DIGESTION AND METABOLISM OF CARBOHYDRATES AND FATS IN
RUMINANTS
A Review
By
Yidersal Erega (MSc, Animal Nutrition)
May, 2020
Woldia University, Ethiopia
TABLE OF CONTENTS
TABLE OF CONTENTS................................................................................................................ii
LIST OF TABLES.........................................................................................................................iii
LIST OF FIGURES........................................................................................................................iv
1. INTRODUCTION....................................................................................................................1
2. DIGESTION AND METABOLISM OF CARBOHYDRATES AND FATS IN
RUMINANTS..................................................................................................................................3
2.1. Carbohydrates and Fats in Ruminants Nutrition............................................................................3
2.2. Carbohydrate Digestion in Ruminants...........................................................................................5
2.3. Carbohydrate Metabolism in Ruminants.......................................................................................8
2.3.1 Volatile fatty acid production in the rumen................................................................................10
2.3.2 Gluconeogenesis.........................................................................................................................12
2.4. Fat Digestion in Ruminants.........................................................................................................13
2.5. Fat Metabolism in Ruminants......................................................................................................16
2.5.1 Fatty acid synthesis.....................................................................................................................16
2.5.2 Lipolysis......................................................................................................................................18
3. CONCLUSIONS....................................................................................................................20
4. REFERENCES.......................................................................................................................21
ii
LIST OF TABLES
Table Page
iii
LIST OF FIGURES
Figure Page
iv
1. INTRODUCTION
Ruminants are cloven-hoofed mammals of the order Artiodactyla, obtain their food by browsing
or grazing, subsisting on plant material. Today, 193 species of living ruminants exist in 6
families: Antilocapridae, Bovidae, Cervidae, Giraffidae, Moschidae and Tragulidae. The number
of wild ruminants is about 75 million and of domesticated about 3.6 billion (Hackmann and
Spain, 2010). Approximately 95% of the population of domesticated ruminants constitute
species: cattle, sheep and goats, all of them belong to the Bovidae family. Cattle, goats and sheep
are the most numerous species and cattle is of the most economic importance. The economic
value of milk and beef production in the EU is almost 125 billion Euro per year and accounts for
40% of total agricultural production (FAIP, 2003).
An important characteristic of ruminants’ digestive system is the occurrence of the microbial
fermentation prior to the gastric and intestinal digestion activity (Bird et al., 1996). The
feedstuffs consumed by ruminants are all initially exposed to the fermentative activity in the
rumen, the place of more or less complete microbial fermentation of dietary components (Hall
and Herejk, 2001). Digestion of carbohydrates is initiated in the mouth in most species. Limited
digestion of carbohydrates, fats, and proteins occurs in the stomach in most species. Extensive
digestion of carbohydrates, fats, and proteins occurs in the duodenum (DeFrain et al., 2004).
Fermentation of fibrous carbohydrates occurs in the rumen of ruminants and the cecum for those
species with functional ceca. Absorption of degradation products of carbohydrates, fats, and
proteins occurs in the duodenum and ileum. Dietary carbohydrates are the main rumen microbial
fermentation substrates. Microbial yields are related primarily to the growth rate that
carbohydrate permits (Lock et al., 2006).
Major differences in processes of fat digestion and absorption exist between ruminant and non-
ruminant animals, as a result of the profound impact of the rumen on dietary lipids. Bacteria in
the rumen split off the fatty acids (and sugars) from the glycerol backbone. The glycerol and the
sugars released from glycolipids are fermented to the volatile fatty acids (VFA) (DeFrain et al.,
2004). The breakdown of dietary lipids by rumen bacteria generally occurs quite rapidly as the
lipids are exposed during rumination and bacterial digestion of feed particles. The individual
carbohydrates characterized by faster rumen degradation rates result in greater microbial yield
1
(Hall and Herejk, 2001). The enzyme systems produced by microorganisms for carbohydrates
hydrolysis are complex; they usually comprise hydrolases from several families, and there may
be multiple enzymes hydrolysing each polysaccharide. Nearly all carbohydrate digestion occurs
(>90%) within the rumen, but under certain circumstances (e.g., high rate of passage), a
significant amount of carbohydrate digestion can occur in the small and large intestine (Jenkins,
1993). Carbohydrates and fats are commonly perceived as having a primary role in the diet as
suppliers of energy; however, these feed fractions are not uniform in terms of their value as
energy sources (DeFrain et al., 2004). . Also, energy does not come from fats or carbohydrates
in the form of calories but as absorbable nutrients that the animal may metabolize to yield energy
or use for anabolic purposes. The diverse fractions of carbohydrates and fats have potential to
provide specific nutrients that support both microbial and animal needs in ruminants (Lock et al.,
2006).
Fat is an important energy component in the diet of ruminants and over the last decade fat
supplementation has become a common practice to increase the energy density of the diet for
high producing dairy cows. Fat and fatty acid metabolism and digestion in ruminants is of
considerable interest at the present time, both to scientists and the agricultural industry (DeFrain
et al., 2004). . The subject has received renewed interest for a number of reasons; first, the use
of dietary fat supplements has increased, and will continue to do so, as nutritionists strive to
increase the energy density of diets to meet requirements of the high producing dairy cow, and
second, we now recognize that fatty acids, both of dietary and rumen origin, can have specific
and potent effects on ruminant metabolism and human health (Lock et al., 2006).
