ADVANCED RUMINANT NUTRITION: COLTECH APAP6204 (40H:25/5/10)

COURSE OUTLINE

Chapter 1: Particularity of the digestive tract of ruminants

1.1 The rumen
1.2 The reticulum
1.3 The omasum
1.4 The abomasum

Chapter 2: Digestion in ruminants

2.1 Digestion processes in ruminants
2.2 Digestion of carbohydrates
2.3 Digestion of proteins and nitrogenous compounds
2.4 Digestion of lipids
2.5 Absorption and metabolism of digestion products

Chapter 3: Factors that influence the efficiency of the rumen

3.1 Temperature
3.2 Oxygen concentration
3.3 Moisture content
3.4 Fibre concentration
3.5 Diet stability
3.6 Energy and nitrogen ratio
3.7 Composition of feed
3.8 Modification of diet composition

Chapter 4: Feed composition and digestibility

4.1 Feed Composition
4.2 Digestibility of feed
4.3 Measures of digestibility in animals

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4.4 Use of markers in animal nutrition

Chapter 5: Energy systems in nutrition

5.1 Gross Energy (GE)

5.2 Digestible Energy (DE)

5.3 Metabolisable Energy (ME)
5.4 Faecal Energy
5.5 Heat increment
5.6 Net Energy
5.7 Total Digestible Nutrients
5.8 Physiological Fuel Values (PFV)

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 CHAPTER ONE: Particularity of the digestive tract of ruminants

Athough they are called polygastrics, ruminants have only one but complex stomach. However, this
stomach is made up of 4 compartments:

- The rumen or paunch

- The reticulum or honeycomb
- The omasum or manyplies
- The abomasum or true stomach

The first 3 compartments are also called the “pre-stomachs”. The true stomach is the abomasum and is the
only compartment which contains gastric glands.

1.1 The rumen or paunch

It is located at the left side of the animal
It is the largest compartment makes up about 90% of the total stomach volume (250-300L) or 70-75%
of the digestive tract volume.
It serves as the storage and fermentation vat for feed.
It has 2 openings: an entry orifice which is narrow but very elastic, attached to the oesophagus. It is
called the cardia. There is also an exit orifice which is very large; located between the rumen and the
reticulum. These orifices are linked by a flap which when upon contraction, feed (such as milk) leaves
the oesophagus directly to the reticulum, by-passing the rumen. The rumen is loaded with billions of
micro-organisms which ferment the feed. Most of the volatile fatty acids produced during fermentation
are absorbed across the walls of the rumen. The walls of the rumen strong muscles which contract
rhythmically to ensure adequate grinding of the feed. The walls are also supplied with rich ramification
of blood vessels for sufficient absorption of the products of digestion. Numerous tiny projections called
“papillae” are present which increase the surface area and absorption capacity of the rumen.
1.2 The reticulum or honeycomb
Is before the rumen, in the forward area of the body cavity.
Its wall has a pouch-like structure with tissues arranged in a network resembling a honey comb.
A small fold of tissue lies between the reticulum and the rumen, but these 2 are not actually separate
compartments. Collectively, they are called the “rumeno-reticulum”. Foreign materials that are ingested
accidentally (such as metallic objects) remain in this compartment. If such materials penetrate deeper in
the tissues (walls) of the reticulum, they can reach the envelopes of the heart and cause death of the
animal.
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1.3 The omasum or manyplies
Is more voluminous than the reticulum.
It has a globe-shaped structure and contains many leaves of tissue arranged in a parallel fashion (like
the pages in a book) and feed passes through theses leaves. This compartment serves as a filter,
permitting only feed materials of certain dimensions to pass from one leaf to the other.
It absorbs water and other substances resulting from digestion/fermentation processes. Feed materials
between the leaves will be drier than materials from the other compartments.
1.4 The abomasum or true stomach
It is the only compartment with a glandular lining.
It contains digestive glands which secrete gastric juices (containing enzymes) and rennin (in the case of
calves).
Hydrochloric acid is also secreted into the abomasum from the walls.
A lot of water and minerals are absorbed from here into the bloodstream.
It is comparable to the stomach on non ruminants.

NB. Among these four compartments, only the abomasum is well developed in calves and lambs (milk and
water pass directly from the oesophagus to the abomasum, by-passing the rumen).

The small intestines measures about 20 times the length of the animal. It comprises the duodenum, jejunum
and ileum. It receives the secretions of the pancreas and gall bladder which aid in digestion. Most of the
digestive processes are completed here and many nutrients are absorbed through the many villi (small
finger-like projections) into the blood and lymphatic systems.

The caecum is located at the junction of the small and large intestines which some previously undigested
fibre may be broken down.

The large intestine is the last segment of the tract through which undigested feedstuff pass; some bacterial
digestion of undigested feed occur here. However, water absorption is the primary digestive activity that
occurs in the large intestine.

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 Chapter 2: Digestion in ruminants

2.1 Digestion processes in ruminants

Digestion processes ensure that complex insoluble compounds present in feeds are
transformed into simple soluble nutrients that can be absorbed across the intestinal walls
into the body. This process of simplification or degradation involves various processes
that complement each other and can be grouped into 3 major actions:

Mechanical: chewing, softening and grinding

Biological: microbial fermentations

Chemical: action of digestive enzymes

2.1.1 Mechanical degradation of feeds

Chewing: of the feed is accomplished in stages: the first stage of chewing is fast (70 to
90 movements per minute for large ruminants and 125 to 150 movements per minute for
small ruminants). There is very little grinding of the feed, which is stacked up in the
rumen together with water and saliva. The second stage of chewing is characterised by
the animal bringing back the feed from the rumen (and reticulum) into its mouth for
further chewing and softening before it is sent again to the rumen for fermentation. This
is called rumination (or chewing of the cord). Rumination has two main advantages: it
facilitates microbial fermentation and digestion of all the components of the feed by
breaking down cell walls/membranes thereby exposing the cell contents for enzyme
action. Secondly, it provokes excessive production of saliva.

Softening of the feed: this is ensured thanks to the production of excess amounts of
saliva. Saliva production is highly developed and is on a continuous basis in ruminants
(large adult cows can produce 100 to 200 litres per day; 10L or more in sheep). Saliva is
a buffer solution (pH is 8.2). In addition to water, it contains urea, mucus (contains
proteins), bicarbonates sodium and potassium phosphates (all of which are used by the
rumen micro-organisms) and salt bases which neutralise the acids produced during
microbial fermentations.

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Prolonged grinding: All fractions of the feed (liquid or solid) enter into the rumen after
chewing. Certain feeds can undergo grinding several times before they are sent to the
reticulum. The duration of stay in the rumen varies depending on the nature of the feed;
it can be as short as 1.5 hours for young plants to as long as 5 days for straw. An
exception to this is in the case of new-born ruminants where the liquid food (milk and
water) provokes a reflex causing the oesophagus to close up such that the food goes
directly to the omasum and then to the abomasum (thus by-passing the rumen). After 4
weeks of age, this reflex is no longer provoked by water but by milk only (pure or
diluted).

All the compartment of the stomach participate in grinding of the feed. In the rumen, the
feed floats first on top of the liquid, then gets immersed in it for softening and
homogenisation. The prolonged stay here (30 to 70 hours) permits destruction of the cell
walls and liberation of the cell contents. Grinding is ensured by contractions of the
muscular walls of the rumen and to a lesser extent by the cilia of protozoa and the release
of fermentation gases. Contractions also take place in the reticulum and abomasum. The
omasum distends when the rumen is contracting thus permitting the feed to descend
through its filter made of many small leaf-like blades.

