth factors
7. immunoglobulin - IgA
8. ABO blood factors (in persons who are secretors)
D. pH of saliva = 6-7.4 (excellent for ptyalin activity)
E. ions in saliva:
1. potassium (30 mEq/L, 7X plasma)
2. bicarbonate ion (50 - 70 mEq/L, 2-3X plasma)
3. sodium & chloride (15 mEq/L, 1/7 - 1/10 of plasma)
4. concentration of ions change with maximal salivation and
aldosterone secretion
 F. stages of salivary secretion:
1. stage 1 - from acini
a. secrete primary secretion
b. contains ptyalin and/or mucin
c. ions - similar to extracellular fluid
2. stage 2 - from salivary ducts (ductile) (amylase)
a. Na+ actively reabsorbed from all salivary ducts
b. K+ actively exchanged for sodium
Therefore, the sodium ion concentration of the saliva becomes
greatly reduced, whereas the potassium ion concentration becomes
increased. However, there is excess sodium reabsorption compared
with potassium secretion, which creates electrical negativity of
about −70 millivolts in the salivary ducts; this negativity in turn
causes chloride ions to be reabsorbed passively. Therefore, the
chloride ion concentration in the salivary fluid falls to a very low
level, matching the ductal decrease in sodium ion concentration.

c. chloride passively reabsorbed

d. bicarbonate ion secreted by ductal epithelium into the lumen of
………………………….the duct.

End
Piece

Cl-
HCO3-
Na+
K+

3. oral hygiene
 a. flow of saliva helps to wash away pathogenic bacteria and food
b. contains antibacterial agents:
1. thiocyanate ions
2. proteolytic enzymes - ex. Lysozyme
3. protein antibodies
G. nervous regulation of salivary secretion
1. salivary glands controlled mainly by parasympathetic nervous signals
from the superior and inferior salivatory nuclei of brain stem
a. at juncture of medulla and pons
b. excited by taste and tactile stimuli
c. excitation from tongue, mouth and pharynx
d. "sour taste" = sapid taste" & smooth objects  copious
secretion of saliva
e. rough objects  reduce/inhibit salivation
2. salivatory nuclei and appetite area of anterior hypothalamus effect
salivation (smell/eat favorite food, salivation is greater)
3. reflexes from stomach and upper intestines
a. irritating foods
b. nausea
c. GI abnormality
4. sympathetic stimulation BF saliva (hence dry mouth)
a. less powerful than parasympathetic
b. originates from the superior cervical ganglia
c. travels along blood vessels to salivary glands
5. blood supply to glands
a. acts as secondary factor
b. secretion requires nutrition
c. parasympathetic nerve signals dilate blood vessels
d. saliva dilates blood vessels
e. kallikrein
1. dilates vessels
 2. splits a blood protein to bradykinin (vasodilator)
H. functions of saliva:
1. chewing
2. swallowing
3. speech
4. dissolves components of food
5. enhances taste
6. cleans oral tissue (brushing teeth is still necessary)

III. ESOPHAGEAL SECRETION

A. mucoid
B. provide lubrication for swallowing
C. simple mucous glands in body of esophagus
D. compound mucous glands in the gastric end and initial portion of esophagus
1. prevents mucosal excoriation (initial segment)
2. protect esophageal wall from digestion by gastric
juices ( distal segment)
E. initiates starch digestion
F. aids in excretion of heavy metals
1. iron
2. lead
3. others
IV. GASTRIC SECRETION
A. mucous-secreting cells
1. line entire surface of stomach between glands
2. secret viscid mucus - mainly insoluble
3. coats mucosa with gel layer of mucus
4. protection for stomach - lubrication for food
5. alkaline pH
6. stimulus:
a. food
b. irritation of mucosa
Cells within the Gastric Glands and their Secretory Products
Gland Area Cell Secretory Products

Cardiac Mucous Mucus, HCO3-, pepsinogens

(group II)
Endocrine
Oxyntic Parietal (oxyntic) HCL, intrinsic factor
Chief Pepsinogens (groups I and II)
Mucous neck Mucus, HCO3 -, pepsinogens
(group I and II)
Enterochromaffin Serotonin
Endocrine
Pyloric Mucous Mucus, HCO3-, pepsinogens
(group II)
G cell Gastrin
Enterochromaffin Serotonin
Other endocrine

