Chapter Ten
Gastrointestinal physiology
1
Digestive System
• The digestive system consisting of the digestive tract (GIT)
and accessory organs
• The alimentary canal or gastrointestinal (GI) tract is
composed of the organs mouth, pharynx, esophagus,
stomach, small intestine, large intestine and anus
• Accessory digestive organs – teeth, tongue and glandular
organs, such as the salivary glands, liver, and pancreas
2
The Components of the Digestive System
3
Functions of gastrointestinal Tract
• Ingestion: taking materials into the digestive tract
• Propulsion: swallowing and peristalsis
–Peristalsis – waves of contraction and relaxation of muscles
in the organ walls
• Mechanical digestion: chewing, mixing, and churning food
• Chemical digestion: catabolic breakdown of food with the
help of enzymes
• Absorption: movement of nutrients from the GI tract lumen
into the blood or lymph
• Secretion: release of water, acids, enzymes, buffers, and salts
• Defecation: elimination of indigestible solid wastes
4
Layers of the GIT
1. Mucosa: Inner layer of the luminal surfaces
- protection, secretion & absorption
2. Submucosa: blood & lymph vessels, nerve plexuses
- necessary for motility + secretion
3. Muscularis externa : contains circular & Longitudinal
smooth muscle, mix and propel the chyme
Circular : Contraction causes a decrease in diameter of the
lumen of the GI tract
Longitudinal : Contraction causes shortening of a segment
of the GI tract
4. Serosa: outer most protective layer, consists of CT
• Secretes a slippery fluid that lubricates and prevents
friction
5
6
Receptors of the GIT
• Mechano- ,osmo- and chemo receptors respond to stretch and
pH, presence of substrate, and end products of digestion
respectively
• They initiate reflexes that:
–Activate or inhibit digestive glands
–Mix lumen contents and move them along
7
Innervation of the GI tract
1. Extrinsic NS
• originate outside the digestive tract and innervate the various
digestive organs
= parasympathetic and sympathetic nervous systems
a. Parasympathetic nervous system
• Excitatory on the functions of the GI tract
• Promotes digestive & absorptive processes
8
b. Sympathetic nervous system
• Inhibitory on the functions of the GI tract
• Slows processes
2. Intrinsic NS
• Lie entirely within the digestive tract wall
=enteric nervous system
• Coordinates and relays information from the
parasympathetic and sympathetic nervous systems to the GI
tract
• Controls motility and secretion
• Shows local, intrinsic regulation
9
a. Myenteric plexus (Auerbach’s plexus)
• Primarily controls the motility of the GI smooth muscle
b. Submucosal plexus (Meissner’s plexus)
• Primarily controls secretion
• Types of Neurotransmitters: Several e.g., Ach, NE, E, NO
vasoactive intestinal peptide (VIP), etc.
10
GI Motility
A. Peristaltic (propulsive) movements:
• Consists of a progressive wave of strong contraction followed
by relaxation
• The main stimulus to cause peristalsis is distension (stretch)
of the gut wall.
Propel or push the contents forward through the digestive tract
• Transit of food through the esophagus is rapid
• In the small intestine—the main site of digestion and
absorption—the contents are moved forward slowly
allowing time for the breakdown and absorption of food
11
a. Mixing movements:
• Occurs due to local contractions taking place in SI segments
Mixing food with the digestive juices (promote digestion of
the food)
Facilitate absorption by exposing all parts of the intestinal
contents to the absorbing surfaces of the digestive tract
The law of the gut movement always occurs from mouth-
to- anus unless pathological
12
Digestion
• Two parts:
1. Mechanical digestion:
• Is the physical grinding of food to smaller units without
altering their chemical composition.
• Helps to expose and increase the surface area for enzymatic
attacks
2. Chemical digestion:
• Is hydrolytic breakdown of nutrients (CHO, fat, protein etc)
by different chemical reactions into simpler forms.
• This processes changes the original composition of the
nutrient.
13
The Mouth (oral cavity)
• The entrance to the digestive tract
• Parts of the oral cavity: cheeks, lip, tongue, hard and soft
palate, teeth
• Is responsible for mechanical digestion of solid food by
mastication
Functions of chewing are:
• To grind and break food up into smaller pieces
–Facilitate swallowing
–Increase the food surface area on which salivary enzymes
will act
• To mix food with saliva
• To stimulate the taste buds
14
Saliva
• Three major pairs of salivary glands
a. Parotid glands (25%)
Secrete mainly serous watery fluid rich in ptyalin
b. Submandibular glands (70%)
Produce both serous and mucous fluid
c. Sublingual glands (~5%)
Secrete mainly thick mucous with little serous fluid
• Components:
–H2O (99.5%)
–Electrolytes(principally Na+, Cl-, and HCO3
-), antibodies
(IgA) , lysozyme, amylase, thiocyanate, mucus..(0.5%)
–Total secretion = about 1-1.5 L/day
15
Saliva functions
a. Digestion: CHO-digestion begins in saliva . The enzyme
ptyalin breaks starch- to-maltose. Lingual lipase begins
fat digestion in the mouth.
b. Protection: has anti-microbial actions (contains IgA,
lysozyme & thiocyanate that kills microbes).
c. Involved in speech: Clear & fluent articulation is possible
in the presence of saliva.
d. Secretes HCO3
- : neutralize acids in food as well as acids
produced by bacteria in the mouth, the neutral pH is good
for ptyalin action.
e. Lubrication: Mucus found in saliva facilitates moistening
and swallowing of food.
16
Control of Saliva Secretions
A. Parasympathetic fibers: causes copious secretion of saliva
(Cholinergic).
B. Sympathetic fibers: Causes small and insignificant secretion
which is viscous (Adrenergic).
Salivary reflex
• Sight, smell, and test or thinking of food  Receptors in oral
cavity or smell  Sensory fibers from the tongue to the
salivatory nuclei in MO  Parasympathetic fibers (act on
salivary glands)  increase salivary secretion.
Salivation can also be controlled by higher centers like
hypothalamus which has nerve connections with salivatory
nuclei in the Medulla oblongata (MO)
Higher centers like appetite area in the hypothalamus are also
involved in reflex control.
17
Phases of Salivary secretions
3-phases of salivary secretions include
1. Cephalic (brain) phase: triggered by thought, smell, or sight
of food
2. Oral phase: triggered by food that stimulate touch & test
receptors in the mouth
3. Gastric phase: triggered by substances which stimulate the
gastric mucosa (acids or sour tastes) in the stomach.
