R uminations

Questions & Answers

and other thoughts on the role of
rumen microbes in dairy cattle

Written by:
James B. Russell
Produced and distributed by:
U.S. Dairy Forage
Research Center
USDA Agricultural Research Service
Madison, WI
R uminations
Questions & Answers
and other thoughts on the role of
rumen microbes in dairy cattle
by James B. Russell, PhD

About the Author:

James B. Russell was a Research Microbiologist with the USDA
Agricultural Research Service until his untimely death in
September of 2009. His laboratory was located in Ithaca, NY
and he was associated with the U.S. Dairy Forage Research
Center (USDA-ARS) in Madison, WI. He joined the ARS in 1981.
Dr. Russell was widely recognized as a leading authority on
ruminal microbiology. In 2004 he was promoted to Super
Grade by the ARS; less than 1% of ARSs Category I scientists
are promoted to this rank.
He received a B.S. Degree from Cornell University and M.S. and
Ph.D. degrees from the University of California at Davis. James B. Russell
1951 to 2009

Produced and distributed by:

U.S. Dairy Forage Research Center
USDA-Agricultural Research Service
1925 Linden Dr. West, Madison, WI 53706; phone 608-890-0050
www.ars.usda.gov/mwa/madison/dfrc
Email contact: Lori.Bocher@ars.usda.gov
This material may be downloaded, printed and/or copied for educational purposes.
It is based on a series of articles, written by James B. Russell, that originally ran in Farm Progress publications.
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 Table of Contents
What is a ruminant? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Are rumens really full of bacteria and other bugs? . . . . . . . . . . . . . . . . . . . . . 2
Did dinosaurs have rumens? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Do belching cows contribute to global warming? . . . . . . . . . . . . . . . . . . . . . . 6
If a cow can digest cellulose, why cant I feed her sawdust? . . . . . . . . . . . . . 7
Why do cattle waste so much pasture protein?. . . . . . . . . . . . . . . . . . . . . . . . 8
Can too much starch in the diet kill my cow? . . . . . . . . . . . . . . . . . . . . . . . . . 9
What role do rumen bacteria play in grass tetany?. . . . . . . . . . . . . . . . . . . . 10
What do cattle and cigarette smokers have in common?. . . . . . . . . . . . . . . 11
How much urea can cattle safely use?. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
If cows utilize non-protein nitrogen, why do we feed true protein? . . . . . . . 13
Are rumen protozoa bad for my cow? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
What do rumens have in common with athletes foot? . . . . . . . . . . . . . . . . . 15
Do rumen bacteria cause polio in cattle? . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
What do rumen bacteria do with nitrate? . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Can ruminants be inoculated with better bacteria? . . . . . . . . . . . . . . . . . . . 18
Can bacteria prevent kidney stones and oxalate poisoning?. . . . . . . . . . . . 19
Do rumen bacteria cause liver abscesses? . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Q: What is a ruminant?
Man has long depended on rumi- until 1884 when H. van Tappeiner added antiseptics

A: nant animals for food. Prehistoric

hunters often followed herds of
grazing ruminants, and domestica-
tion of these herbivores allowed
early man to more fully exploit the photosynthetic
potential of the temperate grass-
to rumen fluid and demonstrated an inhibition of
fiber digestion.

Websters dictionary describes ruminants as any of

a group of four footed, hoofed, even toed, and cud
chewing mammals as the cattle,
lands. Domesticated ruminants were buffalo, bison, goat, deer, antelope,
a symbol of economic status and . . . four-footed, hoofed, camel, giraffe, llama, etc., which
well being, and ancient Jews were even-toed, and cud chewing have a stomach consisting of four
quick to recognize the comparative divisions or chambers, the rumen,
mammals . . . which have a
safety of ruminant meat. Whatever reticulum, omasum, and abomasum;
divides a hoof, thus making split stomach consisting of four the grass etc. that they eat is swal-
hoofs, and chews the cud, among the divisions or chambers. lowed unchewed and passes into the
animals, that may you eat (Le- rumen or reticulum from which it is
viticus 11:3). Aristotle, a scholar of regurgitated, chewed and mixed with
many disciplines, described the four compartments of saliva, again swallowed, and then passed through the
the ruminant stomach, but the role of microorganisms reticulum and omasum into the abomasum where
in the ruminant style of digestion was not recognized it is acted on by gastric juice. This definition is an
anatomically detailed description, but it
lacks any mention of microorganisms or
fermentation.

