Announcements

Office hour/exam review immediately after

class in INGR 2001
Cheat sheets for final exam
8.5 inches X 11 inches
HANDWRITTEN
Content on one side only
(On the back, write your name and a brief
description of yourself)
 14.1 Introduction: Feeding
Strategies and Evolution
 Nutrient molecules needed for energy production and
as building blocks.
The digestive system contributes to homeostasis
by transferring nutrients, water and electrolytes
from the external environment to the internal
environment.
Most animals commit considerable time and
energy to acquiring and digesting food.
Carbohydrates, fats and proteins are the main
fuels.
Broken down into absorbable units (simple
sugars, fatty acids and amino acids)
14.1 Introduction
More complex animal
groups have a complete
digestive system, a tube
with an entrance (mouth)
and exit (anus).
The lumen is part of
the external
environment.
The gut tube is divided
into three regions:
foregut, midgut and
hindgut.
A one-way
digestive tract
allows
simultaneous
operation of
sequential stages
in the processing of
food and reduces
mixing of digested
and undigested
matter. The crop is
a storage region
found in some
animals.
14.2 General Aspects of Digestion
Four basic digestive processes:
1) Motility
Muscular contractions mix the
contents of the digestive tract and
move them forward.
Smooth muscle in the walls of the
digestive organs is regulated by
complex autonomic mechanisms.
Skeletal muscle at the two ends of
the digestive tract is under
voluntary control.
14.2 Motility cont’d
Autonomous smooth muscle function in the
digestive tract.
Slow-wave potentials (basic electrical rhythm,
BER) are fluctuations in membrane potential.
Sheets of smooth muscle cells are connected by
gap junctions, forming a functional syncytium.
If threshold is reached, a volley of action
potentials is triggered.
Opening of voltage-gated Ca2+ channels
results in Ca2+ influx into smooth muscle
cells.
Intensity/strength of contractions depends on
the number of action potentials produced.
Summary
of
pathways
controlling
digestive
system
activities.
The enteric
(intrinsic)
nervous
system of a
rat’s
stomach
14.2 Four basic digestive processes cont’d

2) Secretion
Digestive juices are secreted
into the lumen of the digestive
tract by exocrine glands.
Each secretion contains water,
electrolytes, and organic
molecules important in the
digestive process.
14.2 Four basic digestive processes cont’d

3) Digestion
Complex food molecules
are broken down into
smaller absorbable units
by enzymatic hydrolysis.
Structure and
hydrolysis of
common
dietary
carbohydrates
14.2 Four basic digestive processes cont’d

4) Absorption
Small absorbable units, along with water,
vitamins and electrolytes, are transferred
from the digestive tract into blood.
Involves specialized transporters in
epithelial cells
Surface area is greatly increased to
enhance absorption.
Some aquatic species can absorb small
nutrient molecules through the gills or
epidermis.
14.2 Digestive tract of vertebrates

The tube
Mouth
Pharynx
Esophagus
Stomach (and rumen in ruminants) or
proventriculus-gizzard complex in birds
Small Intestine
Large intestine
Anus
14.2 Digestive tract of vertebrates
continued
Accessory organs
Salivary glands
Exocrine pancreas
Biliary system (liver and
gallbladder)
Digestive
tract of a
frog
14.3 Mouth
 Obtaining and receiving food
Palate separates the mouth from the nasal
passages.
Hard palate (in front) is made of bone.
Soft palate (in rear)
Tongue is a voluntarily controlled skeletal muscle.
Used to capture food, guide food within the mouth, taste,
vocalize, and synthesize an antibiotic peptide.
Specialized mouth shapes
Snakes have hinged jaws and powerful muscles in the
cheek and throat for swallowing prey whole.
Beaks of birds have been molded to conform to particular
feeding habits.
Muscular lips in mammals aid in seizing and conveying
food to the mouth (prehension).
14.3 Mouth cont’d

