School of Veterinary Medicine

Isabela State University- Echague

CPSG,DVM

CARBOHYDRATES

Objectives:
o To familiarize with the water nutrients, there components functions and
main role in animal growth.
o To know the different classification of carbohydrates and its composition.
o To know the process of carbohydrates digestion in ruminant and non
ruminant animal.

Content:
o Introduction to Carbohydrates
o Functions of Carbohydrates
o Classification of Carbohydrtaes
▪ Sugar
- Monosaccharides
- Oligosaccharides
▪ Non Sugar
- Polysaccharides
- Complex Carbohydrates
o Carbohydrates digestion in the rumen
o Carbohydrates Metabolism in ruminant
o Digestion of Carbohydrates in non ruminant
o Carbohydrates Metabolism
o Glucose Metabolism

Introduction

Carbohydrates may be defined as polyhydroxy aldehyde, ketones or acids and

their derivatives or compounds that yield these derivatives on hydrolysis. The
carbohydrates are neutral chemical compounds containing the element carbon,
hydrogen and oxygen, with the last two elements present in the same proportion
as in water mostly, but not at all. One of the example of carbohydrate where such
ratio is not found in the sugar deoxyribose which is a constituent of DNA.
Whereas acetic acid and lacticacid can be represented as hydrates of carbon but
are not carbohydrates.
The carbohydrates serve as both structural and reserve material in the plant. The
animal body contains less than 1 percent carbohydrate, which are present in
blood, muscles and liver. The carbohydrate present in animal body is also known
as animal starch or glycogen.

Based upon their digestibility and solubility, the carbohydrates can be divided
into two groups.

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(a) Soluble carbohydrates: They are called nitrogen free extract (NFE) and
include simple sugar, starch and hemicellulose, which are easily digestible in the
body.
(b) Insoluble carbohydrates: They include hard fibrous substance like crude
fibre, cellulose and lignin. They are less digestible by non-ruminants and easily
digested in ruminants by rumen microflora and microfauna.

Functions of Carbohydrates:

1. Carbohydrates serve as a major source of energy in animal body.

2. They are essential components of production, temperature control and proper
functioning of the different parts of the animal body.
3. They are essential components of milk as lactose.
4. They are stored as glycogen, excess of carbohydrates in the diet is converted
into fat and stored in the fat depot. These are reserve energy materials of the body
in liver and muscles of animals and starch in plants.
5. Carbohydrates are helpful in absorption of calcium and phosphorus in younger
animals.
6. They help the secretion of digestive juices in gastrointestinal tract.
7. They provide suitable environment for the growth of rumen bacteria and
protozoa.
8. They help in peristaltic movement of food.
9. They maintain the glucose level of plasma.
10. They are also component of several important bio-chemical compounds such
as nucleic acids, coenzymes and blood group substance.
11. They play a key role in the metabolism of amino acids and fatty acids.

Classification Examples
Monosaccharides
Trioses (C3H6O3) Glyceraldehyde and dihydroxyacetone
Tetroses (C4H8O4) Erythrose
Pentoses Aarabinase, xylose, xylulose, ribose, ribulose, and 5
(C5H10O5) deoxyribose
Hexoses Glucose, fructose, galactose, and mannose
(C6H12O6)
Heptoses Sedoheptulose, mannoheptulose (in avocados), and L-
(C7H14O7) glycero-D-manno-heptose
Oligosaccharides
Disaccharides Sucrose (D-α-glucose and D-α-fructose), lactose (milk sugar;
D-α-glucose and D-α-galactose), maltose, isomaltose,
cellobiose, ,α-trehalose, α,β-trehalose, and β,β-trehalose)

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CPSG,DVM

Trisaccharides Raffinose, kestose, maltotriose (three units of glucose),

planteose, and melezitose (in sweet exudates of many trees
and in insects), and panose (synthesized by microbes)
Tetrasaccharides Stachyose and lychnose (1-α-galactosyl-raffinose)
Polysaccharides
Homoglycans Pentosans (C5H8O4)n, for example, arabans and xylans
Hexosans (C6H12O6)n, for example, starch, cellulose,
mannans, levans and glycogen.
Heteroglycans Hemicelluloses, pectins, exudate gums, seaweed
polysaccharides (algin, carrageenans, agar,
aminopolysaccharides [e.g., chondroitin and hyaluronic
acid], and sulfated polysaccharides [e.g., chondroitin sulfate)
Conjugated carbohydrates
Glycolipidse Glyceroglycolipids and sphingolipids
Glycoproteins Mucins, immunoglobulins,
Mucins, immunoglobulins,
and membrane-boundand hormone
receptors membrane-bound hormone
receptors

