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Digestion & Metabolism of Carbohydrate, Fat and Protein in Non-Ruminants
Digestion & Metabolism of Carbohydrate, Fat and Protein in Non-Ruminants
• Non-ruminants 80 to 120 mg %.
• Glycogen reserve is short lived. A 24-hour fast will reduce the levels
to nearly zero. Glycogen stores have to be constantly replenished.
Glucose to Fat
• The ability of the liver and other tissues to store sugar as glycogen is
limited.
• When the carbohydrate intake regularly exceeds the current need of the
body for energy purposes, sugar is transformed into fat .
• The formation of body fat from carbohydrate food was first demonstrated
by Lawes and Gilbert in 1859 by means of slaughter experiments.
Lactate dehydrogenase
NAD+
Lactate
• Lactate diffuses from the muscle cell into the blood, carried to the
liver. It is converted back to pyruvate by lactate dehydrogenase.
• Pepsin cleaves bonds adjacent to aromatic amino acids and splits the
polypeptide into large peptides and relatively few amino acids.
• In duodenum the ingesta come in contact with the pancreatic and bile
secretion.
• Neutral amino acids are absorbed rapidly than basic amino acids.
• Vitamin B6 also appears to greatly enhance intestinal transport of
amino acids.
• This competition disappears, when amino acids are absorbed as
oligopeptides.
• All essential amino acids are not absorbed with equal
efficiency,
• For the young animals receiving milk, the stomach is substantially more
important
• Protein reserves are not distinct like glycogen for carbohydrates and depot
• But protein reserves are available from practically all body tissues for the
• Liver is the key organ- that synthesizes proteins, supplies amino acids to the
excess.
• The nitrogenous products are digested and enter the blood stream
The amino acids in the blood constitute the amino acid pool.
Metabolism of amino acids
- decarboxylation.
• most fishes excrete ammonia directly through the gill tissue into their
water environment.
• In fowl, uric acid is synthesized, from glutamine, glycine and aspartic acid
with the help of xanthine oxidase.
• Urea is not synthesized in birds. The only urea in chicken urine comes from
the breakdown of dietary arginine .
• Note that both the amino groups found in the urea molecule came
from glutamate one from oxidative deamination and the other from
transamination
The following steps are involved in the urea cycle.
1) Formation of carbamoyl phosphate (CP). Enzyme required is
carbamoyl phosphate synthetase.
2) Formation of citrulline from CP and ornithine. Enzyme is
ornithine— f carbamoyl transferase.
3) Formation of argininosuccinate and arginine. A second amino group
is transferred from aspartic acid to carbamoyl keto group of
citrulline, to form the amino acid arginine. Enzymes needed are
argininosuccinate synthetase and lyase.
4) In the presence of an enzyme, arginase and magnesium, arginine
yields one molecule of urea and of ornithine.
• The regenerated ornithine can participate in the next turn of the
cycle. There is a net loss of 1 ATP per mole of urea synthesized from
glutamic acid
• Metabolic precursors of the nitrogen in the purine ring of uric acid-two of these
arise from the amide N of glutamine and the other two from the amine groups of
glycine and aspartic acid.
• The purines and pyrimidine's absorbed from the intestines can be used for the
synthesis of nucleotides and nucleic acids.
• Purine compounds which are degraded in the tissue are readily reused. When
excreted they are converted to uric acid in primates, to allontoin in most mammals,
and to urea in fish. Pyrimidine's are excreted as urea and ammonia
Fate of Carbon Skeletons of Amino
Acids
• The carbon skeletons of the amino acids which have lost their amino
groups enter the citric acid cycle.
• All the amino acid residues which do not enter at either acetyl CoA or
acetoacetate have the potential of being converted to glucose in the
process of gluconeogenesis. These are called glucogenic.
• Those entering at acetyl CoA or acetoacetate could provide ketones.
And are referred to as ketogenic.
• Some are referred to as both glucogenic and ketogenic
Catabolism of Tissue Protein and Amino Acids
• The protein mass of the body, like the adipose mass, is in a continuous
state of flux, with tissues constantly being catabolized and
resynthesized.
• As amino acids are released they become available to the general
amino acid pool and can either be reused for protein synthesis or
utilized as a source of energy.
• Two types of catabolism are observed-one a normal function of tissue
maintenance and renewal, and the other that follows periods of under
nutrition/starvation.
Why Should there be a Continuous
Turnover or Protein renewal
• As protein synthesis requires energy, it would appear that the
protein renewal to be wasteful.
• Yet this continuous turnover of tissue protein represents changing
environment.
• With a continuing protein turnover capability, the animal has the
means, flexibility and speed to adapt to certain subtle or gross changes
in its environment which, in the long run, may affect its ability to
survive.
• The rate of turnover is not similar for all tissues. Renewal of intestinal
mucosa is extremely rapid.
• During periods of under nutrition/starvation, tissues lose their protein
and release amino acids.
• About one fourth of the body protein especially liver, muscle are
depleted and repleted.
• In general the integrity of brain and kidney are maintained.
• Thus, due to this property, the vital functions may be protected up to
30-50 days of total starvation.
• During starvation or fasting, the primary need is energy, particularly
glucose. Met through alanine & glutamine
FAT-DIGESTION AND
METABOLISM
• Dietary lipids consumed include triglycerides primarily, phospholipids
and cholesterol
• Although gastric juice contains a lipase, it is essentially inactive
because of low pH of the stomach.
• This lipase may be much more important in the young ones where
the gastric pH is higher and the fat of milk is highly emulsified.
• The principal site of lipid digestion is the small intestine
• Under the influence of the peristaltic action of the stomach,
duodenum and presence of bile, fat exists in the duodenum as a coarse
emulsion of triglycerides.
• Pancreatic lipase (Ca++ ions and bile salts increase its activity)
and colipase hydrolyze the triglycerides into fatty acids and
monoglycerides and reduce the lipid to a finer and finer emulsion.
ABSORPTION
• After digestion fats are present in the small intestine in the solubilized form
of mixed micelles.
• Efficient absorption requires a rapid movement of the highly hydrophobic
molecule through the unstirred water layer adjacent to the mucosa.
• This is the rate limiting stage of absorption.
• The mixed bile salt micelles, with their hydrophilic groups, aid this
process.
• Absorption across the brush border membrane of the intestinal cells is by
passive diffusion and is at its maximum in the jejunum.
• The bile salts are absorbed by an active process in the distal ileum.
• Following absorption there is a resynthesize of triacylglycerol's,
• a process that requires energy, and they are formed into chylomicrons
(minute fat droplets), which then pass into the lacteals of the villi,
enter the thoracic duct and join the general circulation.
• Medium- and short-chain fatty acids, such as those occurring in
butterfat, require neither bile salts nor micelle formation
• as they can be absorbed very rapidly from the lumen of the intestine
directly into the portal bloodstream.
• The entry of these fatty acids is sodium-dependent and takes place
against a concentration gradient by active transport.
• In fowls, the lymphatic system is negligible and most of the fat is
transported in the portal blood as low-density lipoproteins.
• Depot fats are formed from ingested fats and carbohydrates.
• That is why the nature of the fat deposited can be markedly affected
by the character of its food source.
• The influence of the kind of the food fat upon the character of the
body fat is striking-iodine number of food fat is proportional to that of
body fat
• When the ration contains much unsaturated fatty acids in the form of
oils, the body fat is also soft, i.e. of low melting point (higher iodine
number)
• Adipose tissue is not static and there is a constant exchange of fatty
acids between the body fat and fat in the blood. Hence deposits of
soft fat can be modified by a change in diet.
• When a ration which will produce hard fat is given after a period on
feeds rich in unsaturated fat, the deposited fat gradually becomes
harder. This process is called "hardening off”,
• It is noted that rations containing cottonseed oil produce lard (pig fat)
graded as hard lard, and those rich in highly unsaturated fat e.g. maize
oil or soybean oil produce soft lard.
• The "hardening off" process is taken advantage of in feeding practice
in finishing pigs for market.
• Anderson and Mendel (1928) showed that the process takes place
more rapidly where the animal was fasted for a period before the
hardening ration was given.
• Catabolism of Fat and Fatty Acids
• The end result of catabolism of fats and fatty acids is the production
of ATP, CO2 and H2O with the liberation of excess heat.
• The initial degradation of fat leads to the formation of glycerol and
acetyl CoA.
• In the ruminant absorbed acetate, butyrate and ketone bodies
are also available tor immediate catabolism.
• Mobilization and Oxidation of Fat
• It is well known that between meals the blood free fatty acid levels
are elevated as a result of the mobilization of fat.
• Upon eating, they drop promptly.
• Triglycerides are released from the adipose tissue under
hormonal control which activates adenylyl cyclase which in turn
causes the synthesis of cyclic AMP .
• Triglyceride gives glycerol and fatty acid.
• β-Oxidation of Fatty Acids
• Fatty acids are combined with an albumin and circulate as an albumin
fatty acid complex.
• In the cell, fatty acid oxidation begins in the extra mitochondrial
cytoplasm with the formation of fatty acyl CoA (fatty acid + coenzyme
A -> fatty acyl CoA).
• Fatty acyl CoA needs a special carrier mechanism in the form of
carnitine to pass into the mitochondrion.
• Knoop proposed that fatty acids were oxidized physiologically by
β-oxidation.
• In the mitochondria, fatty acyl CoA is successively dehydrogenated,
hydrated, dehydrogenated again, and cleaved to acetyl CoA and a fatty
acid shorter by two carbon atoms.
• This process is continued stepwise each sequence
producing a molecule of acetyl CoA.
• Acetyl CoA enter the TCA cycle and oxidized to C02 + H20.
• The acetyl CoA that is formed here may also follow the below
mentioned fates.
- It may condense to form acetoacetate and ketone bodies.
- It may be converted to malonyl CoA as in fatty acid
synthesis, or
- It may react with acetoacetyl units in sterol synthesis.
Conversion of Fat into Glucose
• Animals cannot convert fatty acids into glucose.
• The two carbon atoms of the acetyl group of acetyl CoA enter the
citric acid cycle but two carbon atoms leave the cycle in the
decarboxylation's catalyzed by isocitrate dehydrogenase and a-
ketoglutarate dehydrogenase .
• Consequently, oxaloacetate is regenerated but it is not formed de-
novo when the acetyl unit of acetyl CoA is oxidized by the citric acid cycle.
• Anyway the glycerol moiety of fat can yield glucose and thus 'fat'
can contribute somewhat to the glucose pool.
Interrelations with Proteins and Carbohydrates
• The metabolism of fatty acids and that of carbohydrates and proteins are
intimately related.
• The constituents are in a constant state of flux. Even the structural
proteins, carbohydrates and storage lipids are constantly broken down
and rebuilt.
• The status of a particular animal is the net result between the rates of
synthesis and of breakdown of its body constituents.
• Glycerol is the only component of lipids that is involved in the synthesis of
carbohydrates.
• On the other hand, lipids can be formed from carbohydrate in many ways.
• Indeed, the metabolism of lipids, carbohydrates and proteins is
metabolically dynamic.
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