B. tubular glands
1. oxyntic (gastric) glands - body and fundus of stomach (80% of stomach)
a. acid forming
b. secrete HCl, pepsinogen, intrinsic factor and mucous
c. cell types:
1. mucous neck cells
a. mucus
b. pepsinogen
2. peptic (chief) cells
a. pepsinogen
1. several different types
2. activated by HCl
3. forms pepsin
a. active proteolytic enzyme
b, active in high acid environment
c. pH 1.8 -3.5 optimum
d. > pH 5, is inactive
e. necessary for protein digestion
 Gastric Surface
Pit Mucous Cells
Oxyntic
Cell
Isthmus
Mucous Neck
Cells
Neck
Endocrine
Cell

Base Chief Zymogenic

Cells
3. parietal (oxyntic) cells
a. HCL
1. pH = 0.8
2. formed in membranes of canaliculi
3. fig. 64-6
b. intrinsic factor - B12
d. other enzymes:
1. gastric lipase
a. = tributyrase
b. acts on butterfat
2. gastric amylase
a. digestion of starches
b. minor role
3. gelatinase - liquefy proteoglycans in meats
4. intrinsic factor
a. essential for vit. B12 absorption in ileum
b. deficiency = pernicious anemia
2. pyloric glands - antral portion of stomach
 a. mucous (lubrication)
b. pepsinogen
c. gastrin - controls gastric secretion
d. cell types:
1. structurally similar to oxyntic glands
2. few peptic cells
3. no parietal cells
4. mostly mucous cells

C. regulation of gastric secretion by nervous and hormonal mechanisms

1. acetylcholine
a. bind with receptor on secretory cells
b. excites all secretory cells in gastric glands
c. released at parasympathetic nerve endings with vagi stimulation

2. gastrin
a. bind with receptor on secretory cells
b. stimulates secretion of acid by parietal cell
c. little effect on other cells
d. secreted by gastrin cells (G cells) in pyloric glands

e. large peptide
1. 6 different molecules
2. 4 AA at end are active site

f. secreted in 2 forms:
1. G-34
a. large form
b. contains 34 amino acids

2. G-17
 a. smaller form
b. contains 17 amino acids
c. most abundant form
g. absorbed into the blood
h. carried to oxyntic glands in body of stomach
i. stimulates parietal cells (strongly) and peptic cells (weakly)
j. increases HCl secretion by parietal cells
k. stimulated by vagi
l. may  HCO3- release from liver and pancreas
m. gastric motility
n.  pressure of LES
o. open ileocecal valve

3. histamine
a. bind with receptor on secretory cells
b. stimulates secretion of acid by parietal cell
c. little effect on other cells
d. amino acid derivative
e. formed continually in gastric mucosa
f. in the presence of acetylcholine will enhance acid secretion
g. necessary cofactor for exciting significant acid secretion
h. type H2 receptors on parietal cells

4. other substances that stimulate gastric secretions:

a. amino acids
b. caffeine
c. alcohol
d. food in stomach - causing distention

Neurocrine
Paracrine Endocrine

Histamine ACH Gastrin

H Atropine
2
Ca++
Cimetidine Proglumide

camp
ATP pK ADP

protein protein

5. nervous stimulation:
a. ½ of signals originate in the dorsal motor nuclei of vagi
1. pass by way of vagus nerves
2. to enteric nervous system
3. then to gastric glands
4. release ACH
b. ½ of signals are generated by local reflexes
1. occur within the wall of stomach
 2. release ACH
c. in pyloric glands
1. intermediate neuron serves as final path
2. secretes gastrin-releasing peptide as neuro-transmitter
3. probably the peptide bombesin

6. signals originate in
a. brain
b. stomach
1. long vagovagal reflexes
a. from stomach mucosa to brain stem
b. back to stomach through vagus nerves
2. short reflexes
a. originate locally
b. transmitted through local enteric nervous system
7. stimuli:
a. distention of stomach
b. tactile stimuli on surface of stomach mucosa
c. chemical stimuli
1. amino acids
2. peptides
3. acid
d. when acetylcholine, gastrin, and histamine are all present,
copious amounts of acid are secreted

D. regulation of pepsinogen secretion

 1. stimuli:
a. stimulation of peptic cells by ACH from vagus nerves
& enteric nerves
b. stimulation of peptic secretion by acid in stomach (indirectly)
c.  acid   pepsinogen secretion
2. when pH falls below 3.0 , stimulation for gastric secretion becomes
blocked   pepsinogen secretion
3. pH 3.0 = optimal pH for peptic enzymes