Classification of Carbohydrates: The carbohydrates are usually divided in to two

major groups:

I. Sugars: The term sugar is generally restricted to those carbohydrates, which

contain less than ten monosaccharide residues. Sugars are divided into two
groups.
o Monosaccharides
o Oligosaccharides

MONOSACCHARIDES:

Monosaccharides: The simplest sugars are the monosaccharides and they can not
be hydrolysed into smaller units under reasonably mild conditions. They are
divided into sub-groups depending upon the number of carbon atoms present in
the molecule.
e.g.
o Triose (C3H6O3)
o Tetroses (C4H6O4)
o Pentoses (C5H10O5)
o Hexoses (C6H12O6)
o Heptoses(C7H14O7)

The 4 common hexoses:

• D-Glucose

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• D-Mannose
• D-Galactose
• D-Fructose

Sugar containing an aldehyde (CHO) group are classed as aldose.

e.g. glucose, mannose and galactose.

Whereas sugars containing a ketone group are classed as ketoses.

e.g., fructose.

Presence of active aldehyde and ketone group in monosaccharide act as reducing

sugar substances. They can be oxidized to produce number of acids like gluconic
acid, glucaric acid and glucoronic acid. The reducing properties of these sugars
are usually demonstrated by their ability to reduce certain metal ions such as
copper or silver in alkaline solution.

PENTOSES:

Pentoses have the general formula C5H10O5. The most important member of this
group are the aldoses, L-arabinose, D-xylose and D-ribose, and the ketoses, D-
xylulose and D-ribulose.

▪ L-Arabinose: Occurs in pentosans as arabans. It is a component of

hemicellulose and gum and present in silage.
▪ D-xylose: Also occurs in pentosans in the forms of xylans. These
compounds form the main chain in grass hemicellulose and xylose along
with arabinose produce in considerable quantities when herbage is
hydrolysed with normal sulphuric acid.
▪ D-Ribose: It is present in all living cells as a constituent of ribonucleic acid
(RNA) and it is also a component of several vitamins and coenzymes.

HEXOSES:

Glucose and fructose are the most important naturally occurring hexose sugar,
while mannose and galactose occur in plants in a polymerized form as mannans
and galcutans.

▪ D-Glucose:

This sugar occurs in plants, fruits, honey, blood and other body fluid. Glucose
is the major component of many oligosaccharide, polysaccharide and
glucosides. In the pure state, glucose is a white crystalline and soluble in
water.

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▪ Fructose or fruit sugar: It occurs free in green leaves, fruit and honey. It
also occurs in disaccharides-sucrose and in fructosans. It differs from other
sugars in being laevo-rotatory and also known as fruit sugar.

▪ Mannose: It occurs in polymerized form as mannan. It does not occur free

in nature.

▪ Galactose: It is a constituent of disaccharide lactose, which occurs in milk

and is also a component of gum, mucilages, pigments etc. It does not exist
free in nature as Mannose.

HEPTOSES:

▪ Sedoheptulose is an important example of a monosaccharide containing

seven carbon atoms. This heptose occurs as the phosphate, as an
intermediate in the pentose phosphate metabolic pathways.

OLIGOSACCHARIDES:

Oligosaccharides: The oligosaccharide (Oligo=few) includes all sugars other than

the monosaccharides. The monosaccharides linked together with a elimination of
water at each linkage and-produces di, tri, tetra or polysaccharide containing 2,3,4
and large number of simple sugar molecules, respectively.
o Disaccharides
o Trisaccharides
o Tetrasaccharides

Disaccharides: The most frequently occurring oligosaccharides in nature are

disaccharides, which on hydrolysis yield two moles of simple sugar.

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Disaccharides consist of two molecules of hexose sugars combine together with

loss of one molecules of water.

2 C6 H12 O6 = C12 H22 O11+ H2O

The most important disaccharides are sucrose, lactose,maltose and cellobiose.

▪ Sucrose, cane sugar, beet sugar or sacchrose: It is the familiar sugar of
domestic use, widely distributed in nature and occurs in most of the
plants. Sucrose is easily hydrolysed by the enzyme sucrase or by dilute
acids and gives one molecule of a-D glucose and one molecule of β-D-
fructose.

▪ Lactose or milk sugar: It occurs in milk only as a product of mammary

gland. Cow's milk contains 4.6 to 4.8 percent lactose. It is not as soluble
as sucrose and is less sweet, imparting only a faint sweet taste to milk.
On hydrolysis it produces one molecule of glucose and one molecule of
galactose.

▪ Maltose or malt sugar: It is produced during the hydrolysis of starch and

glycogen by dilute acids or enzymes or during the germination of barley
by the action of the enzyme amylase. The barley after germination and
drying is known as malt and is used in the manufacture of beer and
scotch malt whisky.

▪ Maltose is water-soluble but it is not as sweet as sucrose. On hydrolysis

it yields two molecules of glucose.

▪ Cellobiose: Cellobiose does not exist naturally as a free sugar, but is the
basic repeating unit of cellulose. It is less soluble and less sweet.

Trisaccharides: The unions of three molecules of hexose sugars.

3 C6 H12 O6 = C18 H 32 )16 + 2H2O

Raffinose distributed widely in plants. On hydrolysis this sugar produces glucose,

fructose and galactose. It is a nonreducing sugar.
Tetrasaccharides: Tetrasaccharides are produce by the union of four hexoses
residues.
4C6 H12 O6 = C24,H42 O21+ 3 H2O

Stachyose is an example of tetrasaccharide, which is a nonreducing sugar, and

hydrolysed to two molecules of galactose, one molecule of glucose and one
molecule of fructose.

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II. Non-sugars: They are tasteless, insoluble, amorphous compounds with a high
molecular weight.

Poly saccharides are divided into two sub groups.

(1) Homopolysaccharides: They are classified according to the kind of sugar,

which produce on hydrolysis. For example, glucans are condensation polymer of
glucose, fructans of fructose and xylans of xylose. This group of polysaccharides
are a polymers of monosaccharides derivatives, such as sugar acid (eg.,
galacturonans) and sugar amines (e.g. glucosaminans).

(2) Heteropolysaccharides: They are mixed polysaccharides, which on hydrolysis

yield mixtures of monosaccharides and derived products.

Homopolysaccharides:

Starch: The reserve materials of most plants consist primarily of starch. When
this is hydrolyzed with acids or enzymes, it is changed into dextrin, maltose and
finally into glucose. In food this exists as a straight chain of glucose units called
amylose, mixed with a branched chain structure called amylopectin. The quantity
of amylose can be estimated in starch by a characteristic reaction with iodine,
amylose produces a deep blue colour while amylopectin solution produce a blue
violet or purple colour.

Amylose + Iodine ------ Deep blue colour

Amylopectin + Iodine ------ Blue violet or purple colour

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CPSG,DVM

Amylose is composed of linear molecules in which the D-glucose residues are

linked between carbon atom 1 of one molecule and carbon atom 4 of the adjacent
molecule whereas amylopectin has a bush-like structure containing primarily -
1,4 linkages but the molecule also contains side chains in which carbon atoms 6
of glucose residues is linked with carbon atom 1 of the other glucose. Starch
granules are insoluble in cold water, but when the suspension water is heated the
granules swell and eventually the granule sacs rupture and a gelatinous is formed.

Amylose Amylopectin
1. α- 1,4 linkage between α- 1,4 linkage in straight chain and α l,6
glucose unit linkage in branched chain are present
2. Only straight chains is there
Straight as well as branched chains are
present
3. Iodine test gives deep blue Iodine test gives blue violet or purple color.
colour.

Glycogen: The small amount of carbohydrate reserve in the liver and muscles in
the form of glycogen, which is also called “Animal starch”. They form colloidal
solutions, which are dextra-rotatory. Glycogen is the main carbohydrate storage
productin the animal body and plays an essential role in energy metabolism.

Dextrins: These are intermediate products of the hydrolysis of starch and

glycogen.