Therefore, the objective of this paper is to review updated information about participatory
range land management in Africa.
2
2. DIGESTION AND METABOLISM OF CARBOHYDRATES AND FATS
IN RUMINANTS
2.1. Carbohydrates and Fats in Ruminants Nutrition
Historically, carbohydrates and fats were valued on their caloric contributions to diets. Feeding
recommendations for these feed fractions now address inclusion levels as well as consideration
of the positive and negative effects of specific types of these nutrients. Feed carbohydrate
characterization has expanded beyond fiber and nonfiber carbohydrates (Jenkins, 1993). Fiber
now encompasses ADF, NDF, physically effective fiber, and fiber digestibility to describe
effects on diet composition, rumination, rumen fill, potential fermentability, and nutrient
contribution (Bird et al., 1996). The nonfiber carbohydrates are now parsed into sugars and
fructans (both in water-soluble carbohydrates), starch, pectins, and others, all of which can differ
in their effects on rumen pH or support of microbial growth (DeFrain et al., 2004).
Dietary fat has the advantage of providing energy without increasing the risk of ruminal acidosis.
However, there are specific considerations for amounts and types fed in high-versus low-forage
diets. Fats can affect ruminal fermentation, having the potential to depress fiber digestion or
affect ruminal methane production. Carbohydrates are the most important source of energy and
the primary precursors of fat and sugar (lactose) in cows' milk (Deiuliis et al., 2010). The
microorganisms living in the rumen allow the cow to obtain energy from fibrous carbohydrates
(cellulose and hemicellulose) which are bound with lignin in plant cell walls or fiber. Since fiber
is bulky it is retained in the rumen where the cellulose and hemicellulose are fermented slowly.
As plants mature, the lignin content of fiber increases and the extent of cellulose and
hemicellulose fermentation in the rumen decreases (Deiuliis et al., 2010).
Fiber in the form of long particles is essential to stimulate rumination. Rumination enhances the
breakdown and fermentation of fiber. The contribution of dietary carbohydrates to the energy or
nutrient supplies of the animal is affected by location and extent of digestion, which alter types
and amounts of products made available to the animal (Jenkins, 1993). Carbohydrates digested in
the small intestine provide monosaccharides, notably glucose from starch. Ruminally, products
of fermentation are much more diverse and come in the forms of gases, organic acids, and
microbial mass. Types of fermentation acids produced include acetate, propionate, butyrate,
3
valerate, and lactate. Molar proportions of acids produced will vary to some degree with
carbohydrate type, with acetate predominating with pectins and butyrate with sucrose and lactose
(DeFrain et al., 2004).
Figure 1. Plant carbohydrates digestible in the small intestine or strictly by gut microbes
(indigestible). (Source DeFrain et al., 2004).
Most of the dietary fat consumed by dairy cows is in the triglyceride form and consists
predominantly of unsaturated fatty acids (FA). The triglycerides are hydrolyzed in the rumen to
glycerol and FA by lipase predominately from bacteria and minor contributions by plant lipases.
The glycerol is metabolized by ruminal microorganisms to propionate, which is absorbed across
the ruminal wall (Jenkins, 1993). Use of stored fat for energy is important for animals during
states of energy deficit due to limited food supply or lack of appetite (e.g., the transition dairy
cow); however, excessive mobilization of FA from adipose tissue increases the risk for fatty liver
and ketosis (Deiuliis et al., 2010). It is important to note that research continues to be done on the
signaling and pathways for FA mobilization from adipose tissue. Traditionally, we considered
that during an energy deficit, hormone sensitive lipase in adipose is signaled to initiate hydrolysis
of FA from triglycerides stored in lipid droplets within adipose tissue. However, it is now
4
understood that adipose triglyceride lipase is the primary enzyme signaled to initiate hydrolysis of
the first FA from the triglyceride (Deiuliis et al., 2010).
2.2. Carbohydrate Digestion in Ruminants
Carbohydrates are as important to the ruminant animal as they are to non-ruminants, since they
provide the glucose necessary for the adequate function of cells. However, in the ruminant,
ruminal fermentation transforms most of the cell wall polysaccharides and all of the intracellular
carbohydrates present in the forage into short-chain volatile fatty acids, which are then absorbed
by the rumen epithelium (Bird et al., 1996). In the non-ruminant, starch digestion occurs mainly
in the small intestine. The situation in the ruminant differs due to the action of microorganisms in
the rumen. Digestion of starch to glucose requires the action of several enzymes produced by the
salivary glands, the rumen microorganisms or the pancreas and small intestine. Amylase secreted
by the nasolabial glands is found at relatively high levels in the saliva of some ruminants, such as
the buffalo. Several species of ruminal bacteria are able to digest starch (Michel, 2010).