18
Swallowing reflex (Deglutition)
• Swallowing is an act of orderly propulsion of food from the
mouth to the stomach.
3-phases of swallowing reflex
1. Voluntary phase in the Oral cavity:
- the tongue rolls and pushes the chyme against the hard and
soft palate in the oral cavity and pushes it to the pharynx.
2. Pharyngeal phase (involuntary): Reflexes whose nuclei are
located in swallowing center of the MO, stimulate the closure
of the nasal and tracheal openings and cause inhibition of
respiration.
19
Steps:
a. The soft palate is pulled upward and closes the nasopharynx.
b. The vocal cords are pulled together and narrow
c. The epiglottis moves upward and covers the larynx
d. This so called “primary peristalsis” in the pharynx starts and
pushes the bolus downward through the upper esophageal
sphincter
20
3. Esophageal phase (involuntary):
• Upper esophageal sphincter closes and the bolus is pushed
down to the stomach.
• If the peristaltic contraction is not adequate to push the bolus,
then distention of the esophageal walls cause the so called
“secondary peristalsis” that is modified by enteric nerves and
strong enough to push the bolus to the stomach.
Secretion of the esophagus
Mucous glands that surround the esophagus secrete mucus that
provide lubrication and protection from gastric digestion (HCl)
that may arise from reflux of stomach contents.
21
Stomach
22
The stomach, functions
Storage (1-3 L)
Mixing (soupy chyme)
Secretion of intrinsic factor and other enzymes
Secretes enzymes that begin protein digestion
Has antiseptic actions (HCl) etc
23
Production of gastric juices in stomach
Cells Secretions
1. Parietal cells HCl + Intrinsic factor
2. Chief cells Pepsinogen (stimulated by HCl)
3. Mucous cells Mucous
4. G-cells Gastrin (at pyloric antrum)
5. D-cells Somatostatin
6. H-cells Histamine
24
• When pepsinogen is first secreted, it has no digestive activity
However, as soon as it comes in contact with HCl, it is
activated to form active pepsin
• Pepsin functions as an active proteolytic enzyme in a highly
acid medium (optimum pH 1.8 to 3.5), but above a pH of
about 5 it has almost no proteolytic activity and becomes
completely inactivated in a short time.
• HCl is as necessary as pepsin for protein digestion in the
stomach
25
• The substance intrinsic factor, essential for absorption of
vitamin B12 in the ileum, is secreted by the parietal cells
along with the secretion of HCl
• Enterochromaffin - like cells (ECL cells), secrete histamine.
• Gastrin is secreted by gastrin cells, also called G cells.
26
Phases of Gastric Secretion
• Gastric secretion is occur in three phases:
- a cephalic phase, a gastric phase, and an intestinal phase.
Cephalic Phase
• Occurs in feedforward fashion (before food reaches the
stomach), especially while it is being eaten.
• It results from the sight, smell, thought, taste of food,
chewing, or swallowing and the greater the appetite, the
more intense is the stimulation.
27
• This phase of secretion normally accounts for about 20% of the
gastric secretion associated with eating a meal.
28
Gastric Phase
29
The gastric phase of secretion accounts for about 70% of the
total gastric secretion associated with eating a meal
Begins when food reaches the stomach
Intestinal Phase
• The intestinal phase has two components that influence gastric
secretion:
• Excitatory
• Inhibitory
Excitatory component
• Occurs in response to the presence of products of protein
digestion in the duodenum
• Intestinal gastrin travels in the blood to the stomach, where it
enhances the secretion of HCl and pepsinogen
• It accounts only for approximately 10% of the acid secretory
response to a meal
30
Inhibitory component
• Has a very strong influence on gastric secretion
31
Pancreas
Major function of the pancreas
1. Involved in digestion processes by producing:
a. Digestive enzymes: necessary to digest
CHO, fat, and protein
b. Bicarbonates : to neutralize the gastric juice
c. Water and electrolytes (Na+, K+ etc):
Location of secretion
1. Digestive enzymes and electrolytes are secreted by acinar
cells of the pancreases
2. HCO3
- and water are secreted by the duct cells lining the
pancreatic ducts.
32
33
• The inactive proenzymes secreted by cells of the pancreas that
break protein, fat, and CHO include:
• Trypsinogen (protein digestion)
• Chymotrypsinogen (protein digestion)
• Procarboxypolypeptidase (protein digestion)
• Pancreatic lipase (fat digestion)
• Alpha amylase (CHO digestion)
34
• All the inactive enzymes (to prevent from digesting the
proteins of the cells in which they are formed) flow through the
pancreatic duct into the duodenum.
• At the beginning, an enzyme called enterokinase
(=enteropeptidase) that is located on the wall of the
duodenum changes trypsinogen to trypsin.
• Trypsin then activates the others as follows (look steps b and
c):
a. Trypsinogen ----enterokinase  Trypsin
b. Chymotrypsinogen----trypsin  Chymotrypsin
c. Procarboxypolypaptidase ---trypsin Carboxypolypeptidase
35
• The most important of the pancreatic enzymes for digesting
proteins are trypsin, chymotrypsin, and carboxypolypeptidase.
• By far the most abundant of these is trypsin.
• Trypsin and chymotrypsin split whole and partially digested
proteins into peptides of various sizes but do not cause release
of individual amino acids.
• However, carboxypolypeptidase does split some peptides into
individual amino acids
36
• The pancreatic enzyme for digesting carbohydrates is
pancreatic amylase, which hydrolyzes starches, glycogen, and
most other carbohydrates to form mostly disaccharide maltose
• The main enzyme for fat digestion is:
Pancreatic lipase hydrolyzes dietary triglycerides into
monoglycerides and free fatty acids, which are the absorbable
units of fat
37
38
Hormonal control of pancreatic exocrine secretion
Phases of Pancreatic Secretion
• Pancreatic secretion occurs in three phases, the same as for
gastric secretion: the cephalic phase, the gastric phase, and the
intestinal phase
Cephalic and Gastric Phases
• The same nervous signals from the brain that cause secretion in
the stomach also cause acetylcholine release by the vagus
nerve endings in the pancreas
• This causes moderate amounts of enzymes to be secreted in the
pancreatic acini
39
• Cephalic phase: approximately 20% of pancreatic secretory
response to a meal
• Gastric phase: 5 to 10% of pancreatic secretory response to a
meal
Intestinal phase
• Most pancreatic secretion takes place
• Secretin(intestinal hormone) stimulates release of a large
volume of pancreatic juice with a high [HCO3-]
• Secretin is released in response to acidic chyme in the
duodenum (maximal release at pH ~ 3.0)
40
• Cholecystokinin(intestinal hormone) stimulates the release of
digestive enzymes from the pancreas
• It is released in response to the presence of the products of
protein and lipid digestion
41
Secretion of Bile by the Liver
• One of the many functions of the liver is to secrete bile,
normally between 600 and 1000 ml/day.
