Overview of
Rumination
The rumen and reticulum (reticulo-rumen)
is the largest compartment of the cow’s stomach. Fiber, starch, sugar, and
proteins are fermented by the microbes to form volatile fatty acids and
microbial protein. Rumen papillae absorb volatile fatty acids. The omasum has
many “leaves” of tissue that absorb water and serve as a filter. The abomasum
is the true stomach where the cow’s own enzymes and acids break down ingested
feed. Rumination is the process by which the cow regurgitates previously
consumed feed and masticates it a second time.
The
ruminant stomach has four compartments: rumen, reticulum, omasum, andabomasum. Ingested feed flows through the rumen and
reticulum first, then through the omasum, and lastly, through the abomasum.
1.
Rumen & Reticulum (also called the “Reticulo-rumen” or simply the “Rumen”):
The capacity of an
adult dairy cow’s rumen is about 184 litres (49 gallons) and the reticulum is
about 16 litres (4.25 gallons). It is one of the most dense microbial habitats
in the world. Microscopic organisms called rumen microbes break down (or
digest) ingested feed by a fermentation process. The rumen is the major site of
fermentation in the cow. There are1,000,000,000 to 10,000,000,000 bacteria per
ml, 1,000,000 protozoa per ml, and a variable amount of fungi. The cow does not
secrete any of her own acids or digestive enzymes in the rumen. Rather, all
rumen digestion is done by the microbes.
It is because of the
rumen and its microbes that dairy cows can digest plant fibers, such as
hemicellulose and cellulose. This allows the dairy cow to convert forages and
industrial byproducts that humans cannot digest into nutritious foods for
humans. Mammalian enzymes and acids cannot digest fiber. The rumen microbes
ferment fiber, starch, sugar, and protein to form volatile fatty acids and
microbial protein.
The
rumen wall (or mucosa) is a major site of nutrient absorption. It is convoluted
to give it tremendous surface area for absorption. Rumen papillae (thousands of finger-like projections
on the inside surface of the rumen) are responsible for absorbing the volatile
fatty acids from the rumen for use by the cow. Volatile
fatty acids (VFA’s) are absorbed from the rumen and used as an
energy source for the cow. Microbial protein(the
actual bodies of the microbes) is not absorbed from the rumen. It absorbed from
the cow’s intestine and is used as a source of quality protein for the cow.
The reticulum is
located in front of the rumen and it is known for its characteristic
“honeycomb” surface. If a cow accidentally eats hardware, such as screws or
nails, it usually ends up lodged at the bottom of the reticulum. If the
hardware punctures through the reticulum wall, it causes the often deadly
“hardware disease”.
2.
Omasum
The omasum is the
third stomach compartment. It holds only about 8 liters (2 gallons) in the
adult cow. It is made of many “leaves” of tissue which serve to absorb water
for the cow. These leaves also work as a filtration system for the cow to only
allow fine particles of digested feed and some fluid to pass on to the
abomasum.
3.
Abomasum
The abomasum is the
fourth stomach compartment. It is also called the “true stomach”. It holds
about 27 liters (7 gallons). This compartment has basically the same function
as the stomach in simple stomached (monogastric) animals, such as pigs and
humans. It is here that the cow’s own stomach acids and enzymes are used to
further breakdown ingested feed before it passes into the small intestine.
Characteristics of the
Rumen Environment
A number of
characteristics about the rumen allow for the growth of the rumen microbes and
for their fermentation process that digests feed and forages for the cow. Any
disruption to the delicate balance of the rumen environment will result in poor
growth of the microbes, poor digestion, and ultimately, lower milk production.
First
of all, the rumen is anaerobic. This
means that there is little or no oxygen in it. The rumen microbes cannot grow
in outside air. They will tolerate a small amount of oxygen so long as the
fermentation is going strong enough so that they can get rid of the oxygen
quickly. Some oxygen does, of course, get into the rumen through feed and
water. Secondly, the rumen temperature is
one degree above body temperature at39°C (102.5°F).
Luckily, it’s fairly easy for us as farmers and nutritionists to maintain an
anaerobic, warm environment deep in the belly of the cow!
Rumen
pH ranges between 5.7 and 7.3. The high side of this pH range (> 7) will be
seen on poor quality forage diets supplemented with urea. In high-producing
dairy cows,acidosis (rumen pH<6.0) is a common problem.
This occurs when the cow eats too much rapidly digestible starch or sugar that
creates acid and overwhelms the rumen’s buffering system. Most of the buffer in
the rumen comes in the form of saliva that is generated when the cow chews her
cud. Inadequate intake of long fiber that promotes rumination (cud-chewing) can
also result in acidosis because it provides less salivary buffer to counteract
the acid produced by grain fermentation. The rumen microbes, especially those
that primarily digest fiber, are acid intolerant. They do not grow well in acid
and they don’t digest feed, especially forages, well under acid conditions.
End Products of Rumen
Fermentation
1.
Volatile Fatty Acids (VFA’s):
The
rumen microbes make three primary volatile fatty acids: acetate (CH3COOH),propionate (C2H5COOH),
and butyrate (C3H7COOH). Acetate is formed primarily
from the fermentation of fiber. Large amounts of propionate are formed from
grain fermentation. Butyrate is produced to a lesser extent than acetate and
propionate. Sometimes lactate is also
formed, especially under acidic conditions in the rumen. VFA are actually waste
products from the rumen microbes but the cow absorbs them from her rumen and
uses them as major source of energy.