Amylolitic organisms are found in larger percentages of the total microbial population when
rations high in starch are fed. Important species that have been enumerated in cattle fed high
grain diets are Bacteroides amylophilus, Butyrivibrio fibrisolvens, Bacteroides ruminocola,
Selenomona lactylitica, Streptococcus bovis, Prevotella ruminocola, Eubacterium ruminantium,
Ruminobacter amylophilus, Ruminococcus bromii, Succinimonas amylolytica and Lactobacillus
sp. The two digestible carbohydrates are starches and sugars, and both of these carbohydrates are
digested, or broken down into their most elementary form, along the gastrointestinal tract
(Michel, 2010). Amylase, an enzyme which breaks apart starches, is found in the mouth and in
the small intestine. Similarly, the three major enzymes which break apart sugars -- sucrase,
maltase and lactase -- are also found in the mouth and in the small intestine. Once these digested
starches and sugars begin to move through the small intestine, they are able to be absorbed.
The major sources of carbohydrates in ruminants are fibrous feeds containing cellulose and
hemicellulose and grains rich in starch are the primary carbohydrate sources. Ruminants have
most of their dietary carbohydrates (e.g., starch and cellulose) fermented in the rumen by
microorganisms, and only 5 to 20% of consumed dietary carbohydrates are digested in the small
intestine. As early as 1883, Tappeiner demonstrated that in cattle dietary cellulose was fermented
5
to volatile fatty acids (VFA), most of which was acetic acid. Numerous ruminant nutritionists
have confirmed these observations and have shown that the major source of energy to the
ruminant is the VFA absorbed from the rumen and other parts of the digestive tract (Michel,
2010).
A major breakthrough in the development of VFA biochemistry in ruminants occurred with the
development of gas-liquid chromatographic quantification of VFA. VFA are also produced from
dietary proteins, pentoses of nucleic acids, and glycerol of glycerol-phospholipids (Hall and
Herejk, 2001). Acetate is a major end product of fermentation, elucidated pathways for synthesis
of each VFA and lactate, showed how diet composition influences proportions of each of the end
products of fermentation, demonstrated that about half of butyrate is converted to β-
hydroxybutyrate during absorption of butyrate from the rumen into blood, and showed that
branched-chain organic acids are derived from amino acid fermentation. High-starch diets
increased the flux of lactate to propionate via the acrylate vs. the succinate pathway of pyruvate
conversion to propionate.
Dietary carbohydrates are the main rumen microbial fermentation substrates. Microbial yields
are related primarily to the growth rate that carbohydrate permits. The individual carbohydrates
characterized by faster rumen degradation rates result in greater microbial yield (Hall and
Herejk, 2001). The enzyme systems produced by microorganisms for carbohydrates hydrolysis
are complex; they usually comprise hydrolases from several families, and there may be multiple
enzymes hydrolysing each polysaccharide. Nearly all carbohydrate digestion occurs (>90%)
within the rumen, but under certain circumstances (e.g., high rate of passage), a significant
amount of carbohydrate digestion can occur in the small and large intestine (Michel, 2010).
The concentration of monosaccharides, glucose and fructose was estimated from 1% to 3% (in
grasses and herbage) and of sucrose from 2% to 8% (Smith, 1973). Sucrose formed from -D-
glucose and D-fructose linked by 1, 2 glycosidic linkage is digested by enzyme sucrose
phosphorylase (Stan-Glasek et al., 2010). Maltose formed from two units of glucose joined with
an α-1–4 bond is digested by enzyme α–glucosidase. Oligosaccharides concentration in the
different plants ranges between 0.3% and 6% and represent a wide diversity of biomolecules
(including stachyose and raffinose), they are chains of monosaccharaides that are two to
approximately 20 units long (Hall and Herejk, 2001). The enzymes belonging to the group of
6
polysaccharide hydrolases which hydrolyse the glycosidic bond between two or more
carbohydrates utilize oligosaccharides (Courtois, 2009).
Structural carbohydrates is less digestible than NFC and is negatively correlated with energy
concentration in the diet for ruminants, but is important for rumination, saliva flow, ruminal
buffering, and health of the rumen wall, however, high dietary concentrations can limit dry
matter intake by increased rumen fill (Michel, 2010). The retention time of plant fiber in the
rumen is sufficiently long (48 h or more in some species) to allow the growth of a fibrolytic
microbial population whose extensive fiber utilization contributes a major portion of the energy
for the animal (Van Soest, 1994). The cellulose fibers are embedded in a matrix of other
structural biopolymers, primarily hemicelluloses and lignin. The high-fibre forage diets
encourage the grown of acetate producing bacterial species, the acetate: propionate: butyrate
molar proportion would typically be in region 70:20:10 (France and Dijkstra, 2005). Fiber
digestion may be reduced due to decreased rumen pH (the fiber digesters are most active at a pH
of 6.2 to 6.8), and the availability of surface area for colonization. Fungi plays an active and
positive role in fiber degradation (Bazooyar et al., 2012).
Starch is a complex of two structurally distinct polymers: amylose and amylopectin (Chesson,
1997). Amylose is chemically composed of -1,4- linked polymers of glucose. It is degraded by
-amylases, which releases oligosaccharides maltodextrins, and - amylases, which remove
maltose units. Amylopectin is a highly branched molecule of -1, 4- linked polymers of glucose
joined 1,6 at intervals along the backbone molecule (Hall and Herejk, 2001). Amylopectin is
degraded to maltose by -amylases action (in 50%), glucanohydrolases and isoamylase. Maltose
and maltodextrins are degraded to glucose by -glycosidase (Michel, 2010). Starch can be
degraded by ruminal microbial enzymes as well as enzymes in the small intestine of the
ruminats. Of the nonfibrous fraction, starches are the highest proportion in the diet, and cereal
grains is the major source of starch in ruminants diet (Bazooyar et al., 2012).
The cereal grains differ in their starch content, with wheat containing (on dry matter basis) 77%
starch, corn 72%, and barley and oats 57 to 58% (Huntington, 1997). Differences exist among
cereal grains in their extents and rates of ruminal starch degradation. In the rumen has been
7
digested from 55 to 70% of corn starch, 80 to 90% of barley and wheat starch, and 92 to 94% of
oats starch (Huntington, 1997). The degradation rates is estimated from 4.0 to 6.4% h-1 for corn
starch and from 14.7 to 24.5% h-1 for barley starch (Michel, 2010). On average, 5 to 20% of
starch consumed is digested postruminally, mainly in the small intestine (from 45 to 85% of
starch entering the duodenum), this capacity is limited by the supply of pancreatic amylase
(Hunhington, 1997).
Cellulose content is in the range from 35 to 50% of plant dry weight (Lynd et al., 2002). It is
chemically composed of a homogenous polymers of β-1,4-D-glucose linked through β-1,4-
glycosidic bonds. Native cellulose exists as fibrils which are composed of amorphous and
crystalline regions formed from cellulose chains, each of them contains between 500 and 14,000
b-1,4D-glucose units (Bazooyar et al., 2012). The digestion of cellulose necessitates a
combination of many classes of cellulases. The digestion process including activity of
endoglucanases, that cut randomly at internal amorphous sites in the cellulose chain;
exoglucanases, that act processively on the reducing or non-reducing ends of cellulose chains,
releasing either cellobiose or glucose as major products; and glucosidases that hydrolyze soluble
cellodextrins and cellobiose to glucose (Lynd et al., 2002).
Hemicellulose concentration varies from 6 to 22% (on dry matter basis) in leaves of grasses and
herbs (Schädel at al., 2010). Hemicelluloses are composed of complex heteropolymers that vary
considerably in primary composition, substitution and degree of branching, and can be grouped
into four classes: xylans, xyloglucans, mannans and mixedlinkage β-glucans (Ebringerova et al.,
2005). Bacteroides (Fibrobacter) succinogenes, Ruminococcus albus, and Ruminococcus
flavefaciens and same strains of Butyrivibrio fibrisolvens and Bacteroides ruminicola are
considered to be the organisms responsible for most of the degradation of hemicelluloses. Rumen
degradation of hemicelluloses varies from 16 to 90%, depending on their composition (Schädel
et al., 2010). Lignin, a complex phenolic polymer, is indigestible by rumen microbes, but their
concentration limits digestibility of structural carbohydrates (Van Soest, 1994). The main reason
for reduction of accessibility for the hydrolases secreted by ruminal microbes is the presence of
strong covalent bonds between lignin and the cell wall polysaccharides (Chesson, 1997).
8
2.3. Carbohydrate Metabolism in Ruminants
In farm animals, dietary carbohydrates provide well over one half of the energy needs for
maintenance, growth, and production. Glucose is a primary energy source for certain animal
tissues and a precursor for lactose synthesis in the mammary gland (Michel, 2010).
Consequently, understanding carbohydrate digestion and absorption, dietary glucose availability,
and the involvement of gluconeogenesis in the regulation of glucose homeostasis is essential for
the manipulation of the production and quality of agricultural foods. Dietary carbohydrates are
mainly starches, disaccharides, monosaccharides, and fibrous carbohydrates (cellulose,
hemicelluloses) (Hall and Herejk, 2001). The luminal phase of small intestine digestion results in
amylase hydrolysis of starches, yielding oligosaccharides (alpha-dextrins, maltotriose, and
maltose, but no free glucose), but the fibrous carbohydrates remain intact. In monogastric
herbivores fed high-fibre diets, the increased bulk accelerates the transit of chime and lowers the
efficiency of luminal digestion of nonfibrous carbohydrates.
Carbohydrates contain carbon and, hydrogen, and oxygen atoms in a ratio equivalent to one
carbon (C) to one water (H2O), or C(H2O). This diverse group of substances includes,
particularly the sugars and their derivatives. The key substance for intermediary metabolism is
glucose. Important roles are also played by fructose and galactose (hexoses), and by ribose and
deoxyribose (pentoses), by glycerol and glyceraldehydes (triose), by sedoheptulose (heptose),
and by erythrose (tetrose). Combined with lipids, carbohydrates form glycolipids, and with
proteins they form glycoproteins. After absorption or gluconeogenesis (see later), glucose can
undergo catabolism to provide energy or metabolites for synthetic pathways. The first stage of
glucose metabolism is glycolysis or Embden-Meyerhopf fermentation (Michel, 2010).
