2.4.

1 Movement of food in the tract

It is similar to that of higher vertebrates, by peristattic waves of ·muscular contraction.
In anterior part of the tract movement is voluntary due to presence of skeletal muscles.
The tongue is not mobile by nself. Many predatory fishes appear to regurgnate large
food nems from the stomach with great facilny. This is possible because of pronounced
development of striated muscles in the wall of the oesophagus to stomach.

2.4.2 Intestinal surfaces

The mouth cavny, oesophagus and stomach are lined with soft mucus membrane as
is in the rest of the tract. There are no salivary glands except in parasnic lampreys.
However, tract wall is liberally supplied wtth glands that secrete mucus which lubricates
passage of food material and protects the gut lining. The gut is highly elastic and
permns larger size food masses. The lining of the small and large intestine are highly
absorptive. The absorptive capacny of these areas is increased by throwing the walls
into lengthwise folds (typhosole), transverse folds (rugae) and finger like projections
(villi). Along the course of the tract there are many gland cells that contribute digestive
enzymes .

2.4.3 Glands and digestive enzymes

There are no digestive juices secreting into the oesophagus. The food passes very
quickly from oesophagus into the stomach. Gastric glands occur in most of the predatory
fishes. These glands secrete gastric juice which contain HCI and pepsinogen, effective
in combination to split large amount of protein molecules. In some carnivorous fishes
gastric acidny of pH 2.4 to 3.6 have been measured. Evidence for stomach enzymes
other than peptidases are not clear. Some fishes (minnows) lack gastric gland and on
this basis, may not possess true stomach. Similarly fish gizzard does not have digestive
glands. HCI produced in the stomach facilitates in

i. disinfecting action killing bacteria

ii. converts disaccharides to mono saccharides.
iii. n activates pepsinogen to pepsin. pepsin is a proteolytic enzyme and hydrolyses the protein
complex food into simple protein molecule in presence of HCI.

Pyloric caeca may have digestive/absorptive function, as absorpture or both. An enzyme

lactase and high levels of saccharase (invertase) has been found in pyloric caeca in
trout, and carp which mainly feed on vegetable matter. Pyloric caeca and intestinal
mucosa are sources of an enzyme lipase which breaksdown fats into fatty acid and
glycerol. The liver is the largest gland of the body which secrete bile - a product of both
excretory and secretory activnies of the organ. The bile is secreted by the hepatic cells into the
bile capillaries and then n is collected into hepatic ducts which join to form a common
bile duct. A cystic duct connects n to the gall bladder. Gall bladder acts as a store
house for continuously secreting bile.
In some fishes, only vestigeal gall bladder is present and in others gall bladder is
completely absent. The bile contains the fat emulsifying bile sa~s alongwtth bile
pigments, biliverdin and bilirubin that originates from the breakdown of RBC and
haemoglobin. This is done in the liver. Biliverdin (green) and bilirubin (red) are the
pigments responsible for the colour of the bile. While bile reaches the intestine, changes
in the bile pigment take place owning to bacterial action. Bilirubin is reduced to
mesobilirubinogen which on further reduction gives stercobilinogen. This on OXidation
gives stercobilin (Brown) and this is responsible for the colour of faeces. The bile sa~s
may not only help to hydrolize the fats but also to adjust the digestive juices of the
intestine to the proper alkalinny for the action of digestive enzymes. (Fat digestion is
facilned by bile juice. Waxes are not digested. Waxes digestion in some birds and
insects is again due to symbiotic bacteria).
9

Besides role in digestion, the liver also act as storage organ for fats and carbohydrates.
It has further important functions in blood cell destruction and blood chemistry, as well
as other metabolic functions such as production of urea and compounds concemed w~h
nitrogen excretion. Liver also acts as a storage organ for fats and vitamin A and D. The
content of vITamins in the liver of tunas is so high that persistant eating of their liver
may lead to hyper-vitaminosis.