2.1.2 Biological degradation of feeds

Ruminants have developed a unique digestive system which permits them to maximize
the digestion of cellulose, a complex carbohydrate predominant in grasses and which is
completely indigestible by most other animals including man. The success of ruminants
in efficiently digesting cellulose in grass is linked to the functioning of its rumen and the
billions of micro-organisms that in there. It is not actually the ruminant that digests the
cellulose but it is the micro-organisms of the rumen that do the digestion (through
fermentation reactions) primarily for their own nourishment and for increase in their
population (growth). Microbial growth in the rumen is very fast; sometimes the
population can double in 20 minutes (generation time). At this growth rate, the microbial
population will become too great for the rumen to contain. Consequently, some of the
micro-organisms move from the rumen to other parts of the digestive tract: they move
first to the omasum, then to the abomasum where they are destroyed due to the high
acidity of the environment. Digestion of the micro-organisms begin here thanks to the

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presence of gastric enzymes. They transit into the small intestines where digestion
continues until the products are sufficiently small to be absorbed through the intestinal
walls and into the blood. Thus the ruminant is nourished by absorbing the nutrients that
result from the digestion (degradation) of micro-organisms of the rumen. Therefore the
ruminant can be regarded as a farmer: it grows micro-organisms in its rumen, provides
grass for the micro-organisms to feed on, and then consumes some of these micro-
organisms daily.

The environment of the rumen is very stable (physically and chemically), which is most
favourable for the survival of the micro-organisms (bacteria, protozoa, fungi). It has the
following ideal characteristics:

Warm temperature: 39 – 40 oC

Anaerobic environment: contains 60-70% CO2, 25-35% methane

pH: relatively constant between 6 and 7. Acids produced during fermentation of fibre are
neutralised by salivary salts, NH3

Supply of finely ground forage (feed) with increased surface area for interactions

Continuous grinding by contractions of the rumen walls.

The micro-organisms of the rumen

Three main groups are present: fungi, protozoa and bacteria. The first group of micro-
organisms to begin degradation of feed is fungi. Their zoospores attach themselves to
feed particles, begin to germinate and produce mycelium which is capable of infiltration
the fibrous structure of forages. They secrete enzymes which degrade fibres. Attack by
fungi reduces the resistance of feed particles and enables bacteria and protozoa to
continue the degradation process.

A large population of ciliated protozoa are also present in the rumen (100 000 to 1
million/cc of ruminal fluid), sometimes making up to 50% of the rumen biomass. Their
cilia contribute in mixing of the feed with ruminal fluid. They feed on bacteria
(phagocytosis), on carbohydrates and feed proteins.

Bacteria are the largest group of microbes in the rumen (1 to 10 billion/cc of ruminal
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fluid). Based on their main functions, they are classified as amylolytic (secrete enzymes
that degrade soluble carbohydrates and starch), cellulolytic (secrete enzymes that degrade
cellulose), and bacteria which synthesize vitamins of the B family.

Example of common types of micro-organisms that carryout fermentation (digestion) in

the rumen include:

Cellulose decomposers: Bacteroides succinogenes

Butyrivibro fibrisolvens

Rumenococcus albus

Starch decomposers: Bacteroides amylophilus,

Streptococcus bovis

Fatty acids decomposers: Selenomonas ruminantium

Megasphaera elsdenii

Methane producer: Methanobacterium ruminantium

Characteristics of microbial digestion

Digestion by micro-organisms has many important characteristics:

Enables grass to be used as food (good exploitation of cellulose)

The chemical and enzymatic digestion mechanisms that take place in man and in most
animals are incapable of digesting cellulose. Cellulose consequently transits through the
digestive tract without any degradation and no nutrient is obtained therefrom. Contrarily,
microbial processes permits the digestion of cellulose to produce energy and other
nutrients (such as proteins) from grass, thus ensuring the survival of the ruminant. This is
very important from the ecological point of view because cellulose is by far the most
abundant of the earth’s biomass. On one hand, an animal that is capable of digesting
cellulose can satisfy its nutritional needs almost anywhere. On the other hand, the source
of cellulose is much more abundant on earth than the other food sources, placing

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herbivores at the base of the food pyramid.

Among mammals the digestive tract in ruminants is the most developed with respect to
microbial digestion. However, not only ruminants carry out microbial digestion. All
herbivores use the same mechanism for the degradation of cellulose to VFAs by
microbial action. The difference between ruminants and the other herbivores is location
of the micro-organisms: for the horse, the micro-organism are located in the colon while
for rabbits and pigs, they are located in the caecum.

The level of exploitation of cellulose varies significantly between the different animal
species as illustrated below:

Digestion of other feed components by micro-organisms

Cellulose is not the only component present in grass that is digested by micro-organisms.
In addition to cellulose, carbohydrates (soluble sugars, starch, etc) and proteins (abundant
in leguminous plants and in many grains) are also present. Thus the major constituents of
feeds undergo microbial digestion, are used by the micro-organisms for their growth
(some are used by the host animal) and the by-products (VFAs) are recuperated by the
animal (for energy production).The only feed components that are not degraded by
micro-organisms are micro-organisms are lipids (which have a negative influence on the
functioning of the rumen).

Microbial digestion, a security factor

A balanced ration must supply not only sufficient quantities of carbohydrates, proteins
and lipids, but the quality of these nutrients must also be adequate so that sufficient
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quantities of essential elements (essential amino acids, essential fatty acids, vitamins) are
also available to the animal. To satify their nutrient needs, ruminants make use of the
products resulting from microbial digestion in the rumen. These micro-organism have
unique potentials that are absent in mammals: they are capable of synthesizing essential
amino acids, essential fatty acids and certain vitamins. Thus ruminants are less
susceptible to nutrient deficiencies than other farm animals because a good proportion of
their nutrient needs is provided thanks to microbial activities. However, ruminants can
still suffer from nutrient deficiencies; micro-organisms cannot supply oligo-elements, for
example.

Poor yield, but still very useful

One significant disadvantage of microbial digestion is that the transformation yield is

comparatively very low: 4 calories from plants will produce 1 calorie of chicken meat but
about 10 to 20 plant calories are needed to produce 1 calorie of beef or mutton.
Production of micro-organisms from feed consumed and the transformation of these
micro-organisms to useful nutrients is accompanied by a big loss in energy in the form of
heat and gas eructed during rumination. An average size cow can eruct about 600L of gas
per day and produce heat equal to that of a 1000W radiator. However, this loss cannot be
compared to the benefits obtained when ruminants are given grass which otherwise will
be useless to man and other animals; and on land which at times cannot be used for
farming. Furthermore, in comparison to poultry or pigs, a ruminant will produce less
meat when fed on cereals.

2.1.3 Chemical degradation of feeds

In addition to mechanical and biological degradation, the feed is also subjected to a series
of actions by enzymes secreted by the digestive glands of the animal and by enzymes
secreted by the micro-organisms in the rumen. The prevailing pH gives rise to 2 types of
environment for degradation to occur: gastric and intestinal environments.

Gastric digestion takes place in an acid environment. In addition to producing HCl

(which acidifies the environment), the gastric glands in the abomasum also secrete pepsin
(degrades proteins to polypeptides), rennin (in young calves which coagulates the casein
in milk) and cathepsin (liquefies the coagulum before it is further degraded by pepsin).

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Intestinal digestion which takes place in an alkaline environment is more complete. Two
sources of enzymes are responsible: (i) enzymes in the pancreatic juice (secreted by the
pancreas) include amylase and maltase which break down starch and maltose and
maltose to glucose, lipase which breaks down lipids to triglycerides, and trypsin which
breaks down proteins to polypeptides and amino acids; (ii) enzymes in the intestinal juice
(secreted by the intestinal glands) include saccharase which break down saccharose and
lactose to glucose and galactase, amylase and maltase, lipase and erepsine which breaks
down polypeptides to amino acids.