Under anaerobic condition, glucose breaks down to lactate, yielding a net 2 mole of ATP per mol
of glucose. Four moles of ATP is generated, but, two is used to phosphorylate first glucose and
then fructose-6-phosphate. Under aerobic conditions, the reduced NADPH plus H+ may be
oxidized via oxidative phosphorylation to produce 3 mol ATP and pyruvate. Pyruvate then enters
the tricarboxylic acid (TCA) cycle (Kreb’s cycle) and becomes oxidized to carbon dioxide and
water generating 15 mol of ATP. During vigorous exercise, muscle cells produce lactate faster
than it can be utilized, via pyruvate, by the cells’ mitochondria (Michel, 2010). The excess of
lactate diffuses into the capillary blood and is carried to the liver to form the basis for
9
gluconeogenesis. This is called the Cori cycle. Another route for glucose metabolism is the
pentose phosphate pathway, which is important in generating the NADPH used in synthesis of
fatty acids in adipose tissue, mammary gland and liver in response to demand. Nutrients or
metabolites differ greatly in their relative efficiency as sources of ATP energy (Hall and Herejk,
2001)..
2.3.1 Volatile fatty acid production in the rumen
During ruminal fermentation, the population of microorganisms (chiefly bacteria) ferments the
carbohydrates to produce energy, gases (methane - CH4 and carbon dioxide - C O2), heat, and
acids. Acetic acid (vinegar), propionic acid and butyric acid are volatile fatty acids (VFA) and
make up the majority (>95%) of the acids produced in the rumen. Also, the fermentation of
amino acids produces some acids called iso-acids. The energy and the iso-acids produced during
fermentation are used by the bacteria to grow (primarily to synthesize protein). The CO2 and
CH4 are eliminated through belching, and the energy of the CH4 is lost. Unless heat is necessary
to maintain body temperature, the heat produced during fermentation is dissipated. The VFA,
end products of microbial fermentation, are absorbed through the rumen wall.
Table 1: Volatile fatty acids produced by ruminal fermentation
Most of the acetate and all the propionate are transported to the liver, but the majorit of butyrate
is converted in the rumen wall to a ketone body called β-hydroxybutyrate. Ketones, are important
sources of energy (fuel for combustion) for most tissues in the body. Ketones come primarily
from the butyrate produced in the rumen, but in early lactation, they also come from the
mobilization of adipose tissue (Hall and Herejk, 2001). Most of the propionate is converted to
glucose by the liver. In addition, the liver can use amino acids for glucose synthesis. This is an
important process because there is normally no glucose absorbed from the digestive tract and all
the sugar found in the milk (about 900 g when a cow produce 20 kg of milk) must be produced
by the liver. An exception arises when cows are fed large amounts of concentrates rich in starch
or a source of starch resistant to ruminal fermentation.
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Then, the starch that escaped fermentation reaches the small intestine. The glucose formed
during intestinal digestion is absorbed, transported to the liver and contributes to the supply of
glucose to the cow. Lactate is another possible source of glucose in the liver. Lactate is found in
well preserved silages, but lactate production in the rumen occurs when there is excess starch in
the diet. This is undesirable because the rumen environment become acidic, fiber fermentation
stops and in extreme cases the cow stops eating (Hall and Herejk, 2001).
11
Figure 2. Carbohydrate Metabolism in Dairy Cows. (Source Michel, 2010)
2.3.2 Gluconeogenesis
Because of the central role of glucose in energy metabolism, the body has mechanisms to keep
glucose concentrations in extracellular fluids in an appropriate range. Excess blood glucose leads
12
to glycosuria and the syndrome of diabetes mellitus. This disorder occurs in pet animals and
horses but rarely in other species. A. greater problem for many animal species, particularly
ruminants, is avoiding the other swing of the pendulum to hypoglycaemia. All species have
metabolic enzyme pathways to synthesize glucose from other substances, but the need of
ruminants is much more chronic and continuous because of the general lack of absorbable
glucose from the intestine (Hall and Herejk, 2001).
Three groups of starting substances are available for gluconeogenesis: carbohydrates, glucogenic
amino acids via deamination, and odd-numbered fatty acids. All species have access to
carbohydrates (lactate, pyruvate, and oxaloacetate). Glucogenic amino acids (especially alanine
and glutamate) can be oxidatively deaminated and transferred to the carbohydrate pathways via
alpha-keto acidswith reduced nicotinamide coenzymes. The reactions are reversible; the
hydrogenases are located inside the mitochondria. The third class of gluconeogenetic substrates
comprises [Link] acids (odd numbers of carbon atoms), which yield odd-number
acyl-CoA derivates, particularly propionate. This is a major and essential contributor to
gluconeogenesis in ruminants.
The utilization of propionate involves its activation to propionyl-CoA and addition of a carboxyl
group to form D-methylmalonyl-CoA (using propionyl CoA carboxylase, which contains biotin).
D-Meth-ylmalonyl CoA is then racemized to the L form, which, in turn, is converted to succinil-
CoA by methylmalonyl-CoA mutase, an enzyme that contains a derivative of cobalamin (vitamin
B12. The cobalamin serves as a source of free radicals that allow the removal of hydrogen atoms
and subsequent intra-molecular rearrangement to succinate. The glucose formed can itself serve
as the source material for the synthesis of other important carbohydrates: glycogen and lactose.
Glycogen is the chief storage carbohydrate. Lactose (milk sugar) is specifically synthesized in
the mammary glands of lactating animals (Hall and Herejk, 2001).