• Bile serves two important functions:
1. Emulsification of fat in the duodenum
=changing greater fat globules into smaller fat-droplets called
micelles
2. Means for excretion of several important waste products from
the blood.
These include especially bilirubin, an end product of
hemoglobin destruction, and excesses of cholesterol
42
• After bile is produced in the liver it will be stored in the gall
bladder
• It is transported toward the small intestine by the hepatic duct
(from the liver) and the cystic duct (from the gallbladder),
which join to form the common bile duct
• The common bile duct and the pancreatic duct join to form the
hepatopancreatic ampulla, which empties into the duodenum
• The entrance to the duodenum is surrounded by the Sphincter
of Oddi
• It is closed between meals and relaxes in response to the
intestinal hormone cholecystokinin
43
44
45
Constituents of bile
• Bile contains the following constituents:
1. Bile salts (bile acids), ~11%
2. Bile pigment (bilirubin), ~1%
3. Others organic constituents like: ( ~3%)
Cholesterol, Lecithin, protein etc
4. Electrolytes (Na+, K+, Ca2+, Cl-, and greater HC03-
than plasma) ~1%.
5. H2O (~ 84%) also takes the higher share of bile
46
Bile pigments (Bilirubin):
• Does not play a role in digestion at all but instead is a waste
product excreted in the bile
• Is the primary bile pigment derived from the breakdown of
worn-out red blood cells
• The end product from degradation of the heme (iron
containing) part of the hemoglobin
• In the intestine, bacterial actions change bilirubin into
urobilinogen. They are further oxidized and excreted in feces
or urine as stercobilin and urobilin.
• Gives faces its brown color
• Excess level of bilirubin in the blood (>18 mg/L) causes
jaundice, that is, yellow coloration of the sclera's, skin etc
occurs.
47
• Gallbladder (GB) is a storage sac for bile.
• GB is stimulated by CCK to contract and release bile.
• Bile emulsifies fat in the duodenum.
• Cholesterol and other substances precipitate in the GB and this
effect favors gall-stone formation.
• 75% of gallstones are derived from cholesterol, other 25%
from bilirubin abnormal precipitation
48
The Small intestine (SI)
• SI is specialized for completion of digestion and absorption
of nutrients.
a. Duodenum: ~25 cm
It receives chyme from the stomach and digestive secretions
from the pancreas and liver and neutralize its acids
b. Jejunum : ~1.5m
Bulk of chemical digestion and nutrient absorption occurs
c. Ileum : ~1.7m
Mainly absorptive
Ileocecal valve: prevent backflow of fecal contents from the
colon into the SI
Ileocecal sphincter: slows emptying of ileal contents into the
cecum 49
50
• The mucosa of SI is well adapted for digestion and absorption
with certain anatomical modifications:
• Plicae circulares (circular folds)
• Villi
• Microvilli
Plicae circulares (circular folds)
• Form internal rings around the circumference of the SI
• Are formed from inward foldings of the mucosal and
submucosal layers of the intestinal wall
• Well developed in the duodenum and jejunum
• Increase the absorptive surface area of the mucosa about
threefold
51
52
Villi
• Millions of smaller projections of mucosa that cover
plica
• Two types of epithelial cells cover the villi:
–Goblet cells (produce mucus)
–Absorptive cells (most abundant)
• Taken together, the villi increase the absorptive surface
area another 10-fold
53
54
Microvilli
• Microscopic projections found on the luminal surface of the
absorptive cells - form the brush border ( contains
enterokinase, disaccharidases (maltase, sucrase,and lactase),
aminopeptidases)
• Increase the surface area for absorption another 20-fold
• Together, these three anatomical adaptations of the intestinal
mucosa — plicae circulares, villi, and microvilli — increase
the surface area as much as 600-fold
55
Motility of the small intestine
• Segmentation and peristalsis take place in the small intestine
• Segmentation: ring-like contractions along the length of the
SI
• Mixes chyme with digestive juices and exposes it to the
intestinal mucosa for absorption
• Causes only a small degree of forward movement of the
chyme along the small intestine
56
• Peristalsis:
• The wave-like form of muscle contraction
• Primarily moves chyme along the intestine and causes only
a small amount of mixing
• Weak and slow in the small intestine so that time is
sufficient for complete digestion and absorption of the
chyme
57
• The motility of the SI may be enhanced during a meal by:
• Distension of the small intestine
–Distension of the duodenum elicit segmentation
contractions in this segment
• Gastrin - causes segmentation of the empty ileum
• Extrinsic nerve stimulation
–Parasympathetic stimulation, by way of the vagus
nerve, further enhances segmentation.
–Sympathetic stimulation inhibits this activity
58
Digestion and absorption in the small intestine
Carbohydrates
• Starch is initially acted upon by amylase
• Salivary amylase breaks down starch molecules in the mouth
and stomach.
• Pancreatic amylase carries on this activity in the small intestine
• Amylase fragments polysaccharides into disaccharides (maltose,
composed of two glucose molecules)
• The disaccharide molecules, primarily maltose, are presented to
the brush border of the absorptive cells.
• As the disaccharides (maltose, sucrose & lactose) are absorbed,
disaccharidases (maltase, sucrase, and lactase) split these
nutrient molecules into monosaccharides (glucose, fructose, and
galactose)
59
• Glucose and galactose enter the absorptive cells by way of
secondary active transport
• Fructose enters the absorptive cells by way of facilitated
diffusion.
• All monosaccharide molecules exit the absorptive cells by
way of facilitated diffusion and enter the blood capillaries
60
61
62
Proteins
• Protein digestion begins in the stomach by the action of the
gastric enzyme pepsin.
–This enzyme fragments large protein molecules into smaller
peptide chains.