In calves, digestive juices are secreted only after 24 to 65 hours following birth. Certain
proteins (such as globulines, antibodies in the colostrum) can be absorbed directly
without digestion, but only for the first 48 hours after birth. Digestion of starch will take
place only after 5 weeks of age. Therefore, flour should not be mixed with milk and
given to the calf before this age.

2.2 Digestion of carbohydrates

The digestion of carbohydrates in polygastric animals is different from digestion in

monogastric animals. In the rumen of polygastric animals, carbohydrates (starch, sugars,
etc.) are broken down very rapidly (by microbial enzymes: amylases) to simple sugars
(mainly glucose). The glucose is either fermented to produce small short chain fatty acids
volatile fatty acids (VFA), energy (ATP) immediately used by the microbes and heat or it
is sent to the small intestines where it is absorbed into the blood.

Cellulose digestion:

Cellulases secreted by the cellulolytic bacteria in the rumen break down complex
carbohydrates (fiber) into glucose and pentose, which can now be fermented. This results
in:

A) Production of energy (ATP) for immediate microbial use.

B) Production of gases: CO2 (60 -70%) and methane (25 -35%). Both gasses are sent out
by eructation although some CO2 is excreted through the lungs. These gases represent
about 8% loss of energy from the ration.
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C) Production of volatile fatty acids (VFAs): These are the main source for the ruminant.
Glucose is oxidized to pyruvic acid accompanied by liberation of energy of energy for
use by the microbes and production of four hydrogen atoms. Under anaerobic condition
of the rumen, pyruvic acid undergoes different oxidation processes each of which results
in the synthesis of 3 different VFAs. The main VFAs produced in the rumen include
acetic acid (has 2 carbon atoms), propionic acid (has 3 atoms) and butyric acid (has 4
atoms).

2 pyruvate + H2O 2 acetic acids + 2CO2 + 2H2O + energy

2 pyruvate + 8H 2 propionic acid + 2H2O + energy

2 pyruvate + 4H 2 butyric acid + 2CO2 + 2H2 + energy

The proportions of these VFAs produced depend on the rumen’s environment. Generally,
when the animal’s ration is of good quality, more propionic acid will be synthesized in
proportion to the same quantity of acetic acid that will be reduced. The quality of VFAs
produced can be improved either by increasing the quantity of easily fermentable feeds
(such as cereals) or by chopping the forage into very small pieces. With a normal ration
and a pH of 6.5, the VFAs produced will be of the following proportions:

Acetic acid: 70%; Propionic acid: 15 – 20%; Butyric acid: 10 – 15%; Other VFAs: 2 –
5%

These VFAs are recycled by the ruminant: they are absorbed through the rumen wall,
transported by the blood to the liver. In the liver, the VFAs are later converted into acetyl
coenzyme A before they are used in the production of energy. Here again the ruminant is
gifted not only in utilizing the micro-organisms in its rumen as a source of nutrients but it
also recycles by-products from microbial metabolism and converts them into energy for
its own metabolism. Residual VFAs that pass unabsorbed (in the rumen) to the large
intestine is also absorbed into the bloodstream at this level.

Variations in the proportions of VFAs synthesized is determined by the rate of

fermentation and the pH of the rumen; the pH is influenced by the feed composition:
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If the main ingredient in the ration is forage, more acetic acid will be produced while
propionic and butyric acids will decrease;

If the ration is rich in concentrates, the starch present in the ration will be rapidly
fermented lowering the pH (this may lead to accumulation of lactic acid causing a health
problem = acidiosis). The low pH will reduce the synthesis of acetic acid while propionic
and butyric acids will increase (whose synthesis will again decrease if the pH is too low).

If the ration is rich in soluble carbohydrates (such as sugars), this will favor the synthesis
of butyric acid while acetic acid will decrease. Butyric acid synthesis promotes the
synthesis of milk fat; consequently, milk production increases.

The presence and the digestion of fibers in the rumen is very important in the ration of
the animals for two main reasons;

Fibers stimulate rumination and facilitate salivation.

Fibers are necessary for the development of a solid layer inside the rumen. The layer is
necessary because it permits regular contraction of the rumen which is important for the
proper mixing of the content inside the rumen.

2.3 Digestion of proteins and nitrogenous compounds

In the rumen, all of the nitrogenous compounds and most (not all) of the proteins
consumed are broken down by microbial enzymes into ammonia. This is called
“microbial proteolysis”. The ammonia produced combines with the carbon chains and the
energy produced as a result of carbohydrate fermentation to form microbial proteins (if
the carbohydrate/nitrogenous matter ratio is adequate). This is called “microbial
proteosynthesis”. Part of the ammonia produced is recycled to the saliva. Another part is
transformed into urea by the liver and eliminated from the body in the form of urine. The
proteins (microbial and feed) pass into the abomasum where they are later broken down
to polypeptides (by gastric enzymes: rennin, pepsin, etc.). Digestion the abomasum is
similar to that which takes place in the stomach of monogastric animals. Then the
degraded material is sent to the small intestines where they are degraded to amino acids
by enzymes secreted by the pancreas, cells of the intestinal walls. If there is any excess of
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ammonia, it passes into the blood to the liver where it is converted into urea (and is
finally excreted out of the body in the form of urine).

The portion of protein consumed that is not degraded in the rumen (relatively smaller
than that degraded into ammonia; it is called “by-pass protein”) passes to the abomasum
where it is broken down to polypeptides then it moves to the small intestines where it is
degraded into amino acids (feed proteins). The amount of by-pass protein included in a
ruminant’s diet is very important especially for animals that are highly productive (eg.
Milk production) because such proteins are very rapidly digestible and will readily
supply the amino acids that are required for rapid milk synthesis.

These amino acids (produced from microbial or feed proteins) are used either for the
synthesis of new proteins by the microorganisms present or are broken to give carbon
dioxide and volatile fatty acids (for energy production when necessary). The
microorganisms in the rumen use the amino acids derived from feed for their growth and
multiplication. They are also capable of synthesizing all the necessary amino acids
(essential and non essential) from the non protein nitrogenous substances present in the
feed. The rate of protein synthesis by the micro-organisms in the rumen depends on their
rate of metabolism of feed ingredients.

In addition, the microorganism synthesizes a wide variety of vitamins that are necessary
for ensuring metabolic processes of the body. Another very important contribution by the
microorganism in the rumen is the secretion of numerous types of enzymes (which are
proteins) that are capable of digesting the fibers consumed in the feed. This is the
fundamental difference in digestion between polygastric and monogastric animals.

Finally these amino acids are used by the animal to satisfy its needs for growth,
maintenance, and/or production.

Therefore, there is a symbiotic relationship that exists between the animals and the
microorganisms in the rumen both benefiting from each other.

Benefits of microorganisms in the symbiotic relationship:

-Gain protection from environmental factors by the host animal

-Ready source of already digested feed nutrients

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-Ideal and constant temperature, pH, etc.

Benefits of animal in the symbiotic relationship:

-Micro-organisms the animal help to digest feed especially cellulose (fiber)

-Source of proteins for the animal (especially when the microbes die)

-Micro-organisms synthesize nutrients not present in animal’s feed eg. Vitamins of B-

complex family.

NB: The micro-organisms destroyed in the digestive tract are very rich in proteins (a
microbial cell is about 80% protein). Therefore, they are also an important source of
amino acids for the ruminant for the synthesis of its own proteins.