2.4. Fat Digestion in Ruminants
Fat digestion and metabolism differ widely between animal species. In ruminants, dietary fats are
hydrogenated in the rumen before intestinal absorption so that absorbed fatty acids (FA) are
more saturated than dietary FA (Deiuliis et al., 2010). In non-ruminants, intestinal FA
13
digestibility depends on the level of saturation of dietary FA. Fat supplementation of the diet of
cows decreases milk protein and has a variable effect on milk fat, depending on the source of
dietary lipids. When encapsulated lipids are used, the linoleic acid content of milk is increased,
but the organoleptic quality of milk may be altered. Supplementary lipids are incorporated into
non-ruminant body fat, whereas lipogenesis is reduced. The digestive utilization of fats by
ruminants is characterized by events in the rumen before they are absorbed in the intestine.
During their stay in the rumen, fats are transformed so that the amount and composition of fat
leaving the rumen differ from intake. Moreover, fat has negative effects on ruminal activity,
mainly on carbohydrate degradation (Doreau and Ferlay, 1995).
Most of the dietary fat consumed by ruminants is in the triglyceride form and consists
predominantly of unsaturated fatty acids (FA). The triglycerides are hydrolyzed in the rumen to
glycerol and FA by lipase predominately from bacteria and minor contributions by plant lipases
(Jenkins, 2006). Fat is an important energy component in the diet of ruminants and over the last
decade fat supplementation has become a common practice to increase the energy density of the
diet for high producing dairy cows. Dietary lipids are extensively hydrolysed in the rumen, by
microbial enzymes originating first from different classes of bacteria (of which the best known is
Anaerovibrio Zipolytica), galactosidases and phospholipases, and lead to the formation of free
fatty acids (FA), without intermediate compounds such as mono- and di-acylglycerols. This
lipolysis is very rapid, in vitro as well as in vivo. Some factors have recently been found to
decrease lipolysis, such as antibiotics and low p H. Lipolysis is reduced with diets rich in starch
(Gerson et al. 1985) (Van Nevel & Demeyer, 1995).
Hydrogenation occurs on free FA. Thus it is necessary that lipolysis has taken place previously.
The first step is an isomerization. Hydrogenases allow the reduction of FA according to different
pathways described by Harfoot and Hazlewood (1988). The endproduct of hydrogenation of C18
FA is stearic acid. However, when large amounts of linoleic acid are available, hydrogenation
generally stops before this final step, leading to various cis and trans isomers of monoenoic FA.
The most important is trans-vaccenic acid. Hydrogenation of polyunsaturated C20 and C22 FA
has been discounted by Ashes et al., (1992) but other results show extensive hydrogenation of
these FA in vivo and in vitro. Hydrogenation generally takes place at a slower rate than lipolysis,
but few polyunsaturated FA are present in the rumen (Deiuliis et al., 2010).
14
Bacteria incorporate FA, but are also able to synthesize a wide variety of FA, those with 15 and
17 C atoms being the more characteristic. Synthesis occurs mainly from volatile FA; branched-
chain FA arise from isobutyrate and isovalerate. Bacterial cis and trans monounsaturated FA may
result from desaturation of saturated FA (Jenkins, 2006). Linoleic acid can also be synthesized
(Demeyer & Hoozee, 1984). The extent of this synthesis is lower than the extent of dietary FA
incorporation and decreases when ruminal FA concentration increases. Protozoa and fungi can
also incorporate and synthesize FA. Synthesized and assimilated FA are esterified as
phospholipids and sterol esters, and constitute structural lipids. When large amounts of FA are
fed, they are stored as free FA in cytosolic droplets. These droplets are especially rich in linoleic
acid whichthus escapes hydrogenation.
Major differences in processes of fat digestion and absorption exist between ruminant and non-
ruminant animals, as a result of the profound impact of the rumen on dietary lipids. As described
above, dairy cows consume a diet that predominantly contains PUFA as part of plant
triglycerides and glycolipids. Bacteria in the rumen split off the fatty acids (and sugars) from the
glycerol backbone. The glycerol and the sugars released from glycolipids are fermented to the
volatile fatty acids (VFA). The breakdown of dietary lipids by rumen bacteria generally occurs
quite rapidly as the lipids are exposed during rumination and bacterial digestion of feed particles
(Deiuliis et al., 2010).
In addition, the process generally is essentially complete so that no monoglycerides or
diglycerides pass to the lower digestive tract. The major exception to this would be when a
highly saturated (or hydrogenated) triglyceride is fed. Because of the very high melting point of
such fats, and their resulting low solubility, bacterial enzymes cannot gain access to the bonds
linking fatty acids to glycerol, and so these would pass to the lower digestive tract.
Unfortunately, the same limitations of solubility and melting point result in poor access of the
animal’s digestive enzymes in the small intestine, and very poor digestibility in that site as well.
The fatty acids released in the rumen are not absorbed from the rumen, but rather will pass to the
abomasum and then the small intestine, which is the primary site for absorption of the fatty acids.
However, the profile of fatty acids that reaches the intestine will be very different from what the
animal has consumed. This is because of the extensive bio-hydrogenation that occurs in the
rumen as a result of bacterial activity (Jenkins, 2006).
15
Unsaturated fatty acids are toxic to many of the species of the rumen bacteria, particularly those
that are involved in fiber digestion. Also, because of the anaerobic environment of the rumen,
there is an excess of hydrogen that the microbial population is continually interested in getting
rid of. (This is the primary reason why methane is produced within the rumen, as it is an
important “sink” for disposal of hydrogens from the rumen to keep rumen fermentation running
efficiently.) Consequently, as unsaturated fatty acids are released from the glycerol backbone,
they are quickly hydrogenated to saturated fatty acids. In cows fed most typical diets, more than
90% of the unsaturated fatty acids will be bio-hydrogenated to produce saturated fatty acids that
flow to the small intestine.