• Digestion is continued in the small intestine by the pancreatic
enzymes trypsin, chymotrypsin, and carboxypeptidase,
which hydrolyze the peptide chains into amino acids,
dipeptides, and tripeptides
• Amino acids enter the absorptive cells by way of secondary
active transport
63
• As the nutrient molecules are absorbed, aminopeptidases split
dipeptides and tripeptides into their constituent amino acids
• The amino acid molecules then exit the absorptive cells by way
of facilitated diffusion and enter the blood capillaries
64
65
(a) Protein digestion
66
(b) Protein absorption
Lipids
• Fat digestion begins in the mouth and stomach by the action of
the salivary enzyme lingual lipase
• However, the role of this enzyme is minor
• Lipids are digested primarily in the small intestine
1. Bile salts cause emulsification, which is the dispersal of large
fat droplets into a suspension of smaller droplets
2. Pancreatic lipase acts on the lipid droplets to hydrolyze the
triglyceride molecules into monoglycerides and free fatty
acids- these are water insoluble
3. Micelles formed by the amphipathic bile salts
• Monoglycerides and free fatty acids are carried in this interior
region of the micelle
67
• Upon reaching the brush border of the absorptive cells, they
leave the micelles and enter the cells by simple diffusion
–This process takes place primarily in the jejunum and
proximal ileum
• The bile salts are absorbed in the distal ileum by way of
secondary active transport
• Within the absorptive cells:
• Monoglycerides + FFAS triglycerides (in endoplasmic
reticulum)
• Triglycerides packaged in a lipoprotein coat (in Golgi
apparatus)
• These protein-coated lipid globules, referred to as
chylomicrons (water soluble)
68
• Chylomicrons leave the absorptive cell by way of exocytosis
• They enter the lacteals, which are part of the lymphatic system
B/c they unable to cross the basement membrane of the
blood capillaries
69
70
(a) Fat digestion
71
Water and electrolytes
• The absorption of nutrient molecules primarily takes place in
the duodenum and jejunum, creates an osmotic gradient for
the passive absorption of water
• Sodium absorption:
• Passive diffusion (through “leaky” tight junctions)
• Na+– Cl- cotransport
• Na+–glucose cotransport
• Na+–amino acid cotransport
• Na+- K+ - 2 Cl–cotransport
72
Large intestine
• From the ileocecal valve (juncture between the ileum
• and the large intestine) to the anus
• Has a larger diameter than the small intestine
• Mucosa:
• Composed of absorptive cells and mucus-secreting goblet cells
• Does not form villi
• Consists of the following structures:
- Cecum - Colon
- Appendix - Rectum
73
• Cecum:
• The most proximal portion of the large intestine
• Receives chyme from the ileum of the SI through the
ileocecal valve
• Appendix:
• a small projection at the bottom of the cecum
• is a lymphoid tissue. This tissue contains lymphocytes and
assists in defense against bacteria that enter the body through
the digestive system
74
• Colon:
• Largest portion of the large intestine
• Consists of four regions:
• Ascending colon (travels upward toward the diaphragm on
the right side of the abdomen)
• Transverse colon (crosses the abdomen under the diaphragm)
• Descending colon (travels downward through the abdomen
on the left side)
• Sigmoid colon (S-shaped region found in the lower abdomen)
75
76
• Rectum:
• Last portion of the digestive tract
• Leads to the external surface of the body through the anus
• Internal and external anal sphincters
• The large intestine typically receives 500 to 1500 ml of chyme
per day from the small intestine
• Chyme consists of indigestible food residues (e.g., cellulose),
unabsorbed biliary components, and any remaining fluid (b/c
most digestion and absorption in SI).
• Therefore, the two major functions of the large intestine are:
• Drying
• Storage
77
• The colon extracts more H2O and salt from the contents. What
remains to be eliminated is known as feces
• The primary function of the large intestine is to store feces
before defecation
Motility of the large intestine
• Normally sluggish
• Two types
–Mixing movements (Haustrations)
• Ring-like contractions (about 2.5 cm) of the circular muscle
divide the colon into pockets called haustra
• Haustrations = bulging of the large intestine into baglike
sacs as a result of circular and longitudinal muscle
contraction
• Serve primarily to move the contents slowly back and
forth, exposing them to the absorptive surface
78
–Propulsive movements (Mass movements)
• Propel chyme from the cecum to the sigmoid colon
• When a mass of feces is forced into the rectum, there is a
desire to defecate
Reflexes Affecting Mass Movements
• Gastrocolic reflex – stimulatory
–Distention of the stomach
• Duodenocolic reflex - stimulatory
–Distention of the small intestine
• Both push the colonic contents into the rectum, triggering the
defecation reflex
79
• Both reflexes transmitted by autonomic nervous system
• Defecation
• When mass movement forces feces into the rectrum
–Immediate desire to defecate
• Reflex contraction of rectrum
• Relaxation of anal sphincter
• Approx. 80 to 200 mL of fecal matter expelled daily
80
81
Defecation Reflex
1. Distension of the rectum.
2. Stimulation of the stretch receptors in the rectum.
3. A. Short reflex: Stimulation of myenteric plexus in sigmoid
colon and rectum.
B. Long reflex: stimulation of parasympathetic motor neurons
in sacral spinal cord.
C. stimulation of somatic motor neurons.
4. Increased local peristalsis, relaxation of internal anal
sphincter and contraction of external anal sphincter.
82
83
• From the spinal cord, defecation signals also have the
following effects
• Taking deep breath
• Closure of glottis
• Contraction of abdominal wall muscles
• Relaxation and movement of pelvis floor downward
84
Secretion of the large intestine
• Large intestine does not secrete any digestive enzymes
• Colonic secretion consists of an alkaline (NaHCO3) mucus
solution
• The mucus provides lubrication to facilitate passage of the
feces
• The NaHCO3 neutralizes irritating acids produced by local
bacterial fermentation
• No digestion takes place within the large intestine because there
are no digestive enzymes. However, the colonic bacteria do
digest some of the cellulose for their use
85
Composition of Feces
• Three-fourths water
• One-fourth solid matter
–30% dead bacteria
–10-20% fat
–10-20% inorganic matter
–2-3% protein
–30% undigested roughage (e.g. bile pigment, sloughed
epithelial cells)
86
Reading assignment
Function of Gastrointestinal Hormones
• Gastrin
• Secretin
• Cholecystokinin (CCK)
• Gastric inhibitory peptide(GIP)
87

A Chapter 10- GIT-1.pptx

  • 1.