2.4 Digestion of lipids

Compared to the other nutrients, lipids are usually present in much smaller quantities in
the ruminant rations. In the rumen, the microbes will breakdown lipids to glycerol, sugars
(easily fermented) and fatty acids. The fatty acids are either saturated (if the lipid is of
animal origin) or unsaturated (if the lipid is of plant origin). The most common type of
fatty acids present are those of C18: linolenic acid in forages and linoleic acid in grains
(unsaturated fatty acids). Lipid fermentation in the rumen (anaerobic milieu) always
results in the production of hydrogen. Some of this hydrogen is evacuated in the form of
methane while the other fraction interacts with the unsaturated fatty acids. The microbes
produce enzymes which are capable of attaching the hydrogen atoms to the double bonds
of the unsaturated fatty acids, thus converting them into saturated fatty acids (this is
called hydrogenation) which are absorbed into the lymphatic system at the level of the
small intestines. Consequently, the proportion of saturated fatty acids in the animal’s
body is much higher than that which was present in its feed.

Thus, one major difference between lipid digestion in ruminants to that in monogratrics
is that the microbes in the rumen are capable of digesting or modifying the lipids in
feeds. In addition, the microbes can also synthesize lipids from simple feed constituents.
The animal uses some of the lipids as a source of energy, for synthesis of body tissues
and/or for milk synthesis. Hydrogenation explains why most of the fatty acids in
ruminant tissues and in the milk are mostly saturated.
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Although lipids are a more concentrated source of energy than the other feed
constituents, large quantities of fatty acids in the rumen will reduce the ability of the
microbes to breakdown the other feed constituents. It is generally recommended that the
proportion of lipids should not exceed 5% in the ruminant’s ration.

2.5 Absorption and metabolism of digestion products

The end products of digestion of feed and microbial cells include:

a) Nutrients: water, vitamins, mineral salts

Simple sugars: glucose, fructose, galactose
Fatty acids (volatile or not), glycerol
Amino acids
b) Undigested materials found in the faeces.

Absorption of nutrients into the body occurs at different mucosal sites:

i) Mucous of the rumen: VFAs (acetic, propionic and butyric) resulting from the
degradation of fibre is absorbed through the mucous of the rumen. Excess ammonia is
also absorbed at this site.

ii) Mucous of the omasum: absorption of lots of water and mineral salts takes place here.
The mucous of the omasum and the abomasum also absorb residual VFAs which were
not absorbed at the level of the rumen (10 – 30%).

iii) Mucous of the small intestine: absorption of all nutrients (glucose, glycerol, fatty
acids, amino acids, minerals, vitamins, water) takes place here (although it may be
residual for certain nutrients).

iv) Mucous of the large intestine: mostly water and mineral salts are absorbed here.

Metabolism of nutrients

Metabolism is all the transformation of nutrients that take place at the level of cells
including all associated energy exchanges that take place. Metabolism comprises 2
inversely related series of reactions:

Anabolism: is the synthesis of new living matter or storage materials. Anabolism requires
large amounts of energy.
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 Catabolism is the degradation of substances (storage materials or cellular constituents),
accompanied by liberation of energy.

VFAs are the main source of energy for ruminants. Thus the blood sugar concentration is
lower for ruminants than for monogastrics whereas the VFAs concentration is much
higher.

VFAs taken to the liver are transformed to glycogen and glucose which will be used as
follows:

- Oxidised by the cells for energy production in the muscles

- Utilised for the synthesis of fatty acids (to synthesise lipids) and as fat for milk
synthesis
- Utilised for the synthesis of carbohydrates in milk.

Amino acids are used for the synthesis of a wide variety of proteins destined for growth,
maintenance/repair, regulation of metabolic processes.

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 Chapter 3: Factors that influence the efficiency of the rumen

Since nourishment of the ruminant depends on the products of microbial degradation, it implies that the
animal can be well nourished only if the rumen is functioning properly. Many factors (both intrinsic and
extrinsic) influence the functioning of the rumen. Some intrinsic factors that must be satisfied in order for
the microbial population to grow rapidly include:

3.1 Temperature: the rumen functions better at a warm and constant temperature. This is provided for by
the animal which has a constant body temperature of about 38.5 oC (for a cow).

3.2 Oxygen concentration: Microbial fermentation in the rumen is more efficient under anaerobic
conditions. The quantity of oxygen that penetrates into the rumen when the animal consumes feed is very
small, thus the rumen environment is almost anaerobic.

3.3 Moisture content: The rumen functions better when the environment has a high moisture content. A
cow drinks large quantities (about 50L) of water daily. In addition, about 200L of saliva is produced by the
salivary glands, some of which remains in the rumen. These sources of water render the rumen very moist;
about 80% of the rumen content is water.

Other conditions that influence the efficiency of the rumen are extrinsic; therefore the farmer can play a
major role in influencing rumen functioning. These include:

3.4 Fibre concentration

The muscular walls of the rumen must regularly contract for the efficient mixing and grinding of the
contents therein (the contractions respect a 3 to 5 minutes cycle). The contractions are provoked by
irritation of the rumen walls by the plant fibres (grass) present in the diet. The nature of the fibre also
influences the frequency and intensity of the contractions (chopped forage is better). Consequently, a
ruminant’s ration must contain a sufficient quantity of forage or straw to supply fibre which will excite the
walls of the rumen, thus assuring an efficient mixing of the rumen contents.

3.5 Diet stability

The rumen’s microbial flora is made of a many different species of micro-organisms. Different species
digest different components of the ration given the animal. When the composition of the ration is modified,
two to three weeks are required for the microbial flora to regain a new equilibrium such that the feed can be
efficiently utilised. Therefore, it is best to maintain the animal’s ration for as long as possible.

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3.6 Energy and nitrogen ratio

For growth, the microbial population needs sources of energy and nitrogen so that they can synthesize the
necessary proteins. However, they need enough energy so that they can be able utilise all the available
nitrogenous matter (as illustrated below); if not the nitrogen will be eliminated in the form of ammonia.

Therefore, the farmer must balance the energy and nitrogen sources (in the diet) in order to avoid wastage.
To maximize utilisation, these two must be supplied simultaneously in the feed.

3.7 Composition of feed

The micro-organisms in the rumen cannot store any available energy or nitrogen: they must be utitlised as
soon as they are produced. If the amount of energy produced is insufficient, it becomes a limiting factor for
the production of microbial proteins (due to wastage of available nitrogen).Therefore it is important to
know the rate at which energy is liberated from the feed give to the animal as well as the quantity of
nitrogen the feed contains. Generally, the proteins in feeds are degraded very rapidly by the micro-
organisms in the rumen. Therefore all the nitrogen they contain is made available (in the form of ammonia)
for microbial use within an hour. However, the rate of liberation of energy varies:

- Feeds rich in soluble sugars, starch and seed cakes liberate their energy with one to hours. They are
called fermentable feeds because they are rapidly fermented by micro-organisms.
- Feeds rich in cellulose must stay in the rumen for 24 to 48 hours to liberate all their energy. They
are called poorly fermentable feeds.

3.8 Modification of diet composition

The micro flora of the rumen comprises diverse species of micro-organisms which have established a
stable yet complex ecosystem, in relation to the diet that the animal is receiving. Any abrupt or rapid
changes in the diet will result in an imbalance of the microbial species present and an upset of this
ecosystem. This will result in a dysfunction the digestive tract. Therefore, it is advisable to allocate a
reasonable transition time during which there is a gradual modification the diet of the animal; and to do
so only when it is absolutely necessary. For example, if the quantity of concentrate must be increased,
do not exceed an additional 250g/day (or 2kg/week).

19
 CHAPTER 4: Feed composition and digestibility

The feed represents the greatest cost of production in cattle rearing, accounting for more than 50% of
the total production left.

Ruminants in general have the particularity that they are capable of consuming forage that is of no
nutritive value to man and to monogastric animals. And they convert the forage to animal products that are
of high biological value to man. It will make little difference if the animal is of very high performing breed
and is not given the proper type and quantity of nutrients for beef synthesis. The animal will be an
inefficient producer.