E. phases of gastric secretion (phases fuse together)

1. cephalic phase
a. occurs before food enters the stomach
b. results from:
1. sight
2. smell
3. thought
4. taste
c. appetite   stimulation
d. neurogenic signals originate in cerebral cortex or appetite
centers of amygdala or hypothalamus
e. transmitted through the dorsal motor nuclei of vagi to stomach
f. accounts for about 20% of gastric secretion
2. gastric phase
a. food in stomach excites:
1. vagovagal reflexes
2. local enteric reflexes
3. gastrin mechanism
b. accounts for 70% of gastric secretion
3. intestinal phase
a. food in small intestine stimulates stomach to secret gastric juice
 b. duodenal mucosa secretes gastrin in response to distention or
chemical stimuli
c. amino acids and other hormones reflexes play minor roles in
secretion of gastric juice

Stimulation of Gastric Acid Secretion

Phase Stimulus Pathway Stimulus to

parietal
cell

Cephalic Chewing, vagus n. to:

Swallowing 1. Parietal ACH
cells
2. G cells Gastrin

Gastric Gastric Local and

vagovagal
reflexes to:
1. Parietal ACH
cells
2. G cells Gastrin

Intestinal Protein 1. Intestinal G Gastrin

digestion cells
products in 2. Intestinal enterooxyntin
F. inhibition of gastric secretion by intestinal factors
1. chyme stimulates gastric secretion during intestinal phase of secretion
and inhibits during gastric phase
a. presence of food in small intestine initiates an
 enterogastric reflex
b. reflex inhibits stomach secretion that is initiated by:
1. distention of small bowel
2. presence of acid in upper intestine
3. presence of protein breakdown products
4. irritation of mucosa
2. presence of acid, fat, protein breakdown products, hyperosmotic or
hypo-osmotic fluids, or irritating factor in upper small intestine
causes release of several hormones:
a. secretin - opposes stomach secretion
b. gastric inhibitory peptide
c. vasoactive intestinal polypeptide
d. somatostatin
3. inhibition of gastric secretion by intestinal factors slows the release of
chyme from the stomach when small intestine is already filled
4. during interdigestive period small amount of nonoxyntic gastric juice
is secreted
a. mainly mucus
b. contains little pepsin
c. almost no acid
d. increases with emotional stimuli
G. chemical composition
1. gastrin
a. polypeptide
b. terminal five amino acids
same as cholecystokinin
c. activity resides in terminal 4 amino acids
2. cholecystokinin
a. polypeptide
b. terminal five amino acids same as gastrin
c. activity resides in terminal 8 amino acids
 3. secretin
a. polypeptide
c. all the amino acids in secretin are essential

4. enzymes -secreted in inactive or zymogenetic form

a. pepsin
1. proteolytic enzyme
2. pepsinogen (zymogenetic) major enzyme
3. pepsinogen  HCl pepsin
4. found in peptic and mucous cells of the gastric glands
5. pepsinogen is activated by HCl
6. pH < 3.0 is needed for activation of pepsinogen
b. gelatinase - helps to liquefy
c. gastric lipase
1. tributyrase (AKA)
2. acts on tributyrin (=butterfat)
3. does not breakdown triglycerides
d. gastric amylase - has minor role in digestion of starch
e. renin
1. chymosin (AKA)
2. not the same as in the kidney
3. not part of the human system
4. originally thought to be in the human GI tract
5. casein (a protein)  paracasein
6. HCl produces curdling of mile in humans in order to
prolong time of in the stomach for protein
digestion
Mechanisms for Inhibition of Gastric Acid Secretion

Region Stimulus Mediator Inhibit Inhibit

Gastrin Acid
Release Secretion

Antrum Acid (pH<3.0) None, direct +

Duodenum Acid Secretin + +

Bulbogastrone + +
Nervous reflex +

Duodenum Hyperosmotic Unidentified +

Jejunum solutions enterogastrone

Fatty acids, Gastric + +

inhibitory peptide
Monoglycerides Cholecystokinin + +
Unidentified enterogastrone +
V. PANCREATIC SECRETION
A. pancreas
1. weighs about 100 gms
2. "sweet breads" = delicacy
3. secretes 1200 - 1500 ml/day of clear, odorless fluid
4. 7.7 - 8.2 pH, alkaline for the neutralization of acid form the GI tract
5. function:
a. endocrine
1. insulin
2. glucagon
3. somatostatin
b. exocrine - GI secretions
6. watery HCO- solution
a. CO2 is taken up by pancreas and mixed with H2O
b. Na and H move counter
c. Na is added to bicarb solution
B. pancreatic enzymes - secreted by acini
1. proteolytic enzymes:
a. synthesized in the inactive forms trypsinogen,
chymotrypsinogen and procarboxypolypeptidase
b. become active in the intestinal tract
c. trypsinogen
1. endopeptidase activated by enterokinase (secreted
by intestinal mucosa) and trypsin to form trypsin
 Trypsinogen