  • 2.
    Digestive System • Thedigestive system consisting of the digestive tract (GIT) and accessory organs • The alimentary canal or gastrointestinal (GI) tract is composed of the organs mouth, pharynx, esophagus, stomach, small intestine, large intestine and anus • Accessory digestive organs – teeth, tongue and glandular organs, such as the salivary glands, liver, and pancreas 2
  • 3.
    The Components ofthe Digestive System 3
  • 4.
    Functions of gastrointestinalTract • Ingestion: taking materials into the digestive tract • Propulsion: swallowing and peristalsis –Peristalsis – waves of contraction and relaxation of muscles in the organ walls • Mechanical digestion: chewing, mixing, and churning food • Chemical digestion: catabolic breakdown of food with the help of enzymes • Absorption: movement of nutrients from the GI tract lumen into the blood or lymph • Secretion: release of water, acids, enzymes, buffers, and salts • Defecation: elimination of indigestible solid wastes 4
  • 5.
    Layers of theGIT 1. Mucosa: Inner layer of the luminal surfaces - protection, secretion & absorption 2. Submucosa: blood & lymph vessels, nerve plexuses - necessary for motility + secretion 3. Muscularis externa : contains circular & Longitudinal smooth muscle, mix and propel the chyme Circular : Contraction causes a decrease in diameter of the lumen of the GI tract Longitudinal : Contraction causes shortening of a segment of the GI tract 4. Serosa: outer most protective layer, consists of CT • Secretes a slippery fluid that lubricates and prevents friction 5
  • 6.
  • 7.
    Receptors of theGIT • Mechano- ,osmo- and chemo receptors respond to stretch and pH, presence of substrate, and end products of digestion respectively • They initiate reflexes that: –Activate or inhibit digestive glands –Mix lumen contents and move them along 7
  • 8.
    Innervation of theGI tract 1. Extrinsic NS • originate outside the digestive tract and innervate the various digestive organs = parasympathetic and sympathetic nervous systems a. Parasympathetic nervous system • Excitatory on the functions of the GI tract • Promotes digestive & absorptive processes 8
  • 9.
    b. Sympathetic nervoussystem • Inhibitory on the functions of the GI tract • Slows processes 2. Intrinsic NS • Lie entirely within the digestive tract wall =enteric nervous system • Coordinates and relays information from the parasympathetic and sympathetic nervous systems to the GI tract • Controls motility and secretion • Shows local, intrinsic regulation 9
  • 10.
    a. Myenteric plexus(Auerbach’s plexus) • Primarily controls the motility of the GI smooth muscle b. Submucosal plexus (Meissner’s plexus) • Primarily controls secretion • Types of Neurotransmitters: Several e.g., Ach, NE, E, NO vasoactive intestinal peptide (VIP), etc. 10
  • 11.
    GI Motility A. Peristaltic(propulsive) movements: • Consists of a progressive wave of strong contraction followed by relaxation • The main stimulus to cause peristalsis is distension (stretch) of the gut wall. Propel or push the contents forward through the digestive tract • Transit of food through the esophagus is rapid • In the small intestine—the main site of digestion and absorption—the contents are moved forward slowly allowing time for the breakdown and absorption of food 11
  • 12.
    a. Mixing movements: •Occurs due to local contractions taking place in SI segments Mixing food with the digestive juices (promote digestion of the food) Facilitate absorption by exposing all parts of the intestinal contents to the absorbing surfaces of the digestive tract The law of the gut movement always occurs from mouth- to- anus unless pathological 12
  • 13.
    Digestion • Two parts: 1.Mechanical digestion: • Is the physical grinding of food to smaller units without altering their chemical composition. • Helps to expose and increase the surface area for enzymatic attacks 2. Chemical digestion: • Is hydrolytic breakdown of nutrients (CHO, fat, protein etc) by different chemical reactions into simpler forms. • This processes changes the original composition of the nutrient. 13
  • 14.
    The Mouth (oralcavity) • The entrance to the digestive tract • Parts of the oral cavity: cheeks, lip, tongue, hard and soft palate, teeth • Is responsible for mechanical digestion of solid food by mastication Functions of chewing are: • To grind and break food up into smaller pieces –Facilitate swallowing –Increase the food surface area on which salivary enzymes will act • To mix food with saliva • To stimulate the taste buds 14
  • 15.
    Saliva • Three majorpairs of salivary glands a. Parotid glands (25%) Secrete mainly serous watery fluid rich in ptyalin b. Submandibular glands (70%) Produce both serous and mucous fluid c. Sublingual glands (~5%) Secrete mainly thick mucous with little serous fluid • Components: –H2O (99.5%) –Electrolytes(principally Na+, Cl-, and HCO3 -), antibodies (IgA) , lysozyme, amylase, thiocyanate, mucus..(0.5%) –Total secretion = about 1-1.5 L/day 15
  • 16.
    Saliva functions a. Digestion:CHO-digestion begins in saliva . The enzyme ptyalin breaks starch- to-maltose. Lingual lipase begins fat digestion in the mouth. b. Protection: has anti-microbial actions (contains IgA, lysozyme & thiocyanate that kills microbes). c. Involved in speech: Clear & fluent articulation is possible in the presence of saliva. d. Secretes HCO3 - : neutralize acids in food as well as acids produced by bacteria in the mouth, the neutral pH is good for ptyalin action. e. Lubrication: Mucus found in saliva facilitates moistening and swallowing of food. 16
  • 17.
    Control of SalivaSecretions A. Parasympathetic fibers: causes copious secretion of saliva (Cholinergic). B. Sympathetic fibers: Causes small and insignificant secretion which is viscous (Adrenergic). Salivary reflex • Sight, smell, and test or thinking of food  Receptors in oral cavity or smell  Sensory fibers from the tongue to the salivatory nuclei in MO  Parasympathetic fibers (act on salivary glands)  increase salivary secretion. Salivation can also be controlled by higher centers like hypothalamus which has nerve connections with salivatory nuclei in the Medulla oblongata (MO) Higher centers like appetite area in the hypothalamus are also involved in reflex control. 17
  • 18.
    Phases of Salivarysecretions 3-phases of salivary secretions include 1. Cephalic (brain) phase: triggered by thought, smell, or sight of food 2. Oral phase: triggered by food that stimulate touch & test receptors in the mouth 3. Gastric phase: triggered by substances which stimulate the gastric mucosa (acids or sour tastes) in the stomach. 18
  • 19.