Starch + Glycogen = Dextrin = Maltose = Glucose

Dextrins are soluble in water and produce gum like solutions. The higher
members of these transitional products produce a red colour with iodine, while
the lower members do not give a colour. The presence of dextrin gives
characteristics flavour to bread crust, toast and partly charred cereal foods.

Cellulose: It is glucan and is the most abundant plant constituent, farming the
fundamental structures of the plant cell walls by farming chemical linkages with
hemicellulose and lignin. Cellulose molecule contains between 1600 to 2700 β-
D-glucose units. Cellulose is more resistant to chemical agents than the other
glucosans. On hydrolysis with strong acid glucose is produced. Enzyme produced
by germinating seeds, fungi and bacteria attack cellulose and produce cellubiose,
which acted upon by enzyme cellubiase and produces glucose. It is fermented in
the rumen by the microbial enzymes and produces volatile fatty acids like acetic
acid, propionic acid and butyric acid.

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CPSG,DVM

Frutosans: It occurs as reserve material in roots, stems, leaves and seeds of a

variety of plants. Fructans are hydrolysed to D-fructose and of D-Glucose. Inulin
is the known polysaccharides belong to this group.

Galactans and Mannans: These are polysaccharides, which occur in cell wall
of plants. It is a component of palm seeds, clovers and Lucerne.

Pectin: The term pectic substance is used to refer to a group of plant

polysaccharides in which D-galacturonic acid is the main constituent in which
some of the free carboxyl groups are esterified with methyl alcohol and others are
combined with calcium or magnesium ions. D-galactose and L-arabinose are also
present as additional components. Pectic substances are found in peel of citrus
fruit, sugar beet pulp. Pectinic acid posses gelling properties and are used in Jam
making.

Chitin: It is a major constituent of the exoskeleton of insects and crustacea. It is

the only known example of a homopolysaccharide containing glucosamine being
a linear polymer of acetyl-D-glucosamine. Next to cellulose, it is probably the
most abundant polysaccharide in nature.

Heteropolysaccharide:

Hemicellulose: The hemicellulose is a group of substances, including araban,

xylan and certain hexosans and polyuronides, which are much less resistant to
chemical agents than cellulose. It is insoluble in boiling water but soluble in dilute
alkali and hydrolyzed by dilute acids to simple sugar and uronic acid such
as glucuronic and galacturonic acid.

Gum arabic: It is a useful plant gum and produced from the wound in the plant,
although they may arise as natural exudates from bark and leaves. Acacia gum
has long been familiar substance; in hydrolysis it yields arabinose, rhamnose
and glucuronic acid.

Mucilages: Mucilages are found in few plants and seeds. Linseed mucilage
produces arabinose, galactose, rhamnose and galacturonic acid on hydrolysis.

Agar: It is sulphated polysaccharides. They are found as constituents of seaweeds

and in mammalian tissues. It is used as a gel-farming agent in microbial studies.
Agar is a mixture of at least two polysaccharides containing sulphate ester of
galactose, glucuronic acid and other compounds.

Hyaluronic acid: It is grouped under amino polysaccharides. It is present in the

skin, synovial fluid and umblical cord. Solutions of this acid are viscous and play

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an important role in the lubrication of joints. Hyaluronic acid is composed of

alternating units of D-glucuronic acid and N-acetyl D- glucosamine. Chondroitin
is chemically similar to hyaluronic acid but contain galactosamine in place of
glucosamine. It is a major component of cartilage, tendons and bones.

Heparin: It is an anticoagulant, which occur in blood, liver and lung. On

hydrolysis heparin yields glucuronic acid, glucosamine and sulphuric acid.

Lignin: The woody parts of plants contain a complex indigestible substance called
lignin. Lignin is a high molecular weight amorphous polymer containing carbon,
hydrogen and oxygen. Lignin is not a carbohydrate but because of its association
with carbohydrate it is usually discussed along with carbohydrates. There is a
strong chemical bonds existing between lignin and many plant polysaccharides
like cellulose. Lignin is resistant to strong acids and microbial action in the
rumen. It is considered to be indigestible by the animals and is responsible for
poor digestion of wheat straw and paddy straw.

Carbohydrate digestion in the rumen:

The major portion of the ruminants diet consist of cellulose, hemicellulose and
other carbohydrates which cannot be hydrolyzed by the enzymes secreted by the
animals in the digestive tract but broken down by enzymes secreted by rumen
microorganisms with the production of volatile fatty acids and gases.