Enterokinase

Proelastase Chymotrypsinogen

Trypsin

Elastase Chymotrypsin

Procarboxypeptidase Carboxypeptidase

d. proelastase (endopeptidase)
e. chymotrypsinogen
1. endopeptidase
2. is activated by trypsin to from chymotrypsin
f. procarboxypolypeptidase
1. A&B forms
2. is activated by trypsin to form carboxypolypeptidase
3. exopeptidase
g. trypsin
1. most abundant
2. split whole and partially digested proteins into peptides
but does not cause release of individual amino
acids
 g. chymotrypsin
1. split whole and partially digested proteins into peptides
but do not cause release of individual amino
acids
h. carboxypolypeptidase
1. split some peptides into individual amino acids
i. elastases
j. nucleases
k. trypsin inhibitor
1. stored in cytoplasm of glandular cells
2. prevents activation of trypsin
3. prevents digestion of pancreas (autodigestion)
4. w/o trypsin inhibitor, pancreatic secretions digest the
entire pancreas within a few hours = acute
pancreatitis
a. often lethal
b. shock
c. not lethal - pancreatic insufficiency
l. ribonucleases & deoxyribonucleases
1. breaks down RNA & DNA
2. prevents ingestion & absorption of
DNA & RNA supplements
2. pancreatic amylase
a. digests carbohydrates
b. hydrolyzes
1. starches
2. glycogen
3. other carbohydrates
4. not cellulose
c. forms disaccharides and few trisaccharides
3. pancreatic fat digestion
 a. pancreatic lipase
1. main enzyme for fat digestion
2. hydrolyzes fat into fatty acids and monoglycerides
3. needs bile and calcium ions
4. main enzyme that attacks fat
b. cholesterol esterase
1. main enzyme for fat digestion
2. hydrolyzes cholesterol esters
c. phospholipase
a. main enzyme for fat digestion
b. splits fatty acids from phospholipids
**** c. only non-proteolytic enzyme that requires
activation by trypsin
C. secretion of bicarbonate ions
1. secreted by epithelial cells of small ductules and larger ducts leading
from the acini
2. mechanism:
a. CO2 diffuses to interior cell from blood
b. CO2 combines with water to form carbonic acid ( in the
presence of carbonic anhydrase)
c. carbonic acid dissociate into bicarbonate ion and hydrogen ions
d. bicarbonate ions are actively transported into lumen of duct
e. hydrogen ions are exchanged with sodium ions in the blood
f. sodium ions are transported into pancreatic duct - providing
electrical neutrality for secreted bicarbonate ions
g. movement of sodium and bicarbonate ions from blood to lumen
creates osmotic gradient
 h.
water moves into pancreatic
duct to create isosmotic

bicarbonate
solution
D. regulation of
pancreatic secretion
1.
stimuli:
a. diet
- change in diet may effect
secretions
b.
acetylcholine
1. released from parasympathetic vagus nerve endings
2. released from other cholinergic nerves in enteric
nervous system
3. stimulate acinar cells more than ductal cells
c. cholecystokinin
1. released by duodenal and upper jejunal mucosa
2. stimulated by the presence of food
3. stimulate acinar cells more than ductal cells

d. secretin
1. released by duodenal and upper jejunal mucosa
2. stimulated by highly acid food in small intestine
3. stimulates large quantities of sodium bicarbonate by
ductal epithelium
4. almost no stimulation of enzyme secretion
 2. multiplicative effects - the effects of stimuli occurring simultaneously
is multiplied vs. the stimuli occurring alone
E. phases of pancreatic secretion - same as gastric secretion
1. cephalic phase
a. vagus nerve
b. smell, sight, thought are stimuli
c. same nervous signals that cause secretion in the stomach
d. acetylcholine released by vagal nerve endings
e. produce 20% of total pancreatic secretion
2. gastric phase
a. same nervous (vagus) signals that cause secretion
in the stomach
b. acetylcholine released by vagal nerve endings
c. produce 5-10% of total pancreatic secretion
3. intestinal phase
a. main control
b. stimulated by chyme entering intestine
** c. secretin (important one)
1. stimulates copious amounts of secretion
2. polypeptide with 27 a.a., mol. wt. = 3400
3. present in S cells of mucosa of upper small intestine
4. prosecretin = inactive form
5. chyme of < pH 4.5 entering the duodenum stimulates
release and activation of secretin
6. HCl, major stimulator of secretin release
7. secretin enters blood and travels to pancreas
8. causes pancrease to secrete bicarbonate ion in water
and sodium fluid with few enzymes