    Swallowing reflex (Deglutition) •Swallowing is an act of orderly propulsion of food from the mouth to the stomach. 3-phases of swallowing reflex 1. Voluntary phase in the Oral cavity: - the tongue rolls and pushes the chyme against the hard and soft palate in the oral cavity and pushes it to the pharynx. 2. Pharyngeal phase (involuntary): Reflexes whose nuclei are located in swallowing center of the MO, stimulate the closure of the nasal and tracheal openings and cause inhibition of respiration. 19
  • 20.
    Steps: a. The softpalate is pulled upward and closes the nasopharynx. b. The vocal cords are pulled together and narrow c. The epiglottis moves upward and covers the larynx d. This so called “primary peristalsis” in the pharynx starts and pushes the bolus downward through the upper esophageal sphincter 20
  • 21.
    3. Esophageal phase(involuntary): • Upper esophageal sphincter closes and the bolus is pushed down to the stomach. • If the peristaltic contraction is not adequate to push the bolus, then distention of the esophageal walls cause the so called “secondary peristalsis” that is modified by enteric nerves and strong enough to push the bolus to the stomach. Secretion of the esophagus Mucous glands that surround the esophagus secrete mucus that provide lubrication and protection from gastric digestion (HCl) that may arise from reflux of stomach contents. 21
  • 22.
  • 23.
    The stomach, functions Storage(1-3 L) Mixing (soupy chyme) Secretion of intrinsic factor and other enzymes Secretes enzymes that begin protein digestion Has antiseptic actions (HCl) etc 23
  • 24.
    Production of gastricjuices in stomach Cells Secretions 1. Parietal cells HCl + Intrinsic factor 2. Chief cells Pepsinogen (stimulated by HCl) 3. Mucous cells Mucous 4. G-cells Gastrin (at pyloric antrum) 5. D-cells Somatostatin 6. H-cells Histamine 24
  • 25.
    • When pepsinogenis first secreted, it has no digestive activity However, as soon as it comes in contact with HCl, it is activated to form active pepsin • Pepsin functions as an active proteolytic enzyme in a highly acid medium (optimum pH 1.8 to 3.5), but above a pH of about 5 it has almost no proteolytic activity and becomes completely inactivated in a short time. • HCl is as necessary as pepsin for protein digestion in the stomach 25
  • 26.
    • The substanceintrinsic factor, essential for absorption of vitamin B12 in the ileum, is secreted by the parietal cells along with the secretion of HCl • Enterochromaffin - like cells (ECL cells), secrete histamine. • Gastrin is secreted by gastrin cells, also called G cells. 26
  • 27.
    Phases of GastricSecretion • Gastric secretion is occur in three phases: - a cephalic phase, a gastric phase, and an intestinal phase. Cephalic Phase • Occurs in feedforward fashion (before food reaches the stomach), especially while it is being eaten. • It results from the sight, smell, thought, taste of food, chewing, or swallowing and the greater the appetite, the more intense is the stimulation. 27
  • 28.
    • This phaseof secretion normally accounts for about 20% of the gastric secretion associated with eating a meal. 28
  • 29.
    Gastric Phase 29 The gastricphase of secretion accounts for about 70% of the total gastric secretion associated with eating a meal Begins when food reaches the stomach
  • 30.
    Intestinal Phase • Theintestinal phase has two components that influence gastric secretion: • Excitatory • Inhibitory Excitatory component • Occurs in response to the presence of products of protein digestion in the duodenum • Intestinal gastrin travels in the blood to the stomach, where it enhances the secretion of HCl and pepsinogen • It accounts only for approximately 10% of the acid secretory response to a meal 30
  • 31.
    Inhibitory component • Hasa very strong influence on gastric secretion 31
  • 32.
    Pancreas Major function ofthe pancreas 1. Involved in digestion processes by producing: a. Digestive enzymes: necessary to digest CHO, fat, and protein b. Bicarbonates : to neutralize the gastric juice c. Water and electrolytes (Na+, K+ etc): Location of secretion 1. Digestive enzymes and electrolytes are secreted by acinar cells of the pancreases 2. HCO3 - and water are secreted by the duct cells lining the pancreatic ducts. 32
  • 33.
  • 34.
    • The inactiveproenzymes secreted by cells of the pancreas that break protein, fat, and CHO include: • Trypsinogen (protein digestion) • Chymotrypsinogen (protein digestion) • Procarboxypolypeptidase (protein digestion) • Pancreatic lipase (fat digestion) • Alpha amylase (CHO digestion) 34
  • 35.
    • All theinactive enzymes (to prevent from digesting the proteins of the cells in which they are formed) flow through the pancreatic duct into the duodenum. • At the beginning, an enzyme called enterokinase (=enteropeptidase) that is located on the wall of the duodenum changes trypsinogen to trypsin. • Trypsin then activates the others as follows (look steps b and c): a. Trypsinogen ----enterokinase  Trypsin b. Chymotrypsinogen----trypsin  Chymotrypsin c. Procarboxypolypaptidase ---trypsin Carboxypolypeptidase 35
  • 36.
    • The mostimportant of the pancreatic enzymes for digesting proteins are trypsin, chymotrypsin, and carboxypolypeptidase. • By far the most abundant of these is trypsin. • Trypsin and chymotrypsin split whole and partially digested proteins into peptides of various sizes but do not cause release of individual amino acids. • However, carboxypolypeptidase does split some peptides into individual amino acids 36
  • 37.
    • The pancreaticenzyme for digesting carbohydrates is pancreatic amylase, which hydrolyzes starches, glycogen, and most other carbohydrates to form mostly disaccharide maltose • The main enzyme for fat digestion is: Pancreatic lipase hydrolyzes dietary triglycerides into monoglycerides and free fatty acids, which are the absorbable units of fat 37
  • 38.
    38 Hormonal control ofpancreatic exocrine secretion
  • 39.
    Phases of PancreaticSecretion • Pancreatic secretion occurs in three phases, the same as for gastric secretion: the cephalic phase, the gastric phase, and the intestinal phase Cephalic and Gastric Phases • The same nervous signals from the brain that cause secretion in the stomach also cause acetylcholine release by the vagus nerve endings in the pancreas • This causes moderate amounts of enzymes to be secreted in the pancreatic acini 39
  • 40.
