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Oleh:
Eko Widodo
INTRODUCTION
Horses and their relatives utilize cellulose
and other fermentable substrates in much
the same way as ruminants, but, lacking
forestomachs, perform fermentation in their
large intestine.
Certain other herbivores have also adopted
this "caudal fermentation" lifestyle, most
notably rabbits and rodents.
Function of the equine
large hindgut
1. The
equine hindgut has a tremendous capacity for
fermentation
With few exceptions, all dietary carbohydrates and proteins can serve as
substrates for microbial fermentation.
The crucial advantage of being a herbivore is the ability to efficiently extract
energy from cellulose and other components of plant cell walls.
Cellulose fibers account for 40-50% of the total dry weight of stems, leaves
and roots. These fibers are embedded in a matrix of hemicelluloses and phenolic
polymers (lignin-carbohydrate complexes) that are covalently crosslinked.
Cellulose itself is a linear polymer of glucose molecules linked to one another by
beta[1-4] glycosidic bonds and herein lies the problem for the vertebrate digestive
system. As far as is known, no enzyme able to hydrolyze beta[1-4] glycosidic
bonds has evolved in vertebrates.
However, a variety of such beta-glucanases are synthesized by microbes.
Thus, the diverse population of bacteria and protozoa in the rumen or
hindgut produce all the enzymes necessary to digest cellulose and
hemicellulose. The glucose released in this process is then taken up and
metabolized by the microbes, and the waste products of microbial
metabolism are passed on to the host animal. Sugars derived from digestion of
soluble carbohydrates such as starch are processed similarly.
The Products of
Fermentation
Fermentation occurs under anaerobic conditions. As a consequence, sugars
are metabolized predominantly to volatile fatty acids (VFAs). Additional major
products include lactic acid, carbon dioxide and methane.
The principle VFAs are acetic, proprionic and butyric acids, which
collectively provide for the majority of a herbivore's energy needs. The ratio of
these VFAs vary with diet, although the majority product is always acetate.
On a diet high in fiber, the molar ratio of acetic to proprionic to butyric acids is
roughly 70:20:10.
Proteins are also important substrates for fermentation.
In caudal fermenters, much of the dietary protein is digested and
absorbed prior to the large gut, but in ruminants, all dietary protein
enters the rumen. The bulk of this protein is digested by microbial proteases
and peptidases. The resulting peptides and amino acids are taken up by
microbes and used in several ways, including microbial protein synthesis.
However, a large quantity of amino acids ingested by fermentative microbes
are deaminated and enter some of the same pathways used for carbohydrate
metabolism. The net result is that much of dietary protein is metabolized to
VFAs.
VFAs
VFAs are the MOST important product of fermentation. These small lipids
are used for many purposes, but the importance of VFAs to herbivores is
that they are absorbed and serve as the animal's major fuel for
energy production, serving much the same function that glucose does in
monogastrics.
The three major VFAs absorbed from the rumen have somewhat
distinctive metabolic fates:
Acetic acid is utilized minimally in the liver, and is oxidized throughout
most of the body to generate ATP. Another important use of acetate is as
the major source of acetyl CoA for synthesis of lipids.
Proprionic acid is almost completely removed from portal blood by the
liver. Within the liver, propionate serves as a major substrate for
gluconeogenesis, which is absolutely critical to the ruminant because
almost no glucose reaches the small intestine for absorption.
Butyric acid, most of which comes out of the rumen as the ketone beta-
hydroxybutyric acid, is oxidized in many tissues for energy production.
What does this mean in
terms of amounts?
Consider a dairy goat weighing about 200 lb and producing 1570
kg of milk in a 305 day lactation.
Her milk was roughly 4% lactose, 3.5% protein and 3.6% fat.
This means that, for the sole task of producing milk, this goat has to
synthesize about 250 grams of lactose and 180 grams of protein and
185 grams of fat every day.
Essentially all the glucose in that lactose is synthesized in the liver
and most of that synthesis is from proprionic acid generated by
fermentation.
Likewise, much of the fat is synthesized from ruminal acetate. When
you consider that synthesis of lactose and milk fat are only two of
many, many processes that are supported by volatile fatty acids, the
process of fermentation in herbivores gains new meaning.
Quote for the Day
I think we're on the
road to coming up
with answers that I
don't think any of us
in total feel we have
the answers to.
Kim Anderson, mayor
of Naples, Florida
Horse Terms
FOAL <1 yr
YEARLING >1yr
LONG YEARLING >1 yr
FILLY <3 to 4 yr
COLT <3 to 4 yr
MARE >4 to 5 yr
STALLION > 4 to 5 yr
GELDING castrated male