9. as more acid enters the duodenum, more bicarb is

released
 10. provides neutral or slightly alkaline environment for
optimum pancreatic enzyme activity
d. cholecystokinin stimulates enzyme secretion
1. polypeptide with 33 amino acids
2. stimulated by the presence of food in the upper
small intestine
a. proteoses and peptones - protein breakdown
products
b. long-chain fatty acids
c. HCl
3. released from I cells in the mucosa of the duodenum
and upper jejunum
4. travels via the blood to pancreas
5. stimulates secretion of large quantities of digestive
enzymes by the acinar cells
6. accounts for 70 - 80% of pancreatic enzyme secretion
after a meal

VI. SECRETION OF BILE BY THE LIVER

A. bile

Bile Salts = steroid acids

precursor = cholesterol

cholic acid chenodeoxycholic acid

glycine or taurine

glycocholic acid
taurocholic acid
1. secreted @ of 600-1200 ml/day
2. contains:
a. no digestive enzymes
b. water
c. bile salts
d. bilirubin
e. cholesterol
f. fatty acids
g. lecithin
h. sodium
i. potassium
j. calcium
k. chloride
l. bicarbonate ion
3. components of bile become more concentrated with secretion
4. starts to empty about 30 minutes after a meal
5. purpose:
a. production of micelles for aiding absorption
b. fat digestion
1. emulsification of large fat particles - increase
surface area
2. transport and absorption of digested fat end products via
micelles to and through intestinal mucosal membrane
c. excretion of blood waste products
1. bilirubin - end-product of hemoglobin destruction
2. cholesterol
 B.
stages of bile
secretion

1.
initial
portion

a. secreted by liver hepatocytes

b. contains large amounts of bile acids, cholesterol, and
organic constituents
c. secreted into the minute bile canaliculi
2. second stage
a. bile flows peripherally toward the interlobular septa
b. canaliculi empty into terminal bile ducts and them larger ducts
c. bile flows to hepatic duct and common bile duct
d. bile empties into duodenum or is diverted to gallbladder
via cystic duct
e. as bile flows through ducts second portion is added
1. watery
 2. sodium and bicarbonate ions secreted by secretory
epithelial cells lining ductules
3. increases bile by 100%
4. stimulated by secretin

C. storage and concentration of bile

1. continual secretion of bile is stored in the gallbladder until

needed in the duodenum

2. maximum volume of gallbladder is 30-60 ml

3. up to 12 hours of bile secretion can be stored in the gallbladder

4. gallbladder concentrates bile that is stored

5. bile constitutes:

a. bile salts - accounts for half of total solutes

b. bilirubin
c. cholesterol
d. lecithin
e. electrolytes of plasma
6. in the concentrating process of the gallbladder, large portions of
electrolytes (except calcium ions) are reabsorbed by the
gallbladder mucosa
D. emptying of the gallbladder
1. stimulated by food in upper GI tract
2. rhythmical contractions occur in the wall of the gallbladder
 3. sphincter of Oddi relaxes
a. cholecystokinin causes relaxation
b. peristaltic waves from gallbladder
c. relaxation phase of intestinal peristaltic waves
4. cholecystokinin - most potent stimulus of gallbladder contractions
5. stimulated by ACH-secreting nerve fibers from vagi and enteric
nervous system
6. in the presence of fat, the gallbladder will empty in about 1 hour
7. 90% of all bile is reabsorbed in distal ilium
E. bile salts
1. 0.6 grams of bile salts formed daily
2. cholesterol, from diet or formed by the liver, is precursor of bile salts
3. actions of bile salts:
a. detergent action on fat particles
1. decreases surface tension
2. allows for agitation to break fat globules
3. emulsifying or detergent function
b. help with absorption
1. fatty acids
2. monoglycerides
3. cholesterol
4. other lipids
5. form micelles
F. secretion of cholesterol; most common gallstone formation
1. bile salts are formed in the hepatic cells from cholesterol
2. 1-2 gm/day cholesterol is secreted into the bile
3. cholesterol is insoluble in water
4. in the presence of bile salts and lecithin, cholesterol is converted into
micelles, which are soluble
5. gall stones:
a. precipitated cholesterol
 b. too much water absorption from bile
c. too much absorption of bile salts and lecithin from bile
d. inflammation of the epithelium of the gallbladder (ex. infection)
e. increased cholesterol due to high fat diet &/or no-fat diet