    • Cephalic phase:approximately 20% of pancreatic secretory response to a meal • Gastric phase: 5 to 10% of pancreatic secretory response to a meal Intestinal phase • Most pancreatic secretion takes place • Secretin(intestinal hormone) stimulates release of a large volume of pancreatic juice with a high [HCO3-] • Secretin is released in response to acidic chyme in the duodenum (maximal release at pH ~ 3.0) 40
  • 41.
    • Cholecystokinin(intestinal hormone)stimulates the release of digestive enzymes from the pancreas • It is released in response to the presence of the products of protein and lipid digestion 41
  • 42.
    Secretion of Bileby the Liver • One of the many functions of the liver is to secrete bile, normally between 600 and 1000 ml/day. • Bile serves two important functions: 1. Emulsification of fat in the duodenum =changing greater fat globules into smaller fat-droplets called micelles 2. Means for excretion of several important waste products from the blood. These include especially bilirubin, an end product of hemoglobin destruction, and excesses of cholesterol 42
  • 43.
    • After bileis produced in the liver it will be stored in the gall bladder • It is transported toward the small intestine by the hepatic duct (from the liver) and the cystic duct (from the gallbladder), which join to form the common bile duct • The common bile duct and the pancreatic duct join to form the hepatopancreatic ampulla, which empties into the duodenum • The entrance to the duodenum is surrounded by the Sphincter of Oddi • It is closed between meals and relaxes in response to the intestinal hormone cholecystokinin 43
  • 44.
  • 45.
  • 46.
    Constituents of bile •Bile contains the following constituents: 1. Bile salts (bile acids), ~11% 2. Bile pigment (bilirubin), ~1% 3. Others organic constituents like: ( ~3%) Cholesterol, Lecithin, protein etc 4. Electrolytes (Na+, K+, Ca2+, Cl-, and greater HC03- than plasma) ~1%. 5. H2O (~ 84%) also takes the higher share of bile 46
  • 47.
    Bile pigments (Bilirubin): •Does not play a role in digestion at all but instead is a waste product excreted in the bile • Is the primary bile pigment derived from the breakdown of worn-out red blood cells • The end product from degradation of the heme (iron containing) part of the hemoglobin • In the intestine, bacterial actions change bilirubin into urobilinogen. They are further oxidized and excreted in feces or urine as stercobilin and urobilin. • Gives faces its brown color • Excess level of bilirubin in the blood (>18 mg/L) causes jaundice, that is, yellow coloration of the sclera's, skin etc occurs. 47
  • 48.
    • Gallbladder (GB)is a storage sac for bile. • GB is stimulated by CCK to contract and release bile. • Bile emulsifies fat in the duodenum. • Cholesterol and other substances precipitate in the GB and this effect favors gall-stone formation. • 75% of gallstones are derived from cholesterol, other 25% from bilirubin abnormal precipitation 48
  • 49.
    The Small intestine(SI) • SI is specialized for completion of digestion and absorption of nutrients. a. Duodenum: ~25 cm It receives chyme from the stomach and digestive secretions from the pancreas and liver and neutralize its acids b. Jejunum : ~1.5m Bulk of chemical digestion and nutrient absorption occurs c. Ileum : ~1.7m Mainly absorptive Ileocecal valve: prevent backflow of fecal contents from the colon into the SI Ileocecal sphincter: slows emptying of ileal contents into the cecum 49
  • 50.
  • 51.
    • The mucosaof SI is well adapted for digestion and absorption with certain anatomical modifications: • Plicae circulares (circular folds) • Villi • Microvilli Plicae circulares (circular folds) • Form internal rings around the circumference of the SI • Are formed from inward foldings of the mucosal and submucosal layers of the intestinal wall • Well developed in the duodenum and jejunum • Increase the absorptive surface area of the mucosa about threefold 51
  • 52.
  • 53.
    Villi • Millions ofsmaller projections of mucosa that cover plica • Two types of epithelial cells cover the villi: –Goblet cells (produce mucus) –Absorptive cells (most abundant) • Taken together, the villi increase the absorptive surface area another 10-fold 53
  • 54.
  • 55.
    Microvilli • Microscopic projectionsfound on the luminal surface of the absorptive cells - form the brush border ( contains enterokinase, disaccharidases (maltase, sucrase,and lactase), aminopeptidases) • Increase the surface area for absorption another 20-fold • Together, these three anatomical adaptations of the intestinal mucosa — plicae circulares, villi, and microvilli — increase the surface area as much as 600-fold 55
  • 56.
    Motility of thesmall intestine • Segmentation and peristalsis take place in the small intestine • Segmentation: ring-like contractions along the length of the SI • Mixes chyme with digestive juices and exposes it to the intestinal mucosa for absorption • Causes only a small degree of forward movement of the chyme along the small intestine 56
  • 57.
    • Peristalsis: • Thewave-like form of muscle contraction • Primarily moves chyme along the intestine and causes only a small amount of mixing • Weak and slow in the small intestine so that time is sufficient for complete digestion and absorption of the chyme 57
  • 58.
    • The motilityof the SI may be enhanced during a meal by: • Distension of the small intestine –Distension of the duodenum elicit segmentation contractions in this segment • Gastrin - causes segmentation of the empty ileum • Extrinsic nerve stimulation –Parasympathetic stimulation, by way of the vagus nerve, further enhances segmentation. –Sympathetic stimulation inhibits this activity 58
  • 59.
    Digestion and absorptionin the small intestine Carbohydrates • Starch is initially acted upon by amylase • Salivary amylase breaks down starch molecules in the mouth and stomach. • Pancreatic amylase carries on this activity in the small intestine • Amylase fragments polysaccharides into disaccharides (maltose, composed of two glucose molecules) • The disaccharide molecules, primarily maltose, are presented to the brush border of the absorptive cells. • As the disaccharides (maltose, sucrose & lactose) are absorbed, disaccharidases (maltase, sucrase, and lactase) split these nutrient molecules into monosaccharides (glucose, fructose, and galactose) 59
  • 60.
    • Glucose andgalactose enter the absorptive cells by way of secondary active transport • Fructose enters the absorptive cells by way of facilitated diffusion. • All monosaccharide molecules exit the absorptive cells by way of facilitated diffusion and enter the blood capillaries 60
  • 61.
  • 62.
  • 63.