A good feeding program complements good breeding and therefore permits the animal to produce
closely to its maximum genetic potential. The nutrients that the animals require are destined for production
activities that include maintenance and repair, growth, production of energy, development of the foetus in
the case of the gestating cows and development of milk production in the case of lactating cows. Forage
serves as the foundation for formulating ration for cattle because they are the cheapest ingredient in the
ration. Therefore, to increase profitability, the animal should be given as much forage as it can consume.

Different types of forage are very significant in their nutritive value, digestibility, palatability, and
physical structure. Digestibility reduces as the forage matures and consequently the nutritive value
decreases. Therefore for efficient utilization, the optimum time of harvest must be established for each type
of forage.

20
 4.1 Feed Composition

The composition of feed is mainly water and dry matter. The nutrients that are in dry matter can be summarized as follows;

Nitrogenous True Proteins

Compounds

Non Proteins
Organic
Matter Lipids (fats and oil)
N2 free
compounds
Dry Matter Non
Carbohydrates
structural;
sugars,
Water solubles starch
Vitamins solublSoluble

Feed Fat Solubles

Inorganic Matter Micro minerals:

Structural
(Fe, I2, Zn, Mn, F, Co, etc) carbohydrates
Water
Macro minerals:

(Na, Mg, Ca, P, K, Cl)

21
 a) Water

It is an essential nutrient like any other nutrient. It is the most crucial nutrient for
survival. Many tissues of the animal will not be developed properly if the water supply is
inadequate. Growth will be significantly reduced because about 80% of muscle tissue is water
and 20% of adipose tissue is water. Water also plays a very important role during digestion,
assimilation and elimination of waste materials from the body. The ruminants need large
quantities of water to facilitate chewing and swallowing of feed. This water is provided in the
form of saliva. Extra quantity of water is added during gestation for foetus development and
during lactation for milk synthesis. The animals also need water to regulate its body temperature.
Some factors that influence the quantity of water the animal needs are;

 Physical effort
 Physiological state (gestation, lactation)

Moisture content in dry materials, environmental temperature/climate.

In tropical countries, the average quantity of water requirement for a cow (400kg) is
about 50l/day. Insufficient water supply is automatically manifested by the decrease in growth
and decrease in milk production.

Good quality water should be available to the animal at all times.

a) Proteins

Proteins are necessary for growth, maintenance, reproduction, lactation, synthesis of

essential body compounds, regulation of water balance, synthesis of antibodies. The quality of
the protein in the feed is important only for the young cows. This is because the microorganisms
in the rumen of the mature animal are capable of synthesizing all the amino acids needed by the
animals even from non-protein substances present.

 Therefore it is recommended to give to the non-adult animals non-protein nitrogenous

compounds because they are cheaper than proteins. A typical example is urea which is
not a protein but it contains nitrogen which can be converted into proteins by

22
 microorganism in the rumen. Under extreme conditions when there is no other source of
energy, proteins willbe de-aminated for the production of energy.

Insufficient supply of protein in the diet will result in;

 Slow growth, low fertility

 Abortions, pre-mature and still births
 Reduce the quantity of milk produced
 Loss of weight
 Increase of disease risks (low resistance to infections)
 Etc.

b) Carbohydrates

They include sugars and starches which are non-structural and structural carbohydrates
also called crude fibre which includes cellulose, hemi-cellulose, pectin, gums, and lignin. The
sole function of carbohydrates is for energy production.

The peculiarity of ruminants is the ability to digest significant proportion of the fibre.
However, fiber digestibility decreases with increase percentage of lignin. Even if the animal is
reared under intensive production system where there is a good proportion of diet, it is
recommended that fiber should constitute at least 15 % of the diet. This is necessary to maintain
the proper functioning of the rumen.

c) Fat (Lipids)

- Lipids comprise fats and oil and they are a concentrated source of energy (main
function). Lipids also contain fat soluble vitamins (lipid soluble vitamins) which are
useful in carrying out the execution of many reactions. They help protect vital organs
(like the heart, kidneys, etc.) from physical shock, they are integral constituents of cell
membranes and some hormones (ex hormones), it is a body regulator (helping to regulate
the uptake and excretion of nutrients by the cells), it is a thermal insulator (helping to
maintain a constant body temperature), it serves as a precursor of prostaglandins

23
 (hormone-like substances capable of simulating muscle contractions). However, it is
recommended that the liquids contained should not exceed 5% because lipids influence
(change) the microflora in the rumen thus negatively affects digestibility. Common
ingredients that are high in lipids include seed cake.

d) Minerals
Minerals are present in the inorganic portion of the dry matter of the feed and it is called
ash. Minerals are essential because they carry out a wide variety of functions in the animals.
Some functions and diseases associated with minerals include;
 Calcium: bone development/maintenance, milk synthesis, development of fetal
skeleton, control of neuro-muscular signals, permeability of cell membranes,
ensures acid/base equilibrium of the blood, promotes blood coagulation (by
catalysing the transformation of prothrombine to thrombine).
 Phosphorus: bone development/maintenance, milk synthesis, development of fetal
skeleton, intervenes in many biochemical reactions: production of energy form
carbohydrates and lipids, metabolism of carbohydrates, acid/base equilibrium,
buffering of the rumen fluid.
Problems associated with insufficient availability of Ca and P in the animal include:
+ diseases related to the skeleton: rickets in young animals (bone deformation, growth
retardation, bent swollen knees, stiff legs, etc), osteomalacia in adult animals
especially dairy cows (soft, fragile easy to fracture bones) due to excessive
mobilisation of the minerals from the bones for milk synthesis and fetal development,
osteofibrosis in adults (sot enlarge bones whose cavities are filled with fibre).
+ diseases related to milk production: hypocalcemia (milk fever) due to excessive
mobilisation of Ca from the bones into milk; a situation where the parathyroid
hormone is required in more quantities than the gland can secrete. hormone :
characterised by reduced body temperature and a reduction of the vital functions of
the body following the peak milk production phase.
+ diseases related to reproduction: low fertility, death of the embryo/fetus.
However, excess P with respect to Ca can cause increased secretion of parathormone
(resulting in osteoporosis), diseases related to reproduction and perinatal death (in the
case of dairy cows), diseases associated to the kidneys (in young lambs).

24
 Sodium: along with potassium and chlorine plays an important role in maintaining
the acid-base equilibrium and osmotic pressure as well as in promoting the
absorption of glucose. Consequences of insufficient Na supply include reduced
growth and reduced utilisation of digested nutrients (amino acids and energy),
 Potassium: also pays an important role in maintaining the acid-base equilibrium
and osmotic pressure, in nerve irritability and carbohydrate metabolism. It is
abundant in both plant and animal feeds.
 Chlorine: found in extra cellular fluid, blood, tissues and urine. Is essential for the
regulation of osmotic pressure, is the main anion of the gastric juice and plays an
important role in the transport of CO2 in the blood.
 Magnesium: Is closely related to Ca and P: almost 70% is found in the skeleton.
Plays an important role in the activation of phosphates, metabolism of
carbohydrates (by activation of many enzyme systems). Deficiency may be due to
low Mg or high inorganic P and K in forage, or a change in the microbial flora of
the rumen (due to diet change). Symptoms include:
+ Grass tetany (nervousness, muscle twitching, paralysis, convulsion and death);
common in dairy cows shortly after calving and feeding only on pasture.
+ Magnesium tetany: common in calves fed milk only for extended periods.
Symptoms are similar to grass tetany.
+vasodilation: results to a reduction in blood pressure.
 Sulfur: is necessary for the synthesis of some amino acids (methionine, cysteine).
It is a component of 2 B-complex vitamins: thiamine and biotin. Its deficiency is
rare.
 Iron: Constituents of some enzymes, part of hemoglobin and myoglobin. Due to
its important role, it is often considered as a macro element. It also interacts with
Co for the synthesis of vit. B12. Deficiency symptoms include:
+ Anemia: can be a reduction in red blood cells, changes in the size of the RBCs
or low haemoglobin content of the RBCs. Anemia can be of hereditory, pathological
and nutritional causes. Nutritional anemia is mostly common in during the suckling
period in mammals when milk (very low in Fe) is the main feed component or entire
[Link] generally do not have Fe deficiency since they begin to eat grass early