VII. SECRETIONS OF THE SMALL INTESTINE

A. Brunners glands
1. compound mucous glands
2. located in first few centimeters of duodenum
3. mainly between the pylorus and the papilla of Vater
4. secrete alkaline mucus (bicarbonate ions - neutralize acid)
5. stimulated by:
a. tactile stimuli or irritation of mucosa
b. vagal stimulation
c. gastrointestinal hormones (mainly secretin)
d. secretin is major stimulus for Brunner's glands
6. function - protection of the duodenal wall from digestion by
gastric juice
B. Crypts of Lieberkhn
1. small pits
2. located throughout small intestine
3. lie between the intestinal villi
4. covered with epithelium containing 2 cell types:
a. goblet cells
1. secrete mucus
2. protect intestinal surfaces
b. enterocytes
1. secrete water and electrolytes (in the crypts)
2. reabsorb water and electrolytes with the end products of
digestion (over the surfaces of the villi)
5. secretions are formed by enterocytes of the crypts @ 1800 ml/day
 6. secretions are almost pure extracellular fluid with pH of 6.5 - 7.5
7. secretion do not contain enzymes
8. fluid from crypts to villi supplies a watery vehicle for absorption
9. mechanism of secretion:
a. active secretion of chloride into crypts
b. active secretion of bicarbonate ions
c. formation of electrical drag of sodium ions
d. osmotic movement of water

C. enzymes of the small intestine

1. final digestion occurs by membrane bound enzymes
2. peptidases
3. enzymes for splitting disaccharides into monosaccharides:
a. sucrase
b. maltase
c. isomaltase =  dextrinase
d. lactase
4. intestinal lipase
5. glucoamylase - breaks maltoligosaccharides
6. tripeptidases
7. dipeptidase
8. lie in brush border
9. catalyze hydrolysis of food on the outside surface of the microvilli
before absorption - absorption - provides aqueous solution
for absorption
10. lifespan of intestinal epithelia cell = 5 days
VIII. SECRETIONS OF THE LARGE INTESTINE
A. has many crypts of Lieberkhn with many goblet cells
B. no villi
C. almost no enzymes
D. epithelial cells are mostly mucous cells that secrete only mucus
E. mucus - main secretion
1. protects against excoriation
2. large amounts of bicarbonate ions
3. regulated by tactile stimulation of mucous cells and local nervous
reflexes to mucous cells in crypts of Lieberkhn
4. secretion with parasympathetic (with extreme parasympathetic
stimulation, ropy mucus bowel movement can occur @ of 1 every
30 min)
5. protects bowel from bacterial activity and provides adherent medium
for fecal matter

F. irritation (enteritis)
1. secretion of large quantities of water and electrolytes with normal
alkaline mucus
2. acts to dilute irritating factors and cause rapid movement of feces
toward anus
3. diarrhea - protective mechanism; loss of large quantities of water and
electrolytes as irritant factor is washed away
FACTORS AFFECTING GALLBLADDER EMPTYING & BILE SYNTHESIS &
SECRETION

Phase of Stimulus Mediating Factor Response

Digestion

Cephalic taste and smell Impulses in branches rate of

of food; food in of vagus nerve; gallbladder
mouth and Gastrin?
emptying
pharynx

Gastric gastric impulses in branches rate of

distension of vagus nerve: gallbladder
Gastrin? emptying

Intestinal fat digestion cholecystokinin rate of

products in bladder emptying;
duodenum  rate of bile acid
secretion

acid in secretin rate of

duodenum secretion of
 bicarbonate-
rich fluid by the
bile duct
epithelium (this
effect strongly
potentiated by
cholecystokinin)

Absorption of High concentration stimulation of

bile acids in of bile acids in portal bile acid secretion
the distal part blood inhibition of bile
of ileum acid synthesis

Inter- low rate of low concentration of stimulation of

digestive release of bile bile acids in portal bile acid
synthesis;
inhibition of bile
acid secretion