The lipids that leave the rumen are predominantly free fatty acids (85-90%) and phospholipids
(10-15%) found as part of microbial cell membranes. In the rumen, most of the free fatty acids
are actually found as potassium, sodium, or calcium salts of fatty acids because of the near-
neutral pH in the rumen contents (6.0 – 6.8). After passing through the acid conditions (pH ~2.0)
of the abomasum, however, the fatty acid salts are dissociated and the free fatty acids will be
found adsorbed (or “stuck”) to the surface of small feed particles that pass as part of the digestive
contents. The fatty acids making up the free fatty acid portion will be predominantly saturated
(80-90%), with about two-thirds stearic acid and about one-third palmitic acid.
It is often considered, in ruminants, that fat supplementation increases fat digestibility. This is a
consequence of the underestimation of fat intake in diets non-supplemented with fat, in which
FA, incorporated in cell structures, are not easy to extract by classical methods, whereas
supplemental lipids, which are generally triacylglycerols, are easier to extract; the fact,
mentioned earlier, that ruminal FA balance is generally positive with non-supplemented diets
and negative with fat-supplemented diets. For this reason, old results obtained by measurement
of digestibility between mouth and faeces, are questionable and should be used only to compare
the digestibility of different sources of lipids given in the same amounts (Deiuliis et al., 2010).
Reliable results have been obtained by measurements between duodenum and ileum or between
duodenum and faeces. A survey of sixty-four data points from the literature showed a wide
variation in intestinal digestibility, from 55 to 92 %. Intestinal digestibility does not depend on
FA intake, and the capacity of FA absorption in the dairy cow can be higher than 1 kg/d with
16
different sources of fat or with rapeseed oil infused in the duodenum within-experiment
comparisons and between experiment values for the same source of fat no clear effect of the FA
composition of the diet can be shown: for example, digestibility of soya bean oil varies between
70% (Jenkins, 2006).
Methodological errors do not explain these differences, since widely different values are often
found in the same experiment. A possible explanation is that micelle formation could depend
either on the proportions of the different FA, or on the production of biliary salts which is related
to the nature of FA. This hypothesis has not been assessed by experimental data but could
explain the observation that the replacement of rapeseed oil by hydrogenated fish oil decreases
the digestibility of all FA (Doreau, 1992). Digestibility depends on chain length but does not
differ between 16 and 18 C atoms: 79 and 77 % on average respectively. Moreover, digestibility
appears to be lower for C20 and C22 FA, but few results are available for these FA. From
literature data it can be calculated that the mean digestibilities of C 18 FA are 77, 85, 83 and 76
% for 0, 1, 2 and 3 double bonds (Doreau & Ferlay, 1994).
2.5. Fat Metabolism in Ruminants
2.5.1 Fatty acid synthesis
Microbial metabolism in the rumen also has consequences for fatty acid synthesis in ruminants,
as essentially all dietary carbohydrates are metabolized to short-chain fatty acids (acetate,
propionate and butyrate. Consequently the liver is committed to gluconeogenesis to make up for
the lack of dietary glucose, and fatty acid synthesis is very low in this tissue. Adipose tissue is
the major site of fatty acid synthesis in ruminants except during lactation when, the mammary
gland becomes the predominant site. Also, acetate rather than glucose is the major precursor for
fatty acid synthesis in ruminant tissues, although there is a requirement for glucose to provide
some NADPH via the pentose phosphate cycle for fatty acid synthesis, while glucose is the only
source of glycerol-3-phosphate in adipose tissue and the major source in the mammary gland for
fatty acid esterification (Jenkins, 2006)..
The major control point of fatty acid synthesis in both ruminants and non-ruminants is acetyl-
CoA carboxylase and it seems that, in adipose tissue at least, the endocrine control is very similar
17
(activation by insulin, inhibition by catecholamine’s) in ruminants and in non-ruminants. In
contrast, in the mammary gland insulin modulates acetyl CoA carboxylase in the rat but has no
effect on fatty acid synthesis in the ruminant. Fatty acids, or at least their acyl CoA esters, inhibit
acetyl CoA carboxylase in adipose tissue and mammary gland from both ruminants and non-
ruminants, but as yet no hormone has been identified which acts directly on the mammary gland
to inhibit fatty acid synthesis in the short term in any species.
Research on lipid metabolism in farm animals in the last 35 y has been focused on studying the
anatomical sites and main carbon and hydrogen sources for lipogenesis in different animal
species. From use of radioactively labeled precursors with in vivo and in vitro assays, it was
found that adipose tissue is the primary anatomical site for fatty acid synthesis in non-lactating
ruminants. Glucose and acetate are the main carbon sources for fatty acid synthesis in porcine
and ruminant adipose tissues, respectively. The substrate specificity can differ with depot site, as
is the case in intramuscular adipose tissue of beef cattle, where glucose rather than acetate seems
to be the primary substrate for fatty acid synthesis.
Glucose was a poor precursor for long-chain fatty acids because of very low activities of citrate
cleavage enzyme (ATP:citrate lyase) and malic enzyme (NADPH-malate dehydrogenase) in
ruminant lipogenic tissues in comparison to those in rats. Therefore, acetate and not glucose was
considered the principal generator of cytosolic acetyl-CoA for de novo fatty acid synthesis,
which occurs in the cytosol. Low rates of glycolysis (glucose → pyruvate) explain the low rates
of glucose use for lipogenesis in ruminant tissues because lactate conversion to cytosolic acetyl-
CoA, which requires citrate cleavage enzyme activity, is greater than glucose use and similar to
that for acetate. Three major pathways can generate cytosolic NADPH for fatty acid synthesis:
the pentose phosphate, malate dehydrogenase, and isocitrate dehydrogenase pathways (Jenkins,
2006)..
Reports suggest that in ruminants, as much as 50–80% of NADPH needed for fatty acid
synthesis in adipose tissue is produced by glucose oxidation via the pentose phosphate pathway.
The rest of the NADPH is generated via the isocitrate dehydrogenase pathway, which can
provide up to 50% of NADPH needed for fatty acid synthesis in the mammary gland. The
significance of the isocitrate dehydrogenase pathway is that it can provide reducing equivalents
18
for fatty acid synthesis from acetate in the absence of glucose oxidation. The contribution of the
malate dehydrogenase pathway to NADPH production in ruminants is insignificant because of
very low malic enzyme activity in ruminants.
2.5.2 Lipolysis
The triacylglycerols of adipose tissue are hydrolysed by hormone-sensitive lipase. This enzyme
is activated by catecholamines and inhibited by insulin; these properties are common to the
enzyme of both ruminant and non-ruminant adipose tissues. Triacylglycerols in both the
mammary gland and adipose tissue are synthesized using fatty acids derived from the
triacylglycerols of plasma chylomicrons. Increased availability of dietary fatty acids suppresses
de novo synthesis, probably by inhibition of acetyl CoA carboxylase (Palmquist, 1984). This
results in some variation in the fatty acid composition of the triacylglycerols, especially in milk
where the proportion of short- and medium-chain fatty acids is reduced (Jenkins, 2006)..
Increasing the proportion of polyunsaturated fatty acids in these triacylglycerols, however,
depends on limiting biohydrogenation of dietary lipids. Giving lipid supplements rich in
polyunsaturated fatty acids results in some escaping hydrogenation and a marginal increase in
the amount of polyunsaturated fatty acids in milk fat, but a substantial increase is dependent on
protecting the lipids (by coating with casein and treating with formaldehyde) against
biohydrogenation. Until recently, research on milk-fat production has been directed at increasing
the amount of milk fat. Diets rich in concentrates are known to tend to depress the milk-fat
concentration with a concomitant increase in fat deposition in adipose tissue (Deiuliis et al.,
2010).
Giving dietary supplements of cod-liver oil also decreases milk-fat production in cattle;
decreased fatty acid synthesis is partly, but not entirely, responsible. Several approaches are
being developed to reduce the amount of adipose-tissue triacylglycerol in farm animals.
Administration of P-agonists such as clenbuterol lead to a diminution in the amount of adipose-
tissue fat. P-Agonists inhibit fatty acid synthesis and stimulate lipolysis in adipose tissue,
probably by the same mechanism as catecholamines (activation of adenylate cyclase and cyclic
AMP-dependent protein kinase leading to phosphorylation and activation of hormone-sensitive
lipase and phosphorylation and inactivation of acetyl CoA carboxylase).
19
Growth hormone also inhibits fatty acid synthesis and promotes lipolysis in adipose tissue of
ruminants but the mechanism is unknown. Activation of adenylate cyclase does not appear to be
involved (Jenkins, 2006). Rather the hormone appears to act as an insulin-antagonist with respect
to activation of acetyl-CoA carboxylase and to increase the responsiveness of the tissue to the
lipolytic effect of catecholamines (McCutcheon & Bauman, 1986). Another approach to
diminish adiposity is to inject antibodies to the adipocyte plasma membrane (Futter & Flint,
1986). This differs from the approaches described previously in that the effect results from the
destruction of adipocytes rather than the manipulation of lipid metabolism; consequently it
provides a potential means of permanently reducing adiposity (Deiuliis et al., 2010).
20
3. CONCLUSIONS
Participatory range land management requires a comprehensive participatory approach,
involving pastoral communities, socio-economists, policy makers, administrators and other
technologists in all phases of planning, implementation and evaluation of activities. In Africa
there are various institutions working on pastoral resource management that perceive the lack of
a comprehensive national policy on rangelands. This should be considered as the first step to
solve this problem. To improve participatory rangeland management for sustainable
development and improve welfare of the local communities, there is a need to increase
production and productivity of rangeland through increased off-take of livestock and livestock
products and increase the income of pastoralists.
In the light of institutional problems among pastoral communities, a participatory approach
involving local communities is recommended in determining the best opportunities available for
the management of rangelands. There is a need to strengthen the management capacity of
rangelands through measures that enhance pastoralists’ control over natural resources. The rights
of pastoralists over rangeland resources must be legally recognized, thus ensuring security of
ownership. Non-Governmental organizations, government and pastoralists working together can
have a positive impact on sustainable use of rangeland resources.
21
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