25
enough after birth( most soils and consequently forage is rich in F.
+ Depraved appetite (pica): abnormally, the animal begins chewing bones.
+ loss of appetite for other normal feeds.
 Copper: It is a component of many enzyme systems, promotes the maturation and
increased lifespan of RBCs and maturation of bones and connective tissues,
maintains the integrity of the nervous tissue, formation of melanin pigment,
reproduction and functioning of the heart. Deficiency symptoms include anemia,
reduced growth, bone disorders, loss of hair pigment, brain lessions, loss of
myelin in the spinal cord, decreased fertility, poor resistance to infections,
paralysis in young animals. Excess Mo and to a lesser degree Mg can limit is
metabolism while a moderate supply of Mn will favor it. Mn-Cu association has
been found to have an anti microbial effect.
 Cobalt: It appears in enzymes involved in the synthesis of DNA and metabolism
of amino acids. Is required by rumen micro flora for the synthesis of vitamin B 12.
This vitamin is essential for the utilisation of propionic acid (an important source
of energy in ruminants)and catalyses the recycling of methionine from
homocysteine. Deficiency symptoms include: loss of appetite, emaciation, rough
coat, scaly skin, anemia, and reproductive failure.
 Manganese: Plays a role in normal skeletal development and growth, reproductive
performance, resistance to allergies and feed intoxications, proper functioning of
the nervous system. It activates many enzyme systems. Deficiency symptoms
include: poor growth, leg disorders, poor fertility, frequent abortions.
 Iodine: Is a component of thyroxine, a hormone necessary for heat regulation,
growth and proper functioning of the live and ovaries. The hormone also regulates
metabolism, circulation and muscle function. Deficiency symptoms include:
goitre, weak or dead newborns, poor fleece (in lambs), suppression of oestrus (in
females) and lack of libido (in males).
 Zinc: Plays essential role in enzyme systems, metabolism of nucleic acids, protein
synthesis and in the metabolism of carbohydrates and vit. A. Deficiency
symptoms include: stunted growth, skeletal deformation, skin disorders
(parakeratosis), low feed efficiency, delayed sexual maturity, high mortality rates,

26
 loss of appetite, vomiting.
 Molybdenum: Plays essential role in enzyme systems involved in the utilisation of
purines and nitrates in the [Link] importance is more associated to its toxic role.
Symptoms of Mo poisoning include: extreme diarrhoea, retarded growth, loss of
weight, anemia, male sterility, dermatosis, decreased lactation. High levels of Mo
in the diet will decrease Cu absorption.
 Selenium: It can perform some of the functions originally attributed to Vit E. Is
required by animals in traces. Symptoms of toxicity include: stiffness of joints,
lameness, loss of hair, skin lesions on the legs.
 Flourine: Traces are needed for the development of normal skeleton. Deficiency
causes dental carries, sometimes bone and joint abnormalities. Is toxic if taken in
excess. It is a cumulative poison.

Based on the quantity required by animals per day, minerals are classified as micro
minerals and macro minerals. Macro minerals are required in large quantities for about 1g/day.
Micro minerals are required in much smaller quantities about 0.01g/day.

Most of the minerals required by the animals are provided directly by forage the animals
consume. However, the mineral content of the forage is greatly dependent on the type of soil on
which the forage is cultivated. Therefore, it is necessary to analyze and establish the different
minerals of different forage at different season of the year. These analysis will provide indication
of the type and concentration of the mineral content of the soil. In case of deficiency, soil
correction can be made using fertilisers.

Ruminant diets (especially forages) are often often deficient in minerals; therefore it is highly
recommended that they be provided as supplements.
For animals that are on zero grazing minerals are supplemented in their diet in the form
of mineral block or directly (as salt NaCl), or incorporated in the concentrates.

The ability of an animal to utilise the minerals it has absorbed (mineral retention) is influenced
by various factors:

A) The mineral must be a form such that it can be assimilated. For example, the rate of
phosphorus in milk is 90% whereas phosphorus in most cereals is less than 40%.

27
B) The animal must have the necessary vitamins. Vitamins A and D are essential for the Ca
fixation (skeletal growth).

C) Proper functioning of hormones. For example: calcitonine (secreted by the thyroid gland)
promotes the deposition of calcium in the bones while parathormone secreted by the (parathyroid
gland) does the contrary (withdrawal of Ca from the bones).

D) The animal must be able to retain the mineral. The absorption coefficient of minerals varies
age (young animals retain better), breed (long slender cow breeds retain better than short
compact breeds).

E) There must be an equilibrium between the minerals. For ruminants, the Ca/P ration in the diet
should be close to 1.5.

e) Vitamins

The most important vitamin needed by ruminants are vitamin A and vitamin D. Most of
the other vitamins can be synthesized by the microorganism in the rumen.

+ Vitamin A: it helps in vision and synthesis of new cells. It is naturally abundant in

green plants. Therefore, it is a problem in the diet of animals in the intensive system only.

Deficiency symptoms of vitamin A include;

 Poor growth
 Malformation of bones
 Infertility
 Abnormal changes in soft skin tissues (keratinisation) of many body parts
 Loss of appetite
 Diarrhea
 Reduced resistance to infection
 Night blindness
+ Vitamin D: is essential for the efficient absorption and utilisation of Ca and P.

Vitamin D is obtained by exposing the animal to sunlight. Therefore vitamin D

28
insufficiency is common to animals that are under zero grazing. Symptoms of vitamin D
deficiency include;
 Rickets (weak soft bones in young animals)
 Osteomalacia (weak easily shattered bones in mature animals)
 Growth retardation

4.2 Digestibility of feed

No matter how highly nutritive the feed is, it will be of no or little value to animals if it is
poorly digestible. Therefore, digestibility influences the overall quality of the feed.

Digestibility can be expressed as the proportion of feed digested on feed consumed.

Quantity of Digested Feed X 100
% Coefficient of Digestibility=_______________________________
Quantity of Consumed Feed
Or

Quantity of Digested Feed Consumed-Quantity of

Nutrients in feces
% Coefficient of Digestibility=_______________________________________________ X 100

Quantity of nutrients in Feed Consumed

Factors that influence digestibility

 Moisture content in the feed

 Time used for digestion (degradation)
 Chemical constituents of the feed ( feed with much cellulose are less digestible)
 Structure or form of feed
 Age
 Breed
 Physiological state of animal

4.3 Measures of digestibility in animals

Digestibility, is the measure of the availability of the nutrients in the feed to the animal.

29
 The main factors that determine the nutritive value of feeds include composition and
digestibility.
Acceptability of the feed is also important because it determines the rate of feed intake. Feed
acceptability is influenced by taste, odour, and texture.
When digestibility is combined with chemical compounds intake data, one can make a fairly
accurate predictions of overall value and hence the potential production which a given feed can
support.
When estimating the nutritive value of a feed, the intake (or acceptability) is the initial factor
to consider because the feed will be useless or of little value if it is not readily consumed by the
animal, even if it is highly digestible.
Generally, more easily digestible feeds are more readily consumed than less digestible ones.
Measures of digestibility are easier and more accurate to obtain than measures of intake in
animal nutrition.
Some common measures of digestibility used in animal nutrition are as follows;
a) Conventional Digestion Trials (in vivo)
 Calls for the use of live animals
 Is the most realistic reliable method of measuring a feed digestibility.
 It is however time consuming, tedious and costly.
 Gives estimation of how the animal digests/utilizes the feed
 It involves feeding known quantities of the feed to an animal. The animal is first kept in
preliminary feeding to permit them adjust to the environment and the routing of holding
and feeding.
 The animal is given a trial or adjustment period during which time its normal regime is
gradually substituted with the experimental feed. This period is generally 10-14 days for
large animals by which time the animal is consuming only experimental feed.
 The animal is restrained in an individual cage (metabolic cage) such that a quantitative
collection of feces (and urine) can be made.
 Then the collection period follows during which time the animal is fed with the test feed
only.
 It also lasts for about 7 days.
 When estimates of N2 balance are desired, urine output is also measured and analysed.