2.
Gases:
Dairy
cows produce 30-50 liters (8-13 gallons) of gas per hour. Carbon dioxide (CO2) (about 60%) and methane (CH4) (about 40%) are the main waste gases produced by
the rumen microbes. Hydrogen (H2) usually makes
up less than 0.05% of the total rumen gases. Gas is primarily found on top of
the solid and liquid contents of the rumen. Methane bacteria actually grow on
CO2 and H2 and other
byproducts from the other bacteria in the rumen. The methane bacteria commonly
produce methane in this way:
4H2 + CO2 ---> CH4 + 2H2O
Cows
get rid of these excess gases by a process called eructation (a kind of silent belching) and by
absorption through the rumen wall and lung exhalation. The cow eructates using
a process which is slightly different from the normal contractions of the
reticulum. Once gas pressure stimulates pressure receptors, the
reticulo-ruminal fold and the cranial pillar stop the rumen digesta from
getting into the oesophagus. The cardia and diaphragmatic sphincters relax, gas
enters the oesophagus and is released through the cow’s mouth.
If
a cow is unable to get rid of excess gases, she can bloat. When the rumen contents foam excessively, the
cow has difficulty getting rid of the gas because it is trapped inside the
foam. The rumen keeps expanding. It creates pressure on the cow’s lungs and the
cow can die of asphyxiation. Many legumes (white clover and alfalfa),
especially those that are young and freshly cut, can cause bloat. A chronic
type of bloat also occurs in beef cattle on feedlots when they are fed a lot of
grain. Oils or low-foam detergents have been used to reduce the stability of
foam in the rumen and prevent bloat.
3.
Microbial Mass
The rumen microbes
grow in the rumen and their bodies are passed down to the intestine of the cow.
Thus, they make up a large portion of the cow’s diet and the largest portion of
the cow’s protein supply. This microbial protein is high quality, meaning that
the amino acid profile is fairly similar to that of milk and meat. Therefore,
the cow can easily and efficiently convert microbial protein into milk and
meat.
4. Heat
As
the microbes digest ingested feed and forages and grow, they release heat. This
is called the “heat of fermentation”. Except for when the cow needs this heat
to warm her body, this heat is a waste of energy to the cow. The fermentation
of forages creates more heat than the fermentation of concentrate feeds.
Rumen Structure and Movements:
Rumination
Rumination,
also called cud-chewing, is the process by which the cow regurgitates (casts
up) previously consumed feed and masticates it a second time. The re-chewed
feed with saliva is formed into a bolus and swallowed a second time. It is the
floating, large particles on top of the rumen which are re-chewed. One purpose
of rumination is to decrease particle size and increase surface area of the
feed. This results in an increase in digestion rate of feed and a decrease in
the lag time prior to fermentation. The fibrous particles will stay in the
rumen longer causing the rumen to feel more full if the cow is not ruminating
enough. This will reduce the total intake of the cow and negatively impact milk
production. Another purpose of rumination is to make saliva (98 to 190 liters
(or 26-50 gallons) per day) to buffer the rumen and decrease rumen acidity.
Cows
usually spend more time chewing during rumination than they do when they eat.
Pressure of coarse material (or “scratch factor”) against the rumen wall stimulates
the cow to ruminate. Therefore, the amount of time a cow spends ruminating is
diet dependent. Feeding a lot of concentrates and/or finely ground feeds
reduces rumination. It is critical, especially in high-producing dairy cows
that consume considerable amounts of concentrate, that there is an adequate
amount of long fiber present in the diet to stimulate rumination.
Stratification of Rumen Contents
Dairy
cows who are fed a diet containing long fiber have a large, dense, floating
layer (or floating mat) located just under the gas in the top portion of the
rumen. This mat contains the more recently consumed feed, especially the
fibrous portion of the diet. As the ingested feed ferments and is ruminated, it
becomes more water-logged and dense and gradually sinks toward the bottom of
the rumen. Smaller, dense particles and liquid is located in the bottom portion
of the rumen. This bottom layer is material that is nearly ready for passage
out of the rumen, into the omasum, and on down the cow’s digestive tract. Diets
high in concentrates or containing only very fine fibrous material may either
have no floating mat or a very small one.
Rumen Mixing
Mixing
of rumen contents is very important to the cow. Contraction and relaxation of
the entire rumen, especially the wall between the rumen and the reticulum,
helps to mix contents. Also, ridged structures called rumen pillars act like
baffles on the inside of a clothes washing machine and help to set up mixing
motion in the rumen.
Rumen
mixing helps to inoculate feed with microbes. It also aids VFA in reaching the
rumen wall for absorption. The combination of mixing and rumination allows the
cow to pass indigestible matter that might otherwise stay in the rumen and
decrease total feed intake of the cow. It is interesting to note that finely
processed diets that result in little mat formation actually increase the rate
at which feed passes and increase the size of particles found in the manure.
Adding
just a small amount of long hay to a diet, such as 1-2 kg (2-4 pounds) can
greatly increase the size of the mat, stimulate rumen mixing, help VFA to reach
the rumen wall for absorption and help to properly sort rumen contents. At same
time, hay promotes rumination and the production of saliva to buffer the rumen.
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