Mastication (chewing)
Purposes
Grind and break food up into smaller pieces to
facilitate swallowing and increase surface area
for mixing with saliva
Mix food with saliva
Stimulate the taste buds
Mastication is regulated by a
chewing center in the medulla
oblongata.
14.3 Mouth cont’d

Mastication continued
Teeth
Exposed part is covered by enamel
(hardest structure in the body).
Teeth of rabbits and lower incisors of
rodents grow continually to replace lost
enamel coating.
Sharks and reptiles replace teeth
throughout life (polyphydontia).
Baleen replaces teeth in baleen whales
for filter feeding.
 14.3 Mouth cont’d
Saliva
Produced by salivary glands
Parotid, sublingual and submandibular glands
in mammals
Contains water, mucus and salivary
amylase
Continuously secreted in mammals
Ruminants produce copious amounts of
alkaline saliva (e.g. cattle produce 140
liters/day).
14.3 Mouth cont’d
Functions of saliva:
Moistening food -- lubrication is
provided by mucus.
Digestion through the action of
salivary amylase and lingual
lipase (most digestion occurs
farther down the digestive tract)
Defense against ingested
bacteria
14.3 Mouth cont’d
Functions of saliva:
Taste -- only molecules in solution reach
taste buds
Neutralization of acid
Thermoregulation -- evaporative cooling in
panting
Poisons (e.g. snake venom)
Anticoagulation in blood-sucking animals
Pheromones
14.3 Mouth cont’d
Salivary reflexes
Simple or unconditioned salivary reflex occurs
when chemoreceptors and pressure receptors in
the oral cavity respond to the presence of food.
Salivary center in medulla oblongata stimulates
autonomic neurons.
Acquired or conditioned salivary reflex
occurs without oral stimulation -- “mouth-
watering” anticipation.
Learned response based on previous
experience
 14.3 Mouth cont’d
Salivary reflexes continued
Autonomic control
Both parasympathetic and
sympathetic neurons stimulate
saliva production, but
parasympathetic dominates.
Sympathetic stimulation
produces more mucus -- dry
mouth sensation during stress.
Control of salivary secretion in a mammal
Quick EC activity

Write your name on a

piece of paper.
Write a question or topic
that you wish had been
covered on Wednesday’s
exam that wasn’t.
14.3 Mouth cont’d

Mastication (chewing)
Purposes
Grind and break food up into smaller pieces to
facilitate swallowing and increase surface area
for mixing with saliva
Mix food with saliva
Stimulate the taste buds
Mastication is regulated by a
chewing center in the medulla
oblongata.
 14.3 Mouth cont’d
Saliva
Produced by salivary glands
Parotid, sublingual and submandibular glands
in mammals
Contains water, mucus and salivary
amylase
Continuously secreted in mammals
Ruminants produce copious amounts of
alkaline saliva (e.g. cattle produce 140
liters/day).
14.3 Mouth cont’d
Functions of saliva:
Moistening food -- lubrication is
provided by mucus.
Digestion through the action of
salivary amylase and lingual
lipase (most digestion occurs
farther down the digestive tract)
Defense against ingested
bacteria
14.3 Mouth cont’d
Functions of saliva:
Taste -- only molecules in solution reach
taste buds
Neutralization of acid
Thermoregulation -- evaporative cooling in
panting
Poisons (e.g. snake venom)
Anticoagulation in blood-sucking animals
Pheromones
14.3 Mouth cont’d
Salivary reflexes
 Simple or unconditioned salivary reflex occurs
when chemoreceptors and pressure receptors in
the oral cavity respond to the presence of food.
 Salivary center in medulla oblongata stimulates
autonomic neurons.
 Acquired or conditioned salivary reflex
occurs without oral stimulation -- “mouth-
watering” anticipation.
 Learned response based on previous
experience
 14.3 Mouth cont’d
Salivary reflexes continued
Autonomic control
Both parasympathetic and
sympathetic neurons
stimulate saliva production,
but parasympathetic
dominates.
Sympathetic stimulation
produces more mucus -- dry
mouth sensation during
stress.
Control of salivary secretion in a mammal
Quick EC activity

Write your name on a

piece of paper.
Write a question or topic
that you wish had been
covered on Wednesday’s
exam that wasn’t.
14.4 Pharynx, Esophagus, and Crop
Pharynx
Cavity at the rear of the oral cavity
Common passageway for both the
digestive system and respiratory system in
vertebrates
Esophagus
Fairly straight muscular tube extending
between pharynx and stomach
Skeletal muscle gives way to smooth
muscle in most vertebrates.
14.4 Cont’d
Crop
Sac-like outpouching of
the esophagus in some
species of birds
Temporary storage of food
Crop milk, high in lipids, is
secreted into the crop of
some birds, regurgitated,
and fed to young.
14.4 Cont’d
 Swallowing is the process of moving food
from the mouth through the esophagus into
the stomach.
 Bolus of food is forced by the tongue into the
pharynx.
 Pharyngeal pressure receptors send afferent
impulses to the medulla oblongata.
 Swallowing center activates a programmed
all-or-none sequence of highly coordinated
activities.
 Swallowing reflex has two phases:
oropharyngeal phase and esophageal
phase.
Oropharyngeal stage of swallowing in humans
 (Whole Milk Dud)

Peristalsis in the esophagus

 14.5 Stomach or Midgut

Functions of the stomach:

Storage of food
Digestion of proteins
(initiation)
Formation of chyme
Chyme is a thick, liquid
mixture of food and
gastric secretions.
Anatomy
of the
stomach
(human)
14.5 Cont’d
Gastric motility
 Filling
 Deep folds in stomach wall flatten as stomach
relaxes with ingestion (receptive relaxation).
 Storage
 Weak contractions in fundus and body allow
storage of food in the body and a pocket of gas
in the fundus.
 Mixing
 Strong peristaltic contractions mix food with
gastric secretions in the antrum.
 With pyloric sphincter closed, chyme tumbles
back for more mixing.
 14.5 Cont’d

Gastric motility continued

Emptying
Small amounts of chyme pass into the
duodenum with each contraction before
the pyloric sphincter closes.
Stomach empties at a rate proportional
to the volume of chyme in it.
Duodenal factors affecting gastric
emptying are fat content, acidity,
hypertonicity and distension.
Gastric emptying and mixing as a result of antral peristaltic contractions in a human stomach.
 14.5 Cont’d
 Gastric secretion
 Cells responsible for gastric secretion are
located on the inner surface of gastric pits
in the stomach lining (gastric mucosa).
 Oxyntic mucosa in the body and fundus
 Mucous cells secrete a thin, watery mucus.
 Chief cells secrete pepsinogen.
 Pepsinogen is inactive until cleaved by HCl or
pepsin to form pepsin.
 Pepsin digests proteins in a low pH environment.
 Parietal cells (a.k.a. oxyntic cells) secrete HCl and
intrinsic factor.
 Pyloric gland area in antrum
 Endocrine G cells secrete gastrin into the blood.
Mechanism
of HCl
secretion
Pepsinogen
activation in the
stomach lumen
(vertebrate)
14.5 Cont’d
 Control of gastric secretion
 Cephalic phase
 Anticipatory secretion of HCl and pepsinogen due
to food-related stimuli
 Stimulated by acetylcholine and gastrin
 Gastric phase
 Stimuli acting in the stomach -- protein (most potent
stimulus), distension, caffeine and alcohol
 Intestinal phase
 Excitatory component -- presence of protein
fragments in duodenum
 Inhibitory component -- same factors that inhibit
gastric motility also inhibit gastric secretion
14.5 Cont’d
 Digestion in the stomach
 Carbohydrate digestion continues due to
swallowed salivary amylase.
 Amylase is destroyed by stomach acid, but
interior of swallowed food masses is protected.
 Protein digestion begins in the antrum with
the action of HCl and pepsin.
 Proteins are hydrolyzed into smaller peptides.
 Stomach lining is protected from digestion
by the gastric mucosal barrier.
 Secretion of bicarbonate-rich mucus.
 Stomach lining is replaced every three days.
Gastric mucosal barrier
14.6 Digestive Accessory Organs:
Pancreas, Liver, Gallbladder
 Exocrine pancreas secretes digestive enzymes and an aqueous
alkaline fluid.

Acinar cells secrete proteolytic enzymes,

amylase, chitinase and lipase.
Proteolytic enzymes are secreted in
inactive form.
Activated in the duodenum by enterokinase
and/or trypsin.
Active forms are trypsin, chymotrypsin and
carboxypeptidase.
Each attacks a different peptide linkage.
Schematic
representation
of the exocrine
and endocrine
portions of the
human
pancreas
14.6 Cont’d

 Exocrine pancreas cont’d

 Pancreatic amylase converts

polysaccharides to disaccharides.
 Pancreatic chitinase (fish and some
marine birds) breaks down chitin.
 Pancreatic lipase hydrolyzes
triglycerides into monoglycerides and
free fatty acids.
 Sodium bicarbonate is secreted by
pancreatic duct cells to neutralize acidic
contents arriving from the stomach.
Hormonal control of pancreatic exocrine secretion (mammal)
 Read on your own.
14.6 Cont’d
 Liver cells (hepatocytes) perform a wide variety
of metabolic and secretory tasks.
 Functions of the liver
 Secretion of bile salts
 Metabolic processing of nutrients
 Detoxification of body wastes, hormones and drugs
 Synthesis of plasma proteins and lipoproteins
 Storage of glycogen, fats, iron, copper and vitamins
 Activation of vitamin D
 Removal of bacteria and worn-out red blood cells
 Secretion of hormones
 Excretion of cholesterol, biliverdin and bilirubin
 Synthesis of ascorbic acid (vitamin C)
 Buoyancy in sharks
 Hepatocytes secrete bile into canaliculi.

Anatomy of the mammalian liver

Hepatic portal
circulation in a
human
14.6 Cont’d
Biliary system
 The liver secretes bile, which
is stored in the gallbladder.
 Sphincter of Oddi
 prevents bile from entering the
duodenum except during
digestion.
 Diverts bile secreted by the liver
into the gallbladder between meals
 Bile salts are recycled
through the enterohepatic
circulation.
Schematic
structure
and function
of bile salts
14.6 Cont’d

Bile salts aid

digestion of
fats
Form micelles
around lipid
molecules for
dispersion
through the
watery
contents of the
lumen
14.6 Cont’d
Biliverdin and bilirubin
Biliverdin (green) is made from
the heme portion of worn-out red
blood cells.
In mammals, biliverdin is
transformed into bilirubin
(yellow).
If bilirubin is formed too quickly it
accumulates, causing jaundice.
Stomach lining is protected from digestion by the
gastric mucosal barrier.

Gastric mucosal barrier

14.6 Digestive Accessory Organs:
Pancreas, Liver, Gallbladder
Exocrine pancreas secretes digestive
enzymes and an aqueous alkaline fluid.
Acinar cells secrete proteolytic enzymes,
amylase, chitinase and lipase.
Proteolytic enzymes are secreted in
inactive form.
Activated in the duodenum by enterokinase
and/or trypsin.
Active forms are trypsin, chymotrypsin and
carboxypeptidase.
Each attacks a different peptide linkage.
Schematic
representation
of the exocrine
and endocrine
portions of the
human
pancreas
14.6 Cont’d

 Exocrine pancreas cont’d

 Pancreatic amylase converts

polysaccharides to disaccharides.
 Pancreatic chitinase (fish and some
marine birds) breaks down chitin.
 Pancreatic lipase hydrolyzes
triglycerides into monoglycerides and
free fatty acids.
 Sodium bicarbonate is secreted by
pancreatic duct cells to neutralize acidic
contents arriving from the stomach.
Hormonal control of pancreatic exocrine secretion (mammal)
14.6 Cont’d
 Liver cells (hepatocytes) perform a wide variety
of metabolic and secretory tasks.
 Functions of the liver
 Secretion of bile salts
 Metabolic processing of nutrients
 Detoxification of body wastes, hormones and drugs
 Synthesis of plasma proteins and lipoproteins
 Storage of glycogen, fats, iron, copper and vitamins
 Activation of vitamin D
 Removal of bacteria and worn-out red blood cells
 Secretion of hormones
 Excretion of cholesterol, biliverdin and bilirubin
 Synthesis of ascorbic acid (vitamin C)
 Buoyancy in sharks
Hepatocytes secrete bile into canaliculi.

Anatomy of the mammalian liver

14.6 Cont’d
Biliary system
 The liver secretes bile, which
is stored in the gallbladder.
 Sphincter of Oddi
 prevents bile from entering the
duodenum except during
digestion.
 diverts bile secreted by the liver
into the gallbladder between meals
 Bile salts are recycled
through the enterohepatic
circulation.
Schematic
structure
and function
of bile salts
14.6 Cont’d

Bile salts aid

digestion of
fats
Form micelles
around lipid
molecules for
dispersion
through the
watery
contents of the
lumen
14.6 Bile pigments
Biliverdin and bilirubin
Biliverdin (green) is made from
the heme portion of worn-out red
blood cells.
In mammals, biliverdin is
transformed into bilirubin
(yellow).
If bilirubin is formed too quickly it
accumulates, causing jaundice.
14.7 Small intestine

Small
intestine is
divided into
duodenum,
jejenum and
ileum
Generalized layers of the digestive tract wall, consisting of four
major layers: from innermost  out: the mucosa, submucosa,
muscularis externa, and serosa
14.7 Small intestine cont’d
Motility in the small intestine
Segmentation both
mixes and slowly
propels chyme.
Oscillating ringlike
contractions of
circular smooth
muscle
Initiated by basic
electrical rhythm
periodically
reaching threshold
Frequency declines
along the length of
the small intestine.
 14.7 Small intestine cont’d
Motility cont’d
After absorption is has
progressed to some point,
segmentation is replaced by
the migrating motility complex.
Weak peristaltic waves sweep
remaining contents toward the
large intestine.
14.7 Small intestine cont’d
 Motility cont’d
Ileocecal valve and
ileocecal sphincter
prevent
contamination of
small intestine by
colonic bacteria.
Gastrin causes
ileocecal valve to
relax. (Gastrin is
secreted in response
to food entering
stomach.)
 14.7 Small intestine cont’d
 The small intestine has a large surface area for
absorption.
Inner surface is folded.
Villi are microscopic finger-like projections.
Each villus contains a capillary network and a
terminal lymphatic vessel (central lacteal).
Microvilli are smaller hair-like projections
arising from the luminal surface of
epithelial cells (form the brush border).
These features increase surface area
about 600- fold.
One of the
circular folds
of the small-
intestine
mucosa,
which
collectively
increase the
absorptive
surface area
3-fold
Villus.
Collectively
the villi
increase
the surface
area
another 10-
fold.
A villar epithelial
cell, depicting
the presence of
microvilli on its
luminal border
to form a brush
border; the
microvilli
increase the
surface area
another 20-fold.
 14.7 Small intestine cont’d
Most absorption takes place in
the duodenum and jejunum.
Absorption of carbohydrates
Disaccharides are hydrolyzed into
absorbable monosaccharides by
enzymes in the brush border.
Glucose and galactose are
absorbed by secondary active
transport with Na+ as a
cotransported species.
Carbohydrate digestion
Carbohydrate
absorption for
vertebrate
small intestine
 14.7 Small intestine cont’d

Most absorption takes

place in the duodenum
and jejunum.
Absorption of proteins
Amino acids are absorbed
by secondary active
transport similar to glucose
transport.
Protein digestion for vertebrate
Protein
absorption
for vertebrate
small
intestine
 14.7 Small intestine cont’d
Absorption cont’d
 Absorption of fat

Once micelles reach the epithelial cells,

monoglycerides and free fatty acids
diffuse across the luminal membrane.
Reassembled into triglycerides inside
epithelial cells
Chylomicrons, containing triglycerides,
are extruded by exocytosis into the
interstitial fluid and picked up by
lacteals.
Fat digestion
and
absorption in
vertebrate
small
intestine
14.7 Small intestine cont’d
 Absorption cont’d
 Absorption of vitamins
Water-soluble vitamins are absorbed passively.
Fat-soluble vitamins are absorbed along with fat.
Vitamin B12 must be combined with gastric intrinsic
factor (secreted by the stomach) for absorption in the
ileum.
 Seven liters of secreted digestive juices must be
absorbed daily.
 Diarrhea is passage of a highly fluid fecal matter
usually due to excessive intestinal motility.
Loss of fluids results in dehydration, loss of nutrients,
and metabolic acidosis
The role of countercurrent
exchange in water absorption
14.8 Hindgut/Large Intestine
Vertebrate hindgut
Consists of the cecum,
colon and rectum or
cloaca
Vermiform appendix in
humans and some apes
stores lymphocytes and
has no digestive function.
The colon is made up of
three regions:
ascending, transverse
and descending colon.
14.8 Hindgut/Large Intestine

Vertebrate hindgut
Carnivores have short, simple
colons; omnivores and herbivores
have longer colons.
In amphibians, reptiles, birds and
some mammals the hindgut
terminates in a cloaca, shared by
the renal and reproductive
systems.
 14.8 Large Intestine
 The primary function of the colon is storage of fecal material.
 The colon of a human
 receives 500 ml of indigestible chyme from the small intestine each day.
 Absorbs water and salt
 Symbiotic microorganisms synthesize vitamins (e.g. vitamin K).
 Undigested cellulose adds to the bulk.
14.8 Hindgut/Large Intestine
Motility
Peristaltic contractions propel the
contents toward the rectum, while
antiperistaltic contractions fill the cecum.
Bacteria accumulate in the large
intestine because of the slow colonic
movement.
After meals, mass movements quickly
drive colonic contents into the distal
portion of the large intestine where
material is stored until defecation.
14.8 Hindgut/Large Intestine
 Defecation
Distension of the rectum stimulates stretch
receptors in the rectal wall initiating a
defecation reflex.
Internal anal sphincter (smooth muscle)
relaxes and the rectum and sigmoid colon
contract more vigorously.
Defecation occurs when the external anal
sphincter (skeletal muscle) is voluntarily
relaxed.
Fecal material consists of undigested food
(e.g. cellulose), bilirubin or biliverdin, small
amounts of salt and water, and bacteria.
14.10 Overview of the Gastrointestinal
Hormones
At least 20 possible signal peptides have
been isolated from the mammalian
digestive tract.
Gastrin -- stimulates secretion of gastric juices
and enhances motility in several areas
Secretin -- acts by several mechanisms to
reduce acidity in the duodenum
Cholecystokinin (CCK) -- inhibits gastric
motility and secretion, stimulates secretion of
pancreatic enzymes and release of bile, and
signals satiety
14.10 Gastrointestinal Hormones cont’d

At least 20 possible signal peptides

have been isolated from the
mammalian digestive tract.
Motilin -- stimulates motility in
the stomach and small intestine
Ghrelin -- stimulates growth
hormone release and increases
appetite
Leptin – suppresses appetite