Modern agricultural practices have placed

new constraints on domestic ruminants
and rumen fermentation. Genetic improve-
ments have produced animals that have
much greater nutritional needs, and the
increased size of farms has intensified the
problem of manure disposal. Following is
a series of articles on rumen fermentation.
Each of these articles will focus on some
aspect of rumen fermentation or a problem
unique to ruminant animals.

Ruminations 1
 Are rumens really full of
Q: bacteria and other bugs?
Yes, and were glad they are. Ru- directly utilize most

A: men microbes could be considered a

cows best friend. Without microbes,
a cows digestive system would shut
down and she would starve to death.

Cows and microbes actually have

feed components, even
simple sugars. She
relies on rumen

microbes to convert
feeds to volatile fatty
Bossy,
lets live
a mutually beneficial relationship. together! acids (VFAs) that the
Microbes give the cow: cow absorbs and uses
z labor to digest feed; to make energy and
z a source of protein; milk. About two-thirds
z a source of volatile fatty acids; of feed digestion takes
z the ability to digest forage. place in the rumen,
On the other hand, cows provide and 90 percent of fiber
microbes with: digestion all with the aid of microbes.
z water;
z warmth; While rumen microbes help cows digest feed,
Come in from
z grinding (cud the cold!
theyre also an important source of feed for cows.
chewing or rumina- Ill grind your Theyre typically about 55 percent protein; on some
tion) of feed; food for you. rations the microbes provide 90 percent of the total
z anaerobic (no dietary protein requirement of the cow. Even more
oxygen) conditions. amazing is the fact that rumen microbial protein has
an ideal mixture of amino acids one that is not
There are three main groups of rumen microbes:
z Bacteria carry out most of the digestion of sugars,
starch, fiber, and protein for the cow.
z Protozoa swallow and digest bacteria, starch gran- Get back in
ules, and some fiber. the rumen
z Fungi make up only a small fraction of the rumen where you
microbial population, but they appear to be important belong!
particularly if the forage quality is poor. They are
though to open plant fibers to make them more easily
digested by the bacteria.

Almost all rumen microbes are anaerobic; they will

only grow in an oxygen-free environment. New
microbes are constantly being produced in the rumen
(under adequate conditions) while old ones are passed
on down through the cows digestive tract. One type of
rumen bacteria can double its population in 24 minutes!

In many ways, when you feed your cow youre really

feeding the microbes in her rumen. The cow cannot We get the food first!

Ruminations 2
 Did you know
easily duplicated in any ration. And no 2 cows have the
rumen microbes are a particularly rich same population of
source of lysine and methionine, two rumen bacteria?
amino acids that are difficult to sup-
plemnt in dairy cattle rations.

Rumen microbes, cows and other ruminants can

make use of feed, like alfalfa and grasses, that
people cant eat. Many of these plants are grown
on land that isnt suitable for other crops. Thus,
cattle produce food from land that might otherwise
be under utilized. Cows are also useful consumers
of by-products from human food production (like
citrus or beet pulp) and from grain ethanol produc- I suppose
tion (distillers grains). you think
yours are
better?

Photographs of rumen microbes

Photo by Lydia Joubert. 1995, Mel Yokoyama & Mario A. Cobos.

Bacteria attacking a strand of fiber that was taken from a This is one of the biggest rumen protozoa. Protozoal
cows rumen. cells are much larger than bacterial cells; some can
almost be seen with the naked eye.

Ruminations 3
 1995, Mel Yokoyama & Mario A. Cobos.