The bacteria, which help in carbohydrate digestion, are as follows:

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The soluble carbohydrates are rapidly fermented, starches are less rapidly
fermented, whereas, the structural carbohydrates like cellulose and hemicellulose
are slowly fermented. All carbohydrates are converted into pyruvic acid.

The bacteria and protozoa mainly responsible for fermentation in the digestive
tract are mainly strict anaerobes although, there may be a small number of
facultative anaerobes. The normal concentration of bacteria in rumen liquor is
1011 bacteria per ml. and protozoa are 106 per ml of rumen content.

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Volatile fatty acid production in rumen: The feeds, which is ingested by the
animals broken down to volatile fatty acids like acetic, propionic and butyric
acids via pyruvic acid. Higher fatty acids like valeric and isovaleric acid etc. are
also formed in smaller amounts. With normal diets the predominant acid is acetic
acid followed by propionic acid and butyric acid. Volatile fatty acids represent in
the following proportions.

1. Acetic acid 60-70 percent

2. Propionic acid 15-20 percent

3. Butyric acid 10-15 percent

4. Valeric and isovaleric acid present in traces.

On an exclusive roughage diet the production of acetic acid is highest. As the

concentrates in the diet are increased, the production of acetic acid reduces and
that of propionic acid increases. Lactic acid is also formed as an intermediate
product but is fermented to acetic and propionic acid. Mature fibrous forage give
rise to VFA mixture with high proportion of Acetic acid (about 70%). Less mature
forage tend to give a lower acetic acid and higher proportion of propionic acid.
On concentrate feeding diet the acetic acid predominates if the rumen ciliate
protozoa survive. The proportion of fatty acids production is changed under
following condition:

1. High ratio of concentrates in the ration.

2. Fine ground forages,
3. Lack of physical fibrousness.
4. Green fodder low in fibre and high in soluble carbohydrates.
5. Pelleted concentrates.
6. Heated concentrates.
7. High starch diet.

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This will bring relatively high ratio of propionic acid to acetic acid. The
conversion of pyruvate into different volatile fatty acids is shown below.

GAS PRODUCTION IN THE RUMEN:

With the fermentation of carbohydrates by the bacteria, gases are also produced.
Carbon dioxide and methane at present as are principal gases. The rate of gas
production in the rumen is most rapid immediately after a meal and in the cow
may exceed 30 Liters/hour. The typical composition of rumen gas is given below:

1. Carbon dioxide 40 percent

2. Methane 30-40 percent
3. Hydrogen
4. Oxygen and nitrogen (small amount ingested from air).

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Carbon dioxide is produced partly as a by-product of carbohydrate fermentation

and partly by the reaction of organic acid with bicarbonate of carbon dioxide by
the methanogenic bacteria. Hydrogen, formate and succinate are hydrogen donors
for this reaction. The quantity of methane gas formed depends upon the type of
food eaten. About 4.5 g of methane is formed for every 100 g of carbohydrates
fermented (digested) and the ruminant loss about 7 percent of its food energy as
methane.

Most of the gases in the rumen is lost by eructation. Under metabolic disorders
the gas is trapped in the rumen and the animal is unable to remove the gases and
a condition known as bloat occurs.

ABSORPTION OF VOLATILE FATTY ACID:

Most of the volatile fatty acids are absorbed directly from the rumen, reticulum
and omasum. Small amount may pass to abomasum and small intestine from
where they are absorbed. Portion of these volatile fatty acids are used by bacteria
and protozoa to synthesize their own polysaccharides or use as a carbon skeleton
for the synthesis of their body protein.

Carbohydrates metabolism in ruminants:

In ruminants considerable amounts of volatile fatty acids (Acetic, propionic and

butyric acids) are produced from the carbohydrate breakdown in the rumen. The
acids then pass across the rumen wall, where a little amount is converted to lactate
and remaining is metabolized in the liver. The net gain of ATP per mole of acetic
acid, propionic acid and butyric acids are 10, 17 and 25, respectively.

Metabolism of volatile fatty acid:

1. Acetic acid Metabolism: It is the major volatile fatty acid present in blood
and absorbed as such. It is utilized for energy and is also a precursor of
fatty acid (Short chain fatty acid of milk fat). It is never converted to
glucose.

