Fisiologi Nutrisi kuda

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

2. The types of substrate and fermentation patterns are

essentially identical for forestomach and hindgut
fermentation

3. The motility functions of the cecum and colon retain

material for fermentation and separate particles by size

4. The rate of fermentation and volatile fatty-acid

production in the equine colon is similar to that in the
rumen
INTRODUCTION
Horses have a simple stomach.
The stomach and small intestine are similar to other
monogastric species.
However, the equine large intestine is massive and
anatomically complex in comparison to most other
animals.
The small intestine empties ingesta into the cecum
through the ileocecal orifice. The cecum also houses
the cecocolic orifice, from which its contents flow into
the colon. The colon itself is huge and, by folding on
itself several times, is divided into segments and
subsegments.
Diagram of
horse GI
THE EQUINE COLON
ascending colon (the "great colon"; by far the largest)
right ventral colon
left ventral colon
left dorsal colon
right dorsal colon
transverse colon
descending colon (small colon)
The cecum and ascending colon have bands of smooth muscle
which cause these organs to form pouches called haustrae. T
The descending colon becomes the rectum at the inlet of the
pelvis.
Some anatomists simplify the segments of the equine large gut
into cecum, ventral colon, dorsal colon and small colon.
Fermentation and Physiology
of the Equine Hindgut
Digestive function in the stomach and small intestine of
horses as in any other monogastric animal.
Dietary protein is digested and absorbed as amino acids and
much of the soluble carbohydrate is hydrolyzed and absorbed as
monosaccharides in the small gut.
Cellulose and related molecules pass through the small gut intact,
although such plant material may be softened and swollen prior to
entry into the cecum.
Structural carbohydrates like cellulose and hemicellulose, along
with starch and other soluble carbohydrates that escape digestion
in the small intestine, flow into the large intestine where it is
subjected to fermentation.
Fermentation and Physiology
of the Equine Hindgut
The large intestine of horses and other hindgut
fermenters is a fermentation system analagous to the
rumen.
The process of fermentation that occurs in the hindgut
is essentially identical to that which occurs in the
forestomachs of ruminants.
Most importantly, horses survive as herbivores because
volatile fatty acids are produced in large quantities,
absorbed through the cecal and colonic epithelium, and
distributed for use throughout the body.
One significant difference from the ruminant strategy is that
that large quantity of microbial protein generated in the
equine large gut is wasted because there is no opportunity
for significant absorption of amino acids.
 The equine gut
The hindgut commences with
the cecum, separated from the
large colon by a well-defined
orifice.

The large colon is folded on

itself three times, forming four
major anatomic divisions:

right and left ventral colon

left and right dorsal colon
segments.

Ingesta enter the right ventral

colon, to the left ventral colon,
from which the material enters
From the left dorsal colon, material moves to the
right dorsal colon before entering the small colon.

Note the tremendous size and volume of the cecum

and colon compared with the small intestine.

The differences in diameter that occur throughout

the colon also occur at the pelvic flexure and at the
junction of the large and small colons.

The sac-like evaginations that occur in the wall of

the cecum and most segments of the colon are
called haustra.

Functionally, the equine hindgut can be divided into

four sections: cecum, ventral colon, dorsal colon,
and small colon.
The equine hindgut has a tremendous
capacity for fermentation

A general function of the cecum and colon is to

recover fluid and electrolytes from ingesta
leaving the ileum.

In many herbivorous species, this function has

been expanded to include fermentative
digestion. Absorptive and fermentation
functions complement each other in the colons
of non-ruminant herbivores, like horses.
 The types of substrate and fermentation patterns are essentially
identical for forestomach and hindgut fermentation
Hindgut fermentation may be aided by prior gastric action, eg acid
exposure.
Some sugars and starches may be digested and absorbed before
material arrives in the cecum.
Substantial amounts of starch and sugars reach the cecum. Even
with a high-grain diet, up to 29% of dietary starch may reach the
cecum and colon.
Protein as well as some carbohydrate, is absorbed in the small
intestine, potentially leading to a deficiency of nitrogen for colonic
microbes. However, there is extensive urea recycling into the colon
and cecum similar to the rumen (see Fig. 30-6).
Thus, urea plus protein escaping small intestinal digestion supplies
the nitrogen needs of the microbes.
In contrast to ruminants, horses do not have an efficient means of
recovering the microbial protein synthesized in the hindgut, and
most of it passes out in the feces.
The motility functions of the cecum
and colon retain material for
fermentation and separate particles by
size
The functions of the equine hindgut in maintaining fermentation are
similar to those of the rumen: favorable conditions must be
maintained to support optimal fermentation. As in the rumen, these
conditions are:
(1) substrate supply
(2) control of pH and osmolality
(3) anaerobiosis
(4) retention of fermenting material
(5) continual removal of waste products and the residue of spent
fermentation substrate. Separation of fermenting material from
residue appears to be accomplished by selective retention of particles
according to size, just as in the rumen; however, the
means by which the cecum and colon accomplish
MOTILITY
A large portion of soluble ingesta reaches the cecum by 2 hours after
ingestion, whereas solids take longer.
The material in the cecum has a high water content and a slurry-like
consistency. Most cecal motility is of a mixing nature, with frequent
low-amplitude contractions that transport ingesta from haustrum to
haustrum and back in a mixing pattern. The mixing action of the
cecum maintains the cecal contents in a homogeneous state.
About once every 3 to 4 minutes, there is a strong contraction of cecal
muscles in a mass-movement type of action in which the body and
apex of the organ shorten and constrict, lifting ingesta into the base.
Constriction of the base forces material through the cecocolic orifice
and into the right ventral colon. The motility pattern functionally
separates the cecum from the ventral colon; there is no retrograde
flow of material from the colon to the cecum.
 Three types of motility patterns exist in the right and left
ventral colon: haustral segmentation, propulsive
peristalsis, and retropulsive peristalsis.
Segmentation serves a mixing function that aids in
promoting fermentation and bringing VFAs in contact
with the mucosa for absorption.
Propulsive activity in the ventral colon originates near the
cecum as a continuation of the cecal mass movements.
Peristaltic activity in the proximal ventral colon propels
ingesta distally into the left ventral colon. In the left
ventral colon, retropulsive or anti- peristaltic movements
are encountered, which resist the flow of ingesta and
result in the retention of
material in the ventral colon, allowing time for microbial
digestion and preventing the washout of microbes.
Large Intestinal Motility
Motility in the equine hindgut provides the same fundamental functions as in the
large intestine of other animals: mixing, retention and propulsion of ingesta.

Motility in the cecum consists of mixing contractions in which the haustra

alternately contract and expand. Additionally, every few minutes the a strong,
mass movement-type contraction occurs that forces some of the cecal contents
through the cecocolic orifice into the ascending colon.

Segmentation and haustral contractions occur within the ascending colon

occurs that efficiently mix ingesta and expose it to the mucosa for:

Absorption of water, electrolytes and volatile fatty acids produced through

fermentation.

There are peristaltic contractions that "fight" with antiperistaltic contractions ,

leading to additional mixing and an overall transit rate that is rather slow (it takes 2-
3 days to traverse to the colon). Within the small colon, the predominant patterns of
motility are peristalsis and segmentation - segmentation contractions assist in
formation of the fecal balls characteristically observed in horses.
MOTILITY
In the distal ventral colon, antiperistaltic activity and the
narrow diameter of the pelvic flexure retard the
movement of material, causing it to be retained in the
ventral colon.
The squeezing action of the pelvic flexure selectively
retains large particulate matter; liquid and small
particles to pass on.
As particle size is reduced by fermentative action and
the mixing activity of the colon, particles eventually
become small enough to flow with the fluid phase and
leave the colon.
Some particulate matter enters into the left dorsal
colon.
MOTILITY
The actions of the dorsal colon are like those
of the ventral colon. Impedance to ingesta
flow is created by the size restriction at the
junction of the right dorsal colon and small
colon. In addition, there may be retropulsive
motility originating in the area of the distal
right dorsal colon, near the junction with the
small colon.
These actions tend to impede the movement
of ingesta through the dorsal colon, subjecting
the material to another round of fermentative
digestion, as occurred in the ventral colon.
The delay in the flow of ingesta created by the
combined actions of the ventral and dorsal
colons results in significant retention of
material, with most particulate matter taking
from 24 to 96 hours to pass the large.
Understanding the motility of the equine colon
is important, because problems of colon
impaction in horses are common. Impactions
usually occur near or within the pelvic flexure,
probably because the pelvic flexure is a site of
flow restriction and differential flow of solid
and liquid material.
WATER FLUX
Large fluxes of water traverse the cecal and colonic
mucosa during the course of digestion. Feed starts to
enter the cecum about 2 hours after eating, and VFA
production rapidly commences.
As ingesta are transported from the cecum, VFA
production continues in the large colon.
During the period of active VFA production, large
quantities of water enter the hindgut from the blood
through the mucosa.
Water flux iis likely a response to direct fluid secretion
from the crypts of the colonic epithelium.
Secretion of sodium-/ bicarbonate-, and chloride-
containing fluid from the colonic mucosa appears to
occur in response to high concentrations of VFA in the
lumen.
This secretory response, in combination with the ileal
secretions, is responsible for buffering of the lumen
contents.
This figure illustrates the magnitude of water fluxes
that occur during hindgut digestion in the pony.
Note that there is considerable inward and outward
movement of water across the mucosa in each of the
major fermentation compartments, ventral and dorsal
colons, and cecum.
Inward (into the lumen) water movement results from
mucosal secretion, whereas outward water movement
occurs in association with absorption of VFA.
FERMENTATION
The rate of fermentation and volatile fatty acid production in the
equine colon is similar to that in the rumen.
In the equine colon, efficient means of buffering and VFA
absorption must be present.
Salivary buffering, as occurs in ruminants, cannot aid in buffering
the colon, because of the changes in ingesta pH that occur during
transit through the stomach and small intestine.
Therefore, in the horse, large quantities of fluid, rich in
bicarbonate and phosphate buffers, are secreted by the ileum and
transferred to the cecum.
There is also direct addition of bicarbonate and other electrolytes
by glands of the colonic mucosa.
FERMENTATION
STRATEGIES
Among the animals that use fermentation, two distinct strategies
have evolved. The chief difference between the two groups is in
positioning of their fermentation vat relative to the stomach and
small intestine:

Cranial fermentors or ruminants have a large, multi-compartmented

section of the digestive tract between the esophagus and true stomach.
These forestomachs house a very complex ecosystem that supports
fermentation. Examples of ruminants are cattle, sheep and deer.

Caudal fermentors, also known as cecal digestors, are similar to dogs

and humans through the stomach and small intestine, but their large
intestine, where fermentation occurs, is complex and exceptionally
large. Examples of cecal digestors include horses and rabbits.
 The process and outcome of fermentation
in the rumen of a cow and the cecum of a
horse is essentially identical .
However, the positioning of the fermentation
vat in relation to the small intestine has
implications for physiology and nutrition.
These similarities and differences are
summarized next.
Function Ruminants Cecal
Digestors
Digest and extract
energy from cellulose Yes Yes

Utilize dietary sugars

sources directly No Yes

Utilize the protein

from fermentative
Microbes Yes No
FERMENTATION
STATEGIES
In monogastrics animals, the small intestine is the only site in the
digestive tract where simple sugars and amino acids can be
absorbed.
Ruminants can utilize dietary starch, but very little of it is
absorbed as glucose.
Instead, in ruminants, starch and other soluble carbohydrates are
fermented to volatile fatty acids in the forestomachs.
What little starch enters the small intestine is poorly digested here
due to a relative deficiency in amylase.
In contrast, starch fed to a horse is digested to glucose by
amylase and maltase in the small intestine, and that glucose
is absorbed across the epithelium into blood.
 The bodies of microbes in the fermentation vat
represent a large source of high quality protein.
In ruminants, those microbes flow into the stomach and
small intestine, where they are digested and absorbed
as amino acids and small peptides.
Because the fermentation vat of a horse is behind the
small intestine, all their microbial protein is lost.
Horses absorb dietary protein before fermentation, i.e.
in the stomach/small intestine,
 Both ruminants and cecal digestors
are very successful in evolution.
Each of these two strategies
predispose these groups of animals to
distinctive diseases.
INTRODUCTION TO
MICROBIAL
FERMENTATION
Fermentation is supported by a rich and dense collection of microbes. Each milliliter of rumen
content contains roughly 10 to 50 billion bacteria, 1 million protozoa and variable numbers of
yeasts and fungi.
The environment of the rumen and large intestine is anaerobic and, as expected, almost all these
microbes are anaerobes or facultative anaerobes.
Fermentative microbes interact and support one another in a complex food web, with the
waste products of some species serving as nutrients for other species.
Fermentative bacteria representing many genera provide a comprehensive battery of
digestive capabilities.
These organisms are often classified by their substrate preferences or the end products they
produce. Although there is some specialization, many bacteria utilize multiple substrates.
Some of the major groups, each of which contain multiple genera and species, are:
Cellulolytic (digest cellulose)
Hemicellulolytic (digest hemicellulose)
Amylolytic (digest starch)
Proteolytic (digest proteins)
Sugar utilizing (utilize monosaccharides and disaccharides)
Acid utilizing (utilize such substrates as lactic, succinic and malic acids)
Ammonia producers
Vitamin synthesizers
Methane producers
 Protozoa, predominantly ciliates, appear to contribute substantially to the fermentation
process. Several experiments have demonstrated that young ruminants deprived of their
ruminal protozoa show depressed growth rates and are relative "poor-doers" compared to
controls with both bacteria and protozoa.
In general, protozoa utilize the same set of substrates as bacteria and, as with bacteria,
different populations of protozoa show distinctive substrate preferences. Many utilize simple
sugars and some store ingested carbohydrate as glycogen.
An interesting feature of some protozoa is their inability to regulate glycogen synthesis: when
soluble carbohydrates are in abundance, they continue to store glycogen until they burst. An
additional feature of protozoa is that many species consume bacteria, which is thought to
perhaps play a role in limiting bacterial overgrowth.
The distribution of microbial species varies with diet. Some of this appears to reflect
substrate availability; for example, populations of cellulolytic microorganisms are depressed in
animals fed diets rich in grain.
Environmental conditions in the fermentation vat also can have profound effects on the
microbial flora. Rumen fluid normally has a pH between 6 and 7, but may fall if large amounts
of soluble carbohydrate are consumed. If pH drops to about 5.5, protozoal populations
become markedly depressed due to acid intolerance. More drastic lowering of rumen pH, as
can occur with grain overload, can destroy many species and have serious consequences to
the animal.
 The forestomach of ruminants and large intestine of caudal fermenters are
continuous flow fermentation systems containing enormous numbers of microbes.
What do these microbes and the process of fermentation provide the
herbivore? Apart from fermentation, the microbes that digest cellulose and
other substrates also provide at least three other major services:
Synthesis of high quality protein in the form of microbial bodies. Caudal
fermentors cannot take advantage of this service, but in ruminants, bacteria and
protozoa are constantly flowing into the abomasum and small intestine, where they
are digested and absorbed. All vertebrates require certain amino acids which their
cells cannot synthesize (the "essential amino acids"). Fermentative microbes can
synthesize all the amino acids and thereby provide them to their host.
Synthesis of protein from non-protein nitrogen sources. Fermentative
microbes can, for example, utilize urea to synthesize protein. In some situations,
ruminants are fed urea as a inexpensive dietary supplement. They also secrete
urea formed during protein metabolism into saliva, which flows into the rumen and
serves as another nitrogen source for the microbes.
Synthesis of B vitamins. Mammals can synthesize only two of the B vitamins and
require dietary sources of the others. Fermentative microbes are able to synthesize
all the B vitamins, and deficiency states are rarely encountered.
Substrates 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