Pancreas is a glandular organ lying close to the duodenum. It is formed of exocrine and
endocrine tissues. The exocrine tissue produce pancreatic juice which is carried by the
pancreatic duct into the duodenum. It is neutral to alkaline and is important in digestion
of food. Pancreatic juice contains enzymes for the digestion of proteins, carbohydrates
and fats and nucleic acids. The pancreatic secretion is a complete digestive juice
because ~ contains CHO-splitting, fat-splitting and protein-splitting enzymes.
The c<:lrbq!lydrate splitting enzymes from the pancreas is pancreatic amylase and it
acts upon starch and glycogen in a similar but even more effective way than salivary
amylase of mammals, completing the conversion of ~tarch iota maltose. The !at splitting
enzyme is pancreatic lipas~ It hydrolyzes each molecule into one molecule of glycerol
and t!}ree-of frf!.e fatty acids. The work of lipa~e is facil~ated by the a~tion of bile. There
are no digestive enzymes in the bile. It breaks up la~ge globules of fat into very small
ones, giving much more surface for the enzyme to act on. The protein splitting enzymes
of the pancreatic juice are trypsin and chymotrypsin. Trypsin is secreted in an inactive
form, trypsinogen. This substance is converted into active trypsin by the action of
another enzyme present in the intestinal fluid called the enterokinase. Trypsin and
chymotrypsiO continue the ~reakdown of large protein and polypeptide molecules into
smaller molecules. This process is by hydrolysis with two enzymes' acting inside the
molecules (endopeptidases) rather than at the ends. Following this action, the polypeptides
are broken into much smaller unrts made up of 2,3,4 or more aminoacids linked together.
This is accomplished through the action of enzyme carboxypeptidase. This enzyme is also
present in pancreatic juice.
(Note : The cellulose digestion in all vertebrates is done due to enzymes of bacterial
origin. No vertebrate is able to produce enzyme cellulase. Rumen in ~ow and buffallows
has got strong actions of bacteria on the cellulose and thus they derive large amount
of energy.
The intestinal fluid contains, in addition to enterokinase mentioned above, several
enzymes which are necessary to complete digestion of food to simple absorbable
substances. The small intestine secretes a group of aminopeptidases and dipeptidases
(Erepsin) which complete the breakdown of proteins into aminoacids, each enzyme
being qu~e specific as to which amino acid IT will split off. The intestinal fluid also
contains three inverting enzymes by which the disaccharides are split into monosaccharides.
They are maltase splitting maltose into glucose, lactase splitting lactose to glucose and
galactose and sucrase, splitting sucrose to glucose and fructose.

2.5 Absorption and assimilation of food

Absorption can be defined as '1he passage of food through the lining of the digestive tract into
the blood". In order for digested food to be absorbed, they must be in aqueous solution,
hence, they themselves must be soluble. The component molecules must further be of a size
that will enable them to cross the membrane of the cells lining the tract, pass into the
Circulatory system and ultimately be carried to and enter cells that need them' and store them.
It is interesting to note in this context that fat absorption is intensified in the pyloric stomach of
some fishes and pyloric caeca of others. Fats have been shown to enter into the lymph ducts
in these regions without being spirt into their component fatty acid and glycerol molecules
upon which intestinal absorption depends. No absorption takes place in mouth. Not much
absorption in the stomach also except for simple molecules of glucose to some extent if at
all they are present.
10

2.5.1 Mechanism of absorption

Nearly all organic and inorganic compounds are absorbed in small intestine. Mod~ications S'.
of gut facilitate the process of absorption. The lining of the small and large intestine are highly
absorptive. The absorptive capacity of these areas is increased by throwing the '.
walls into length wise rugae or villi(typlosole). These folds are covered with epijhelium 00
within which is a net work of blood capillaries and also lymph vessels. In absorption,
material passes from the lumen of the intestine through the epithelium and into the
capillaries or lymph vessels by process known as active transport. The active transport
involves movement of materials against concentration gradient and needs energy. Some •
substances on the otherhand penetrate passively and diffuse through these folds.

2.5.2 Absorption of proteins

The amino acids formed due to digestion of proteins in the intestine, do not accumulate
there but are absorbed into the intestinal capillaries and these enter the portal veins,
to be carried into the general circulation by the way of liver. Absorption is more rapid
than simple diffusion. Some absorption of protein derivatives has been shown to occur.
It has been shown that the smallest units into which the proteins are broken down in
the intestine are dipeptides. They apparently leaves to intracellular digestion for the final
breakdown of proteins into single building blocks (amino acids) from which the fish
proteins are resynthesized. The aminoacids absorbed into the blood diffuse through the
body fluids and so reach all the tissue cells. At the same time, most of the tissue
proteins are continually undergoing disintegration to release amino acids which also
enter the circulation and this become the "amino acid pool". From this pool amino acids
are .taken up by the cells to be bui~ up into the cell structure as required. If the cell
takes up as much amino acid as it looses, it is the state of dynamic equilibrium If the
loss is greater, the cell wastes and if the gain is greater the cell grows.

2.5.3 Absorption of carbohydrate

The absorption of sugars from the stomach and colon is very slight. The proximal part
of the small intestine seems to be the chief sije for absorption of sugars. Simple
diffusion can play some part no doubt in the absorption, when concentration in the gut
exceeds that in blood, but the main features of absorption can only be explained by
active transport, requiring energy. The absorption of sugars also depends upon the
presence of Na+ ions, probably also on N.a: K ratio. However, their remains much to be
found about the details of sugar absorption.

2.5.4 Absorption of fats

Fat absorption from the intestine is not clear. It is shown that long chained fatty acids
are .absorbed very largely into the lymph, whereas, short chained fatty acids enter the
portal blood. It has also been pointed out by some that efficient fat absorption requires
both lipase enzyme and bile salts .