    Proteins • Protein digestionbegins in the stomach by the action of the gastric enzyme pepsin. –This enzyme fragments large protein molecules into smaller peptide chains. • Digestion is continued in the small intestine by the pancreatic enzymes trypsin, chymotrypsin, and carboxypeptidase, which hydrolyze the peptide chains into amino acids, dipeptides, and tripeptides • Amino acids enter the absorptive cells by way of secondary active transport 63
  • 64.
    • As thenutrient molecules are absorbed, aminopeptidases split dipeptides and tripeptides into their constituent amino acids • The amino acid molecules then exit the absorptive cells by way of facilitated diffusion and enter the blood capillaries 64
  • 65.
  • 66.
  • 67.
    Lipids • Fat digestionbegins in the mouth and stomach by the action of the salivary enzyme lingual lipase • However, the role of this enzyme is minor • Lipids are digested primarily in the small intestine 1. Bile salts cause emulsification, which is the dispersal of large fat droplets into a suspension of smaller droplets 2. Pancreatic lipase acts on the lipid droplets to hydrolyze the triglyceride molecules into monoglycerides and free fatty acids- these are water insoluble 3. Micelles formed by the amphipathic bile salts • Monoglycerides and free fatty acids are carried in this interior region of the micelle 67
  • 68.
    • Upon reachingthe brush border of the absorptive cells, they leave the micelles and enter the cells by simple diffusion –This process takes place primarily in the jejunum and proximal ileum • The bile salts are absorbed in the distal ileum by way of secondary active transport • Within the absorptive cells: • Monoglycerides + FFAS triglycerides (in endoplasmic reticulum) • Triglycerides packaged in a lipoprotein coat (in Golgi apparatus) • These protein-coated lipid globules, referred to as chylomicrons (water soluble) 68
  • 69.
    • Chylomicrons leavethe absorptive cell by way of exocytosis • They enter the lacteals, which are part of the lymphatic system B/c they unable to cross the basement membrane of the blood capillaries 69
  • 70.
  • 71.
  • 72.
    Water and electrolytes •The absorption of nutrient molecules primarily takes place in the duodenum and jejunum, creates an osmotic gradient for the passive absorption of water • Sodium absorption: • Passive diffusion (through “leaky” tight junctions) • Na+– Cl- cotransport • Na+–glucose cotransport • Na+–amino acid cotransport • Na+- K+ - 2 Cl–cotransport 72
  • 73.
    Large intestine • Fromthe ileocecal valve (juncture between the ileum • and the large intestine) to the anus • Has a larger diameter than the small intestine • Mucosa: • Composed of absorptive cells and mucus-secreting goblet cells • Does not form villi • Consists of the following structures: - Cecum - Colon - Appendix - Rectum 73
  • 74.
    • Cecum: • Themost proximal portion of the large intestine • Receives chyme from the ileum of the SI through the ileocecal valve • Appendix: • a small projection at the bottom of the cecum • is a lymphoid tissue. This tissue contains lymphocytes and assists in defense against bacteria that enter the body through the digestive system 74
  • 75.
    • Colon: • Largestportion of the large intestine • Consists of four regions: • Ascending colon (travels upward toward the diaphragm on the right side of the abdomen) • Transverse colon (crosses the abdomen under the diaphragm) • Descending colon (travels downward through the abdomen on the left side) • Sigmoid colon (S-shaped region found in the lower abdomen) 75
  • 76.
  • 77.
    • Rectum: • Lastportion of the digestive tract • Leads to the external surface of the body through the anus • Internal and external anal sphincters • The large intestine typically receives 500 to 1500 ml of chyme per day from the small intestine • Chyme consists of indigestible food residues (e.g., cellulose), unabsorbed biliary components, and any remaining fluid (b/c most digestion and absorption in SI). • Therefore, the two major functions of the large intestine are: • Drying • Storage 77
  • 78.
    • The colonextracts more H2O and salt from the contents. What remains to be eliminated is known as feces • The primary function of the large intestine is to store feces before defecation Motility of the large intestine • Normally sluggish • Two types –Mixing movements (Haustrations) • Ring-like contractions (about 2.5 cm) of the circular muscle divide the colon into pockets called haustra • Haustrations = bulging of the large intestine into baglike sacs as a result of circular and longitudinal muscle contraction • Serve primarily to move the contents slowly back and forth, exposing them to the absorptive surface 78
  • 79.
    –Propulsive movements (Massmovements) • Propel chyme from the cecum to the sigmoid colon • When a mass of feces is forced into the rectum, there is a desire to defecate Reflexes Affecting Mass Movements • Gastrocolic reflex – stimulatory –Distention of the stomach • Duodenocolic reflex - stimulatory –Distention of the small intestine • Both push the colonic contents into the rectum, triggering the defecation reflex 79
  • 80.
    • Both reflexestransmitted by autonomic nervous system • Defecation • When mass movement forces feces into the rectrum –Immediate desire to defecate • Reflex contraction of rectrum • Relaxation of anal sphincter • Approx. 80 to 200 mL of fecal matter expelled daily 80
  • 81.
  • 82.
    Defecation Reflex 1. Distensionof the rectum. 2. Stimulation of the stretch receptors in the rectum. 3. A. Short reflex: Stimulation of myenteric plexus in sigmoid colon and rectum. B. Long reflex: stimulation of parasympathetic motor neurons in sacral spinal cord. C. stimulation of somatic motor neurons. 4. Increased local peristalsis, relaxation of internal anal sphincter and contraction of external anal sphincter. 82
  • 83.
  • 84.
    • From thespinal cord, defecation signals also have the following effects • Taking deep breath • Closure of glottis • Contraction of abdominal wall muscles • Relaxation and movement of pelvis floor downward 84
  • 85.
    Secretion of thelarge intestine • Large intestine does not secrete any digestive enzymes • Colonic secretion consists of an alkaline (NaHCO3) mucus solution • The mucus provides lubrication to facilitate passage of the feces • The NaHCO3 neutralizes irritating acids produced by local bacterial fermentation • No digestion takes place within the large intestine because there are no digestive enzymes. However, the colonic bacteria do digest some of the cellulose for their use 85
  • 86.
    Composition of Feces •Three-fourths water • One-fourth solid matter –30% dead bacteria –10-20% fat –10-20% inorganic matter –2-3% protein –30% undigested roughage (e.g. bile pigment, sloughed epithelial cells) 86
  • 87.
    Reading assignment Function ofGastrointestinal Hormones • Gastrin • Secretin • Cholecystokinin (CCK) • Gastric inhibitory peptide(GIP) 87