The large microbe is a type of protozoan. The creature that

looks like a tadpole attached to the side of the protozoan is 1995, Mel Yokoyama & Mario A. Cobos.

a fungal spore. The smaller, rod-shaped beasts lining the Most rumen microbes, like this protozoan, grow by increasing
underside of the protozoan are bacteria. their cell size, then splitting in half to make two cells.
Under ideal conditions, most species of rumen bacteria
can double their populations in a few hours.

1995, Mel Yokoyama & Mario A. Cobos.

The hairy stuff on the outside of this protozoal cell is cilia

Latin for eyelash. These cilia wave back and forth and
either propel the microbe through the water or propel food
This rumen protozoan is covered with chains of bacteria.
toward it. This protozoan has several chains of bacteria
Some protozoal and bacterial cells actually have a symbiotic
that are stuck to it.
relationship, each giving the other something that it needs
to survive.

Ruminations 4
Q: Did dinosaurs have rumens?
With films like Jurassic Park and has another constraint. Acetic acid (the acid found in

A: Lost World, there has been an

increased interest in dinosaurs.
Some of the dinosaurs were meat-
eating carnivores, but the largest
ones were plant-eating herbivores. Because ruminant
animals (cattle, sheep, goats, deer, etc.) utilize ru-
vinegar) is the primary end-product of rumen fermen-
tation, and the ruminants use this acid for energy. If
the rumen retention time gets too long, a bacterium
called Methanosarcina is able to convert acetic acid
to methane and
carbon dioxide.
men microorganisms to digest plant material, it is Since these gas- The problem with dinosaurs
reasonable to ask the question Did dinosaurs have a es are belched having a traditional rumen
rumen? away, the animal is their very large body size.
would loose
The script to Jurassic Park indicated that dinosaurs most of its
were giant cows, but there are scientific reasons for energy. Nature has prevented the problem of acetate
disputing this idea. conversion to gas by making sure that retention time
is never too long and is short
All animals have microorgan- enough to wash Methanosar-
isms in their gastrointestinal cina out of the rumen.
tracts, but the rumen is a
highly specialized organ that The problem with dinosaurs
facilitates the digestion of having a traditional rumen is
fibrous plant materials. Adult their very large body size. The
ruminants have a very large largest plant-eating dinosaurs
rumen compartment, and the weighed more than 25 tons.
rumen usually accounts for Based on the observation that
1/10 to 1/7 of the animals feed retention time in the ru-
weight. By having a large men and body size appears to
rumen, ruminants are able be a linear function, the reten-
to retain feed material for a tion time of a dinosaur rumen
long time, and this gives the could be greater than 1250
microorganisms a greater chance to digest food. hours, a time clearly not compatible with efficient
acetate utilization.
Rumen retention time increases with the size of the
ruminant. The duiker is a very small ruminant that Some large herbivores have a different strategy for
lives in Africa, and mature adults have a weight that digesting plants. The cecum is an enlarged pouch that
is less than 10 pounds. Because duikers have very extends from the intestines, and it too can serve as
small rumens, they are not able to eat as much fiber a habitat for microorganisms. Humans have a very
as large ruminants like cattle and buffalo. Duikers small cecum (the appendix), but horses and elephants
are browsers, and they consume tender leaves that have enlarged ceca. Cecal fermenters do not retain
are rich in sugars and other readily digested carbohy- feed as long as or digest plant materials as com-
drates. pletely as ruminants, but this may not have been a
problem for dinosaurs. Large plant-eating mammals
The strategy of having an ever larger rumen so the had not yet evolved, and plants may have been more
microorganisms will have more time to digest fiber digestible.

Ruminations 5
 Do belching cows contribute
Q: to global warming?
The rapid conversion of fossil fuels tive to chlorinated hydrocarbons like chloroform, and

A: to carbon dioxide has lead many

scientists to believe that the earth
is becoming a global greenhouse.
Because even small changes in
average temperature would have catastrophic effects
on climate, ocean depth and other aspects of our
there is enough chloroform in three medicated cough
drops to completely inhibit the methane production
of a sheep. The ability of chloroform to inhibit rumen
methane production was, however, confounded by
the ability of rumen bacteria to adapt. Ruminal bacte-
ria eventually reