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Lesson 22

Lesson Outline:
General Phylogenetic Trends
Feeding Methods
Specific Features and Phylogenetic Trends
o Food reception
o Food conduction, storage and mechanical digestion (Foregut)
o Chemical digestion and absorption (Midgut)
o Water and ion absorption and defecation (Hindgut)
o Dynamics of gut structure - influence of diet

At the end of this lesson you should be able to:
Describe the different types of feeding methods found in animals and discuss the unique
problems and solutions associated with each
Understand the basic design of the digestive system
Know the three basic types of food processing
Be able to describe the basic functions of each region of the gut
Understand the basic structure/function relationships of different digestive systems
Describe the dynamic component of gut structure/function

Chapter 12: 265-291

Reading for Next Lesson:

Chapter 12: 265-291
Phylogenetic Trends
At its simplest in the protochordates, the digestive system was a tube. These animals used cilia to
create a water current that drew water in through the mouth and over the gills where it could be
strained by the gills. Food particles adhered to mucous lining the gills, and were trapped. The
trapped food and mucous was then
transported to the primitive gut where the
bolus of food was moved along by
peristalsis, and chemical and mechanical
digestion took place, and the digested
substances were absorbed.

In the cyclostomes, the alimentary canal is a straight tube

leading from mouth to anus without coils, or major bends.
The ciliated esophagus runs directly from the pharynx to the
intestine. There is no distinct stomach present (remember they
eat detritus and blood, and small particulate matter rasped from

In the Chondricthys we see the evolution of jaws, teeth, and

active predation. This allowed these fishes to eat large pieces of
food. This requires not just jaws, but teeth. The teeth evolve
from the surface armour. In the Chondricthyes teeth develop
from the dermal denticles. Initially they evolved not for
chewing, but for grasping.

This leads to the evolution of a more complex digestive systems

- a stomach in which to store and mechanically digest the
- a more definitive pancreas to aid in the chemical
digestion of food
- a well developed intestine for absorption of food.

Beginning with the gnathostome fishes, there is considerable

variation in the design of the alimentary canal reflecting the
changes in diet that are seen in individual groups.

The size and length of the stomach, small intestine, and large
intestine reflects the diet of the animal and amount of surface
area required for digestion, and absorption of nutrients and
reabsorption of water.
Feeding Methods
Acquiring food is probably the most time consuming activity of most animals.

A variety of techniques, each with its own associated structures and physiology have evolved for
this purpose.

1) Filter Feeding
Found in animals which ingest aquatic prey (although they don't necessarily live in
aquatic environments).
Food is filtered from water by specialized entrapment devices whether on the body
surface or within it.
Movement of water to the "filter" is usually achieved by muscular movement and
muscular pumps.
In some, the same structure that is used to obtain food for metabolism is also used for gas
exchange for oxygen to metabolize the food.
i.e. - branchial baskets in sea squirts
- gill rakers in tadpoles and fish

In most cases, mucus is used to entrap the food particles at the "filter" and the mucus is
then transported to the gut for digestion - usually by cilia.
Filter feeding supports some of the largest multicellular organisms on earth, such as the
baleen whales.

2) Fluid Feeding
a) Piercing and Sucking (bats, hummingbirds, etc.)
Animals feed on body fluids on animals and plants.
Need adaptations to pierce the epidermal layers of
their food items.
Need a sucking pump to draw the fluids out.
For those that feed on animals, they usually need to
produce an anticoagulant.
Many may inject digestive (proteolytic) enzymes to produce liquefaction of the tissues of
their prey to aid digestion.

b) Cutting and Licking (lampreys, hagfish, etc.)

Cut the body wall of their prey and then lick the body
fluids that leak out from the open wound.
Many have rasping mouth parts.
Many have sharp biting mouth parts to produce
puncture wounds.
Many produce a saliva containing anticoagulants.
Many produce a saliva containing analgesics.
3) Seizing of Prey
a) Jaws, Teeth and Beaks
Found in animals that actually eat their prey.
Are used for holding prey, primarily. Can only be
used for tearing and cutting prey once the
prey item is immobilized.
Swallowing prey whole is very common.
In many, all teeth are the same. In others there is a
high degree of differentiation.
The beaks of birds are as diverse as the teeth of
Many use the crop or gizzard as a secondary
structure for mechanically grinding food.

b) Toxins
To immobilize large prey for subsequent tearing and
dismemberment, many animals use toxins.
Most are neurotoxins acting on the nervous system.
Some are hemolytic - destroy blood cells.
Some are proteolytic.
Must be stored in an inactive form.
Predator must have proteolytic enzymes to digest the toxins
once it ingests the prey.

4) Herbivory and Grazing to Collect Food

Usually requires rasping mouth parts if grazing on encrusting vegetation (not dissimilar to some
cutting and licking strategies).

Requires grinding mouth parts if grazing on taller plants.

Teeth are either very resistant to wear or continuously growing and replaced.
Form and Function of the Digestive System - Specific Features and Phylogenetic Trends

Food Reception
Buccal Cavity
The opening to the alimentary canal.
Consists of organs and structures for feeding and swallowing.
Often must be designed to control the flow of ingested food
and water or air for gas exchange.

Usually receives the secretions of oral glands. Saliva acts as:

- a lubricant (mucus) - aids mechanical digestion
- digestive enzymes - aid chemical digestion
- toxins, anti-coagulants, analgesics

Usually contains chemoreceptors for taste - sensing

what is being ingested and assessing food quality.
The teeth grasp prey and the tongue aids to propel the
food back into the alimentary canal.
In mammals, upper and lower lips meet forward of the
angle of the jaw to form cheeks (and cheek pouches in some species). This allows mammals to
hold and chew food and use the mouth as the first part of the system for chemical digestion.

Behind the buccal cavity lies the pharynx. While these two chambers are distinct in some
animals such as the dogfish) in others they are not and may be collectively referred to as the
buccal-pharyngeal or oro-pharyngeal cavity.
Remember, however, that the pharynx develops from the endoderm of the anterior
foregut while the buccal cavity forms from ectoderm from the stomodeum.
The pharyngeal pouches form in
the walls of the pharynx in cyclostomes Pharyngeal
Reptile Bird Mammal
and fishes. Pouch
While the pharynx is considered 1 Spiracle Tubotympanic Membrane
little more than a passage for air and Thymus
2 Thymus
food in tetrapods, its contribution to Parathyroid
adult structures is huge. 3 Thymus
While it is not essential to know Parathyroid
the details of this table, note that 4 Thymus
derivatives of the pharyngeal arches Parathyroid
become: Thymus
- palatine tonsil (lymphatic 5 Ultimobranchial Body
gland) Thyroid
- parathyroid gland
- thymus gland
- ultimobranchial bodies (all but mammals)
- thyroid gland
The roof of the pharynx gives
rise to the eustacian tubes and
tympanic cavity and the floor
of the pharynx gives rise to the
glottis, larynx and the lung

Two important structures that

appear in the mouth or
oropharyngeal cavity are the
palate and the teeth. We have
discussed these in earlier
lectures and I won't go over
them again now.

Alimentary Canal - Food Conduction, Storage, Digestion

Esophagus and Stomach (Foregut)
The esophagus transports the food to the stomach. The esophagus
opens only to let food into the gut and prevents water from entering
the gut at other times.

Generally only serves as a conducting tube to the stomach.

May contain a crop used to store food in animals that feed

infrequently. Allows animals to store large amounts of food to be
digested slowly at a later time.

The crop may also be used to pre-digest food to be regurgitated for their young.

The stomach stores the meal, adds chemicals to start chemical
digestion and mechanically churns the food to physically break
it down into smaller pieces. The latter increases the surface area
of the food exposed to chemical digestion.

May also serve as a storage site.

Primary site of mechanical digestion.
Frequently the site of the start of protein digestion.

Some animals lack a stomach and food enters directly into the
Carnivores and omnivores have a mono-gastric stomach
consisting of a single chamber. Mechanical digestion is
generally due to muscular action but many animals use other
strategies to assist in the process. Some ingest sand, pebbles, or
stones to aid in grinding food. Some have tooth like structures.
Such stomachs are often referred to as a gizzard.

Herbivores often have digastric stomachs consisting of two

principle chambers.

The first chamber is used to store and pre-process food. Large quantities of food can be eaten
and stored quickly and later regurgitated into the mouth to be chewed more thoroughly aiding
mechanical digestion. This chamber also harbors high
concentrations of micro-organisms. Bacteria and protozoans
break down cellulose to simpler carbohydrates and then ferment
these into butyrate, lactate, acetate and proprionate, also
producing peptides, amino acids and short chain fatty acids in the
process. Most of these compounds can cross cell membranes and
will be absorbed directly through the epithelia of the fore
stomach. The bacteria and protozoans form a symbiotic
relationship with the animal in whose stomach they live - the
animal provides them with mechanically digested plant material
for them to feed on and in return, they digest the plant cellulose,
something that animals can not process well.

In other (non-ruminant) animals, fermentation may occur

simply by storing the food long enough in a crop for micro-
organisms to grow and begin to digest the food before it passes
into the stomach.

The second chamber functions more like the stomach of other

animals secreting digestive enzymes and further breaking
down food substances.

Intestines (Midgut): Chemical Digestion and Absorption

The food is then slowly emptied into the intestines where more digestive chemicals are added.

This is the major site of chemical digestion of proteins, fats and carbohydrates. The duodenum
receives secretions from the liver (the bile) and pancreas. Some of the chemicals that are added
are emulsifiers and these break the fats into small droplets increasing their surface area so that
enzymes can act more readily on them. The other chemicals are enzymes that break apart the
food molecules.
In general, the anterior portions of the midgut are more involved with secretion of digestive
enzymes and chemical digestion and the caudal portions are less involved in secretion and
digestion and more involved in absorption.

Bacteria, protozoans and fungi may also

contribute to chemical digestion here before
being digested themselves. Some are important
sources of vitamins.

Meat requires less processing than plant material

and hence the midgut is shorter in carnivores
than herbivores.

In some air breathing fish, gas is exchanged from

swallowed air bubbles along with the other

Undigested material is eventually voided through the cloaca at the end of the digestive tract.

Intestinal epithelium
The small intestine is the major site of nutrient absorption across the epithelium. Absorption also
takes place in the intestines and it has
adaptations at every level from gross anatomy
to the organelles of individual cells designed
to amplify the surface area available for the
absorption of nutrients.

To begin with, the inner epithelium of the

intestine is folded into circular folds which
increase surface area and slow the movement
of food through the intestine.

The folds, themselves are folded into finger-like extensions

called villi. The villi are the actual absorptive surface of the
digestive epithelium. Each villus contains a network of blood
vessels - arterioles, capillaries, venules and lymphatic vessels
(lacteals). These play an essential role in transporting absorbed
nutrients to the rest of the body.

Each cell of a villus is highly folded producing microvilli. The

fine hair-like surfaces of the cells of the villi are referred to as
the brush-border. They are formed by the cytoskeleton (actin
and myosin filaments) of each cell deforming the plasma

Most of the villus is composed of absorptive cells but there are

also goblet cells which are responsible for producing mucus to
lubricate the digestive epithelium. Adjacent absorptive cells are
held together by desmosomes and tight junctions. The tight
junctions act to prevent any material from entering the body
between the cells of the epithelium.

Only substances that can enter the absorptive cells can enter the
body. The desmosomes act to hold adjacent cells together
despite the strong mechanical forces generated along the gut

In many herbivores, a caecum is usually present at the junction
between the small and large intestines. It contains additional
microorganisms effective in cellulose digestion and provides an
expanded (blind ending) chamber prolonging the time available
for digestion and fermentation. Given its location, beyond the
absorptive region of the gut, in most of these species, the
digested cellulose must then pass through the large intestine
where the water is reabsorbed, and be voided as semisolid feces
to be eaten again so that the nutrients can be absorbed on the
second pass through (the distinction between night and day
feces in rabbits).

Large Intestine (Hindgut): Water and Ion Absorption and

The primary function of the hindgut is water reabsorption. The contents that remain after
nutrients have been absorbed by the midgut consist of indigestible material and both water that
has been ingested but also much water that was secreted to transport digestive enzymes, etc.
along the digestive tract. This is why it is generally referred to as water re-absorption. Animals
living in arid conditions show special adaptations that allow them to re-absorb virtually all of the
water, excreting semi-solid feces.


Over the course of an average day an individual:
Ingest fluid 2L Reabsorb
Saliva 1L Small Intestine 8L
Gastric Juice 2L Large Intestine 0.9L
Bile 1L
Pancreatic Juice 2L Total 8.9L
Intestinal Juice 1L
Total 9L Excreted in Feces 0.1L
The hindgut consolidates the undigested material into feces for defecation.

In many species from arid environments, the hindgut ends in a cloaca rather than an anus. The
distinction here is that the urethra from the bladder terminates in a common chamber with the
undigested food from the intestine, rather than through its own independent opening through the
external genitalia (cloaca is latin for sewer). The advantage of this is that water can then be re-
absorbed from the urine also resulting in the formation of a semi-solid paste and a further
reduction in water loss.

Dynamics of Gut Structure: Influence of Diet

It is estimated that up to 2 x 1010 cells per day are sloughed off and replaced from the human
intestine. The entire midgut lining is replaced every few days. This is expensive. In animals
that are opportunistic eaters, there are strong selection pressures to reduce the size of the gut
during periods of fasting and to increase it following a meal.
Cold and exercise have been shown to increase food intake and with it, the length of the small
intestine in house wrens.

The gut atrophies in hibernating animals and then re-grows in the spring after they come out of
hibernation and start to feed again - the mass of the stomach may increase 3-4 times.

The Burmese python will increase the mass of the small intestine by 40% within hours of eating
a meal after a fast and will double the mass within 2 days. This will increase the surface area for
absorption by up to 24 times.

Even without increasing the length or mass of the gut, animals can alter the number of transport
proteins in the plasma membranes of the absorptive cells to match uptake capacity to the level of
the nutrient intake.

These are all reversible changes designed to provide sufficient uptake capacity when needed but
to reduce the costs of maintaining such an expensive structure when it is not needed.