30
  The feed should also be sampled regularly for analysis throughout the experimental
period. The feed is provided daily and must be constant in quantity and in composition
throughout the study. Mix properly to prevent the animal from selective consumption.
 If there is any feed uneaten between feeding times, it should be completely removed,
weighed and a representative sample analysed. The faeces of each animal should be
collected daily, weighed and to last chemical analysis.
 Do not collect samples from sick or stressed out animals.
 A routine procedure for operations should be outlined and respected throughout the study
including adjustment period.
 The time of day for execution should be the same and be respected throughout.
 For such trials, a minimum of three healthy animals per feed are required, more animals
are preferred.

% Nutrient Digestibility = nutrient intake – nutrient in feces x 100

Nutrient intake

Disadvantages: time, cost, tedious, etc.

b) Prediction of Digestibility from Chemical Composition (In Vitro).

It predicts digestibility based on the chemical compounds of the feed in question.
This involves the development of numerous regression equations relating various
chemical components of the feed to in vivo digestibility.
Generally, the digestibility estimates obtained from prediction equations are not as
precise as desired.
Nowadays, in vitro digestibility measurements are more extensively used to estimate
digestibility.

c) In vitro Fermentation (Degradation)

It is a laboratory technique which simulates the feed degradation process which occurs in
the rumen of live polygastric animals.
It is the most commonly used in vitro technique that is was devised by Tilley and Terry in

31
1963.
It provides a quick, inexpensive and precise prediction of in vivo digestibility in
ruminants also it is done in vitro.
The in vitro procedure does a better job of prediction than chemical composition analysis
because it takes into account all factors that affect digestibility, whether known or unknown,
which is not possible with current chemical methods.
This is accounted for primarily as a result of the use of rumen fluid from a donor animal
as the digestive agent.
The procedure involves moving a small quantity of the feed with rumen fluid that has
been previously diluted with a specified buffer. The buffer is formulated based on the
composition of animal saliva.
The feed/buffer/rumen/fluid mixture is incubated at 39 oC for 96 hours or until gas
production ceases.
During the incubation period, the micro-organisms present in the rumen fluid degrade the
feed by fermentation reactions. One of the by-products of this fermentation is gas production.
The volume of gas produced is recorded at regular intervals and these intervals vary
depending on the nature of the sample. For forages for example gas readings are recorded at 3
hours, 6 hours, 12 hours, 48 hours, 72 hours and 96 hours of incubation.
Gas produced can be fitted into a pre-established mathematical equation which predicts
the rate of degradation.
At the end of the incubation period, the residual mixture is boiled with neutral detergent
solution to dissolve and separate the soluble carbohydrates. Then the resulting mixture is filtered
and the insoluble fraction (NDF) recovered. The NDF fraction is dried and weighed.
Digestibility is calculated as :

% RD = weight of original sample used for incubation – weight of NDF fraction x 100
Weight of sample
One major advantage of this technique is that estimates of digestibility for a large number
of samples can be done simultaneously and the results obtained within a few days.
However, some variables may influence the data collected
i) Variations in the microbial population as a result of the diet of donor animal,

32
 differences between animals, inoculum processing.
ii) Variations due to different storage, grinding and processing techniques during sample
preparation.
iii) Differences attributable to the fermentation medium; buffer sample/inoculum ratio.
iv) Procedural variations such as length of fermentation and laboratory errors.
The variables can be standardized in various ways
 Microbial population; use more animals of same species, feed them with same diet used
for in vitro fermentation, fluid should be collected at a standard time after feeding the
animal.
 Sample variation, use finely ground samples, grind all in similar manner, serve, and
maintain sample weights.
 Fermentation medium; use the same rumen/buffer ratio, maintain fermentation at 39 oC
and at a pH between 6.9 and 7.1.
 Procedural variations; standardize incubation and time of fermentation, always include a
blank and a standard forage with each in vitro analysis in order to determine the validity
of your sample.
 All things being equal, the in vitro digestibility technique is the best means of laboratory
evaluation of digestibility available today because of its convenience and precision.

d) Nylon bag Digestibility Techniques

It is another technique method for estimating the digestibility of feeds. In this procedure
specially manufactured nylon bags are filled with a small known quantity of the experimental
feed (0.5 – 1g) in question and then incubated in the rumen of cannulated (or fistulated) animals.
The bags should not be too large or too small (5 x 15 cm) and special nylon material is
used such that it is not affected by the content of the rumen.
Also, the pore size of the bag should be small enough to prevent the passage of the feed
from the bag but large enough to permit microbial and other components of the fluid entry.
The sample bag ratio is also important and a ratio of approximately 10 mg/cm 2 of bag
surface is probably adequate.
The sample should be dry, finely ground, and sieved through a 2.5 mm screen.
After filling the bags with the sample, the bags are attached to a weighted cord to

33
prevent floating in the rumen and ensure adequate exposure to microbial digestion.
Then the weighted cords containing the bags is directly inserted into the rumens of the
previously cannulated animals.
Sheep are most appropriate for such experiments because they are easier to handle.
Care must be taken so that the animal is not affected in any way by the cannula. Regular
cleaning of the opening is crucial for the health of the animal. The animals must not be inserted
into the bags if it manifests signs of malaise or ill health or loss of appetite.
The bags are incubated inside the sheep for 72 hours, then removed, washed, dried and
weighted.
The difference in weight of bags before and after incubation represents the amount of
adequate feed degraded.
Percentage degradation is calculated using the formula
P= a + b (1- e-ct)
P= Percentage degradation at time t
a= water soluble fraction
b= insoluble but potentially degradable fraction
(a+b)= Potential Degradability
c= Rate of Degradation of insoluble fraction
A disadvantage of this technique is that only a limited number of samples can be run at
a time.
Most of the same variables which affect the in vitro degradation technique (diet of
animal, time of fermentation) will also affect the nylon bag technique. So measures should be
taken to minimize these variables.

4.4 Use of markers in animal nutrition

Indigestible markers or reference substances are used in animal nutrition.
They can be used to determine digestibility, intake, rate of passage of nutrients through
the GIT, site and extent of digestion and microbial protein synthesis in ruminants.
The ideal marker should have the following characteristics;
 Should be inert, with no toxic physiosecretion, digestion, absorption or motility of
content inside GIT.

34
  Should have physicochemical properties which allow for precise, quantitative analysis
and it must not interfere with other analysis.
 logical effect on the animal or microflora.
 Should not be absorbed or metabolized within the GIT.
 Should be physically similar to or intimately associated with the material it is to mark.
 Should not influence gastro intestinal
Unfortunately, none of the markers currently in use satisfy all these criteria. Therefore,
proper selection of marker must be made based on specific experimental conditions.
Markers have been divided into two groups; internal and external.
 Internal markers are integral components of the feed. Example, lignin, digestible ADF,
and NDF. However, lignin is the most widely used.
 External markers are indigestible substances added to a feed. Example chromium
sesquioxide (Cr2O3), cerium, dysprosium, etc.
Special markers used in determining the amount of microbial protein synthesized in
various diets include; diaminopimellic acid (DAP) and ribonucleic acid. DAP is found only in
bacteria, so if the DAP content of the digesta is determined and the DAP N ratio is known, we
can estimate the portion of N2 in the digesta that is of microbial origin.