2. Propionic acid Metabolism: Propionic acid, which is produced in rumen,

is carried out to the liver where it is changed into glucose.

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The energy balance sheet is as

Mole ATP
+ -
2 moles propionate to 2 moles succinyl COA - 6
2 moles succinyl COA to 2 moles malate 6 -
2 moles malate to 2 moles phosphoenol pyruvate 6 2
2 moles phosphoenolpyruvate to 1 mole glucose - 8
1 mole glucose to CO2 and H2O 38 -

Net gain ATP per mole 50 – 16 /2 = 17

So 17 moles of ATP are produced per mole of propionic acid.

3. Butyric acid Metabolism: It is absorbed as aceto acetic acid and b - hydroxy
butyric acid in its passage across the ruminaland omasal walls. It is ketogenic in
nature and utilized for synthesis of long chain fatty acid of milk fat.

Digestion of carbohydrates in non-ruminants:

1. Digestion in the mouth:

o Food, mixed with the saliva, which contains the enzyme ptyalin (a-
amylase). This enzyme hydrolyzes starch into the maltose and
isomaltose. But the food remain in the mouth for a short time and
about 3 to 5 percent of the starch hydrolyzed into maltose.
o The amylase enzyme hydrolzyes the a-1, 4- glucosidic bond in
polysaccharide. When amylose, which contain a-1,4- glucosidic
bond is attacked by a-amylase, random cleavages of these bonds
give rise to a mixture of glucose and maltose.
o Amylopectin, on the other hand contains in addition to a-1, 4-D-
glucosidic bond, a number of branched a-I, 6-D-glucosidic bonds

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which are not attacked by a-amylase and the product includes a

mixture of branched and unbranched oligosaccharides (dextrin) in
which 1, 6-D-glucosidic bonds are abundant.

2. Digestion of carbohydrates in the stomach:

o The action of a-amylase enzyme of saliva continues for about 30 to
50 minutes after the food has entered the stomach, that is, until the
content of the fundus are mixed with the stomach secretions.
o Then the acid of the gastric secretion blocks the activity of the
salivary amylase. The acid of the stomach juice can hydrolysed
starch and disaccharides to a slight extent.

3. Digestion of carbohydrates in intestine:

o Pancreatic secretions contain large quantities of a-amylase which is
capable of splitting starch into maltose and isomaltose in intestine.
In general, the starch is almost totally converted into maltose and
isomaltose before they have passed beyond the Jejunum. The
epithelial cells of the small intestine contain the four enzymes

▪ Lactase, which split lactose into glucose and galactose.

▪ Sucrase, which split sucrose into glucose and fructose.
▪ Maltase, which split maltose into two molecules of glucose.
▪ Isomaltase, which split isomaltose into two molecules of glucose.

o Thus the final products of carbohydrate digestion that are absorbed

into the blood are all monosaccharides. The enzyme hydrolyses
starches into glucose and other carbohydrates into final products.

Starches

Salivary L-amylase Pancreatic L-amylase

Maltose + Isomaltose

Lactose Sucrose
Lactase Maltase Sucrase
Isomaltase

Galactose Glucose Fructose

DIGESTIVE JUICES AND THEIR FUNCTIONS:

Secretion Gland Principal Components Functions
Saliva (Salivary Water, Mucous Salt Soften and lubrication
gland) of food. Provide neutral
medium for action of

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salivary amylase and

help to preserve teeth
against acid formed by
bacteria.
Salivary Amylase Act on starch and split
into dextrin
and maltose.
Gastric juice Water, Mucous Further soften of food.
(Gastric gland) Prevent gastric juice
from damaging the
stomach wall
Hydrochloric acid Stop the action of
salivary amylase
and allow pepsin to
work and kills
microorganisms.
Pepsin (Secreted as Protein moieties of the
a pepsinogen) food are hydrolysed
into proteases, peptone
and polypeptides and
curdle of milk
in adults when rennin
enzymes are
absent.
Rennin Milk casein is
converted into curds
such as paracassinate,
which is easily
attached by other
protein digesting
enzyme.
Bile Juice (Liver) Water Waste materias excreted
with faeces or absorbed
and re-excreted later.
Bile pigment, Bile salt Alkaline therefore,
neutralize acidity of
chyme and stop action
of pepsin but allow
action of intestinal
enzymesemulsify fat.
Pancreatic juices Water, Alkaline salt, Help to increase
(Pancreas) Pancreatic lipase, Pancreatic alakalinity in intestine
amylase, Trypsin, and combined with fatty

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Chymotrypsincarboxypoly acid to form soap splits

peptidase fats into fatty acids and
glycerol. Splits certain
proteins, proteases and
peptone into shorter
polypeptide chains and
liberate some amino
acid.
Interstinal Juice Water Mucus Protect intestinal
(Duodenal gland and mucosa. Activates
goblet cells) trypsinogen forming
trypsin , trypsin then
activate chymotrypsin.
Enterokinase Split amino acid
Peptidase Split maltose into
glucose
Carboxypeptidase Split sucrose into
glucose and fructose
Amino peptidase, Split lactose into
Dipeptidase,Maltase,Sucrase, glucose and galactose
Lactase

Absorption of carbohydrate in non-ruminants:

The final product of carbohydrate digestion in non-ruminants is glucose,

galactose and fructose. Their absorption is an active process utilizing a specific
carrier protein that trans locates the molecules across the brush border membrane
of small intestine. This is energy dependent process and also required Na+ and
K+ ions. The rate of absorption of monosaccharides is also different. Galactose
is fastest absorbed than glucose, fructose, mannose and slowest pentose sugar.

Factors affecting digestion of carbohydrates in nonruminants:

1. Particle size: If particle size is reduced, than digestibility will be increased
because of increase in surface area for digestion. Grinding broken down
the cell wall so that cell contents come in contact with digestive enzymes.
2. Form of starch: Soluble starch is more digestible than insoluble form i.e.
amylose is more digestible than amylopectin.
3. Processing: It improves the digestibility of starch by breaking down the
cell wall. Cooked starch is more digestible than uncooked.
4. Fibre content: If fibre content is increased more than a level, it reduces the
digestibility of carbohydrates.

PRINCIPLES OF ANIMAL NUTRITION 18

 School of Veterinary Medicine
Isabela State University- Echague
CPSG,DVM

5. Enzyme Inhibitors: Presence of enzyme inhibitors like saponin, tannins etc.

reduces the digestibility of starch.

Factors affecting fibre digestibility in ruminant:

Following factors affect the digestibility of crude fibre (cellulose + hemicellulose

+ lignin).

1. No. and type of microbes present in rumen: If number of microbe is more,

digestibility of crude fibre increase. If cellulolytic bacteria are there, cellulose
digestion is more.
2. Relative proportion of fibre component: If hemicellulose is more, digestibility
of crude fibre is more. Lignin proportion is inversely related with fibre
digestibility.
3. Protein content in diet: Increased protein level in diet stimulates microbial
growth and improves digestibility of crude fibre.
4. Fat content in diet: Increased fat content in diet gives a protective layer on feed
particles, which depress the fibre digestibility.
5. NFE: CF ratio: If NFE content is increased, then digestibility of crude fibre is
decreased. Because NFE represents the soluble carbohydrates in feed i.e. starch
which is a more available source of energy.
6. Supplementation of green forages: It stimulates digestion of crude fibre
because they supply vitamins and some nonspecific factors required for microbial
growth.
Carbohydrates metabolism:

The metabolic processes in the body are of two types. The degradation of complex
compounds to simpler materials is called catabolism. Whereas those metabolic
processes in which complex compounds are synthesised from simpler substances
are called anabolism. As a result of the various metabolic processes; energy is
made available for mechanical and chemical work. The end products of
carbohydrate digestion in the simple stomach animals are glucose, galactose and
fructose. Energy is produced when these are burnt to carbon dioxide and water.
The energy released during metabolic processes in the cell is stored in the form
of high-energy bonds particularly those found in adenosine triphosphate (ATP)
and creatinephosphate (CP).

Glucose metabolism:

The degradation/ synthesis of carbohydrates in the cells is done by a number of

enzymes, which are mostly specific. The major pathway whereby glucose is
metabolized to give energy is a two-stage process.

PRINCIPLES OF ANIMAL NUTRITION 19

 School of Veterinary Medicine
Isabela State University- Echague
CPSG,DVM

1. Glycolysis (Anaerobic cycle, Embden-Meyerh of Paranes pathways).

2. Tricarboxylic acid cycle (Aerobic cycle, kreb' s/ citric acid cycle).

1. Glycolysis. In this process glycogen, glucose, galactose and fructose are

broken down to pyruvic acid and lactic acid in the absence of molecular
oxygen.

The sequence of reactions:

o The ATP production in the glycolytic pathway: Two moles ATP are
used in the initial phosphorylation of steps 1 and 3 and fructose-1-6-
diphosphate so formed then break down to yield two moles of
glyceraldehyde-3 phosphate. Subsequently one mole of ATP is
produced directly at each step 6 and 9. Four moles of ATP produced
from one mole of glucose. Since two moles of ATP are used up, the
net production of ATP from ADP is two moles per mole of glucose.
o Glyceraldehyde-3 phosphate is converted to 1,3 - diphosphoglyceric
acid in the presence of glyceraldehyde-3- phosphate dehydrogenase
enzyme and reduced NAD+ is produced and it may be oxidised via
the oxidative phosphorylation pathways, with the production of
three moles of ATP per mole of reduced coenzyme. Under aerobic
conditions, therefore, glycolysis yields eight moles of ATP per mole
of glucose.

2. Tricarboxylic acid cycle: The next stage in the degradation of glucose is

conversion of the two pyruvic acid molecules into two molecules of
acetyl coenzyme A (Acetyl CoA) as following reaction.

2 Pyruvic acid + 2 Coenzyme A= 2 Acetyl Co-A + 2C02 + 4 H.

▪ From this reaction it can be seen that two carbon-dioxide molecules and
four hydrogen atoms are released, while the remainders of the pyruvic acid
molecules combines with coenzyme-A to form two molecules of acetyl Co-
A. In this conversion, no ATP is formed, but six molecules of ATP are
formed when the four hydrogen atoms are oxidised in oxidative
phosphorylation system. So citric acid cycle (Tricarboxylic acid cycle or
kreb's cycle) is a sequence of chemical reactions in which the acetyl portion
of acetyl coenzyme-A is degraded to carbon dioxide and hydrogen atoms.
Then the hydrogen atoms are subsequently oxidised, releasing still more
energy to form ATP.
▪ The TCA cycle involves four dehydrogenations, three of which are NAD+
linked and one is FAD linked, resulting in 11 moles of ATP being formed

PRINCIPLES OF ANIMAL NUTRITION 20

 School of Veterinary Medicine
Isabela State University- Echague
CPSG,DVM

from ADP. In addition one mole of ATP arises directly with the change of
succinyl coenzyme-A to succinic acid. The oxidation of each mole of
pyruvate thus yields 15 moles of ATP. The total ATP production from the
oxidation of one mole of glucose is given below:
o 1 mole of glucose to 2 moles of pyruvate produces 8 Moles ATP
o 2 moles of pyruvate to CO2 and water produces 30 Moles ATP
=Total ATP per mole of glucose 38

▪ Gluconeogenesis: It is ·the process of synthesis of glucose from the

sources other than carbohydrates. When the body stores of carbohydrates
decrease below normal, moderate quantities of glucose can be formed from
amino acid, glycerol portion of fat, and from propionic acid. This process
is known as gluconeogenesis. Approximately 60 percent of amino acids in
the body protein can be converted into carbohydrates.
▪ Glycogenesis and glycogenolysis: Glycogen synthesis from simple sugars
in the body tissues is known as glycogenesis. Glucose, galactose, fructose
and mannose are readily converted to glycogen by various stages in which
various enzyme systems are involved.
▪ Similarly the process of degradation of glycogen to glucose- 1- phosphate
in the cells is known as glycogenolysis.
▪ Lactose synthesis: Lactose is formed by condensation of one glucose and
one galactose molecule. It is formed by the action of the UDP - D -
galactose with glucose in the presence of the lactose synthetase.

UDP – Galactose + D - glucose Lactose synthetase UDP + Lactose

▪ Fat synthesis from glucose: When the carbohydrate intake exceeds the
requirement of the body for energy purposes, sugar is transformend into
fat. It involves the synthesis of two components, fatty acid and glycerol,
which combine with each other to give fat.

PRINCIPLES OF ANIMAL NUTRITION 21