PHYSIOLOGY OF DIGESTION IN SHELLFISHES

In the aquatic environment, particularly in the sea, the crustacea have exploijed every type of
niche and this ecological diversity is paralleled by the diversity of food eaten. The micro-
crustaceans typically feed on microalgae, whereas larger crustaceans range from burrowing
detrijus feeders to active predators of molluscs and fish. Most of our knowledge of digestive
physiology is derived from the larger decapoda, although there has been appreciable research
on the Amphipoda and Isopoda. Earlier knowledge of function was fragmentary, but over the
last 3 decades experimental research on digestive physiology, using modern methods has
grown considerably. However, there are still very large gaps in our knowledge and much
remains to be done.

The principal regions of the gut and their general functions

There is a considerably body of older I~erature describing the anatomy of the gut of Crustacea.
Many recent publications, however, describe detailed investigations of regions of the gut,
particularly foregut and the unrastructure or histochemistry of the midgut. The gut in Crustacea
is basically a simple tube running virtually the length of the body, from the anteroventral mouth
to the anus at the end of the last body somite. The foregut and hindgut, derived from
embryonic ectoderm are lined with cuticle and only the midgut, derived from embryonic
endoderm, has cells in direct contact with the lumen of the gut. Storage, tr~uration and early
digestion may therefore take place in the foregut. It has been reported that up to 12% sugars
may pass through the walis of foregut but it is probable that most absorption normally occurs
via the midgut. Various glands associated w~h oesophagus have been described, but their
function appears to be lubricatory rather than digestive. Secretion of enzyme is lim~ed to the
midgut, while faecal formation and defecation is the usual role of the hindgut. In larger
Crustacea, a mechanism is needed to mix the much larger mass of food in the foregut w~h the
digestive enzymes from the midgut. If the food particles are large, digestive efficiency will be
improved by trituration and complex gastric mills have evolved in the Decapocla.

The foregut: Structure and Function

Structure: The foregut or proventriculus is always divided into an anterior distendable part
that usually serves as a crop in macrophagus feeders. The posteriorend of this chamber
constructs as a gastric mill, comprised principally <i median dorsal ossicle and two lateral
ossicles. The gastric mill leads into posterior part of the proventriculus, which is in turn divided
into dorsal and ventral chambers. The dorsal chamber, which bears lateral grooves, leads into
the midgut or directly into the hindgyt. The ventral chamber contains the filter press (Wshaped)
which leads into the digestive gland. The floor of the anterior proventriculus bears a
median groove and two ventro-Iateral grooves, with fringing dense setae. The ventro-Iateral
grooves lead to the finer press. There is a complex system of muscle attachment over the
surface of the proventriculus, particularly around the gastric mill.

Function: As the food enters the anterior chamber of the proventriculus it is penetrClted by
the fluid from the digestive gland that flows towards dorso-Iaterally in grooves in the posterior
chamber. Trituration and further mixing with fluid occurs at the gastric mill ossicles. The food
mass is continually being manipulated by the lateral plates of the anterior chamber and forced
into the gastric mill. Eventually the fluid passes from the food mass into the ventral grooves
of the anterior chamber. Dense setae exclude larger particles and the fluid passes backward
through the filter-press which excludes particles above 1 um and finally into the openings of
the digestive gland. Fluid from the digestive gland is pump~ dorsally into the dorso-Iateral
grooves, joined by the fluid squeezed from the food mass in the posterior chamber. Some fluid
is also pumped in and out of the anterior diverticulum of the rTlidg,IJt. The combined fluid then
pass forwards to the anterior chamber. The circulation is driven by the pumping action of finer
press and associated structures. We do not know what happens to the fluid that enters
digestive gland. Probably dissolved nutrients are absorbed and the fluid with add~ion of more
enzymes returns to proventricular circulation. This will be the fru~ area of research to be
investigated.

3.3 The midgut: Cell structure and Function

The digestive gland is comprised of a large number of simple, fragile tubules, each tubule is
invested w~h only a thin layer of connective tissue. (therefore autolysis is rapid). Because the
midgut serves the dual role of enzyme secretion and absorption of digested food. The ep~helial
layer is differentiated into two cell types one w~h microvilli border i.e. absorptive cells (R) and
other w~hout i.e. secretory cells (F). Both these cells actually arise from E cells (embryonic
cells).
R Cells (absorptive)
E (Embryonic cells)
F Cells (Secretory)
Sometimes the F cells appear to develop into B cells with a large single vacuole containing
digestive enzymes. R-Cells absorb and store nutrients and F-Cells synthesise digestive enzymes.
The secretion is of holocrine in nature (whole things liberated into lumen).

3.3.1 Digestive enzymes

Proteolytic enzymes: Crustaceans contain proteolytic enzymes similar to mammalian
enzymes. There are reports of absence of some enzymes e.g. Chymotrypsin but
controversy still exists. There are reports about the presence of trypsin. They are
similar to mammalian trypsin in many characters. The crustacean enzymes have an acidic
iso·electric point and they are irreversibly inactivated in acid solutions, they do require ca+ for
stability and they are resistant to autodigestion. The crustacean trypsin like proteinases differ
from mammalian trypsin in that they attack undenatured protein. Apart from trypsin there are
other proteinases in crustacea but very less work on this has been done. In Homarus seven
proteolytic enzymes have been reported. The presence of various peptidases has been reported
mainly based on studies with synthetic substrates and crude mixtures. Presence of
earboxypeptidases, aerylamidase, and dipeptidase have been reported in the foregut and
digestive gland. The dipeptidases may be associated with the brush border of the absorptive
cells in the digestive gland, zymogens. No zymogenS of crustacean proteolytic enzymes have
been found. If this so, how do the secretory cells protect themselves from proteolytic activity.
Whether zymogens are synthesized in crustacean is still open question.

3.3.2 Lipid digestion

The digestion of lipids has been given less attention than protein digestion. Lipase and
esterase activ~ies have been demonstrated in many crustaceans. Lipase has been
partly purified from the foregut of lobster and shown to have molecular wt.43,OOO which
is similar to that of hog pancreatic lipase. Lipase helps to breakdown long fatty acids
into ~·diacyglycerol and x·monoacyglycerol, in this form only probably they are absorbed
in the gut. Micelles of sterols and lecithins and readily formed by a fatty aeyltaurine
thus promoting their absorption.

3.3.3 Carbohydrate digestion

Hydrolysis of carbohydrate is a complex process involving transglyeosylation and
condensation. Some glycosidases have been also determined. Thus many of the different
carbohydrases that have been reported as present in the digestive tract of Crustacea
may not exists. The glycosidases actually attack dissacharides and oligo· saccharides
and make them absorbable.

Starch and similar compounds: x·Amylase activ~y has been demonstrated in all
Crustacea. For digestion of starch and glycogen two other enzymes viz oligo x-1-6
glucosidase and maltase have been isolated.
1
Cellulose : Cellulase enzyme hydrolysases the cellulose component. The main
components are endo ~ 1.4 glucanase and exo ~-1.4 glucanase and ~-glucosidase.
Detailed studies have not been made on cellulolytic enzymes in Crustacea. Probably
many Crustacea synthesize cellu/ases but cellulase activity may be due to microorganism
in the gut and in food prior to ingestion. Cellulase may have two functions in digestion
1) to convert cellulose to glucose as energy source and 2) to enable other digestive
enzymes to penetrate a plant cell wall.
Glucans : Glucans are structural polymers present in many algae. fungi. protozoans.
therefore poten~al source of energy for crustaceans that feed on algae or micro-organism.
Glucanases enzyme occurs in crustacea together with cellulases. amylases and
chitinases for digestion of carbohydrates.
Chitin: Chitin is the main structural carbohydrate in crustacean skeleton and many of
them they eat their own exuviae. This is digested by an enzyme chitinase and chitobiase.
The removal of digestive fluid together w~h rise in pH and association of other compound
such as mucopolysaccharide with peritrophic membrane may inhibit residual chitinase
activity.

3.3.4 Micro-organisms in the digestive tract

Bacteria are found in the digestive tract of most crustaceans and they derive very little
benefit from them. Micro-organism may be a good sources of digestive enzymes for
polysaccharides that can not be digested by animals rtsen . Other possible function of
microbial activrty in the digestive tract is the supply of vitamins and essential amino
acids.

3.3.5 Peritrophic membranes

Peritrophic membranes are formed in large number of crustacea. In decapoda the
cylindrical peritrophic membrane is secreted by midgut eprthelia cells just behind the
digestive gland opening and contains the feces in a characteristic long pellets. It is
shown that peritrophic membrane is thought to protect the gut from abrasive material
in the feces. Usually the membrane disintegrates soon after its egestion.

3.4 The Hindgut

In higher crustacean the hindgut is a long tube and has internal longitudinal folds containing
longitudinal muscle bundles with an outer layer of cuticular muscle. In penaeid shrimp
sixsmooth surface pads containing spongy tissue fill the lumen of the gut. These pads taper
posteriorly to longrtudinal ridges of the muscle. The obvious function of the hindgut is
defecation. In shrimp the congrtudinal pads grasp the fecal pellet in its peritrophic membrane
and rhythmically expel it. The muscular hindgut also pump water into the gut. Anal drinking of
water has been also observed in many crustacean. This is mainly to assist defecation process
but it also assist in removal of electrolytes excreted into the gut. Further research is needed to
assess the exact functions.