Use of Markers to Determine Digestibility

Estimate of total tract or specific site of digestibility are possible with many markers.
The digestibility of a given nutrient can be determined by applying the following
equation:

% Nutrient Digestibility = 100 - [100 x (% marker in feed X % nutrient in feces)]

% marker in feces X % nutrient in feed

Similarly, estimates of digestibility in specific sites of the digestive track can obtained
by slightly modifying the equation

% Nutrient Digestibility in rumen = 100 - [100 x (%marker in feed X %nutrient in

duodenum)]

35
 %marker in duodenum X %nutrient in feed

36
Chapter 5: Energy systems in nutrition

The ability of feed to supply energy is very important in determining its nutritive value.
When the animal does not eat, it still requires energy for functions such as:
- Essential muscular activity
- Chemical work such as movement of dissolved substances against a concentration
gradient (active transport)
- Synthesis of used body constituents such as enzymes and hormones
Energy required for these processes in a starving animal is obtained from the breakdown of body
reserves, first glycogen, then fat and protein. When the animal is fed, the energy from the feed is
used primarily for maintenance, thus preventing the breakdown of body tissues.
Energy in excess of that needed for maintenance is used for various production functions such
as:
- Growth in young animals (muscle tissue)
- Fattening in adult animals (adipose tissue)
- Milk synthesis in lactating animals
- Work for animals used for traction
- Synthesis of wool (or eggs, etc)
Energy represents the ability of an animal to perform work and other productive processes. All
forms of energy can be converted into heat. As related to body processes, energy is expressed as
heat (calories).
Energy is not a nutrient but rather a property of nutrients. It is a principal requirement
for living organisms and as such most nutrients function to a large content as sources of energy.
5.1 Gross Energy (GE)
The quantity of chemical energy present in feed is measured by converting it into heat energy
and determining the heat produced. When a substance is completely burned to its ultimate
oxidation products of CO2 and H2O or other gases, the heat given off is termed gross energy or
heat combustion. Gross energy is generally measured in an O2 bomb calorimeter and is the
starting point for determining the energy value of feedstuffs. It can also be called the heat of
combustion of the feed and is measured in units of calories. A calorie is the amount of energy
needed to raise 1 gram of water by 1 oC (e.g from 14.5 to 15.5 oC). Heat can also be measured in

37
units of joules. One calorie is equal to 4.183 joules.
If a number of purified nutrients were used to determine gross energy, we will find that
on the average, fats contain about 8.9-9.6 Kcal/g of GE, carbohydrates contain 3.7-4.4 Kcal/g
and proteins 5.4 – 5.9 Kcal/g, cellulose contains 4.18 Kcal/g; maize contains 4.43 Kcal/g and hay
contains 4.51 Kcal/g (N.B. per g of dry matter). Therefore, gross energy can be calculated from
the proximate composition of feeds. This is done by multiplying the percentage of each nutrient
by its appropriate energy value (4.15, 9.4, 5.65 Kcal/g) followed by a summation of these values.
However, living organisms are not capable of capturing all of the energy in the in the
foods they consume due to digestive and metabolic inefficiencies.
Variable amounts of the GE in feeds are actually used in productive body functions.
The energy content of feeds can be expressed in various ways
5.2 Digestible Energy (DE): It is the first major energy loss that occurs in digestion because all
the feed is never completely digested. It is calculated as the amount of energy lost in feaces
when subtracted from gross energy.
Digestible energy = Gross energy – Faecal energy.
Fecal energy losses represent the largest loss of GE. It amounts to about 40-50% in
cattle and sheep feed roughages and 20-30% in cattle and sheep fed concentrates.
5.3 Metabolisable Energy (ME): Accounts for energy losses in feaces, in urine and as methane
(gas) from rumen fermentation. The principal components of urinary energy are in completely
oxidized nitrogenous compounds (mainly urea) and endogenous nitrogen constituents (mainly
creatinine).
ME = GE in feed – (GE in faeces + GE in urine + GE in gasses)
Urinary energy amounts to about 4-5% of GE in cattle.
Gaseous losses are quite small in most animal species (pigs, dogs, chicken) but are
considerable (7-10% of GE) in ruminants. Therefore, ME values are higher for non-ruminants
than for ruminants.
5.4 Heat increment: it is the amount of energy spent in chewing, mastication, digestion, enzyme
secretion in the alimentary tract, and microbial breakdown of crude fibre in the rumen,
absorption and utilization of feed. Heat increment comprises of two components: Heat of
fermentation and heat of nutrient metabolism. It is a loss of heat from the body. It is not
visible like feces and urine but may amount to 15-40% of GE. Heat fermentation is actually a

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 digestive loss that results from chemical inefficiency of fermentative processes.
5.5 Net Energy: It represents the portion of gross energy which is actually useful for body
functions. It is generally used in the body either for production (tissue growth, fat deposition,
milk production, foetal development, work) or for maintenance (basal metabolism, sustaining
body temperature, etc). It is calculated as the gross energy of feed minus the losses of energy
in faeces, urine, gasses and heat increment.

5.6 Total Digestible Nutrients (TDN): This is another approach devised to determine the usable
energy content of feeds (i.e. net energy). It uses digestibility data to account for fecal energy
losses in animals. It therefore provides values comparable to digestible energy.
TDN is defined as a measure of the digestible energy content of a feed valued on a carbohydrate
equivalent basis. Carbohydrate equivalent basis means all nutrients are scaled to the energy
equivalent of carbohydrates.
For lipids, the scaled equivalent factor is 2.25 and is 1 for proteins.
To obtain TDN, the proximate composition of the feed is determined and the value
obtained for each nutrient is multiplied by its respective digestion coefficient and by the
carbohydrate equivalent factor.
The digestion coefficients are standard published values which have been
internationally accepted.
An example of TDN calculation is as follows:
Proximate Percentage Digestion Factor Value
components coefficient %
Crude Protein 10 65 1 6.5
Crude Fiber 30 50 - 15
NFE 40 90 - 36
Ether Extract 5 90 2.25 9
TDN = 66.5

e.g. CP = 10 x 65 x 1 = 6.5
100

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 In species which procedure minimal quantities of gases, TDN is very similar to
metabolisable energy.

5.7 Physiological Fuel Values (PFV): They are caloric values for nutrients developed by
W.D Atwater. He studied the caloric value of each nutrient (carbohydrates=4.5; lipids=9.4;
proteins=5.65), the digestibility of the nutrient in human and the energy lost in urine with
protein. Then, from these, he calculated the average caloric values for the three classes of
nutrients.
Because urinary excretion is considered, PFV’s are quite similar to metabolisable
energy.
An example for calculation of PFV is as follows:

Source GE Kcal/g Human % Urinary PFV’s Rounded

Apparent Energy Loss Kcal/g PFV’s
Digestibility Kcal/g
Carbohydrates 4.5 97 0 4.03 4
Fat 9.4 95 0 8.93 9
Protein 5.65 92 1.25 4.05 4

A flow chart illustrating energy distribution in body processes is as follows:

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 Production :–tissue,
growth, fat, etc; - stored
in products (milky eggs);
- work

Net Energy
Maintenance:
–basal metabolism;
-activity at
maintenance;
Heat increment: - -sustaining body
heat of digestive temperature
Metabolisable
fermentation and
Energy
action;

-heat of nutrient Energy wasted as

metabolism heat

Digestible Urinary Energy losses: -

Energy residues of imperfect nutrient
metabolism (N2-compounds);

–endogenous catabolism
Gross Energy (creatinine)
(GE)
Gaseous Energy losses:

–gaseous products of
Fecal Energy Lost via bowels or
fermentation (CH4)
Undigested feed residues belching
Metabolic products (enz,
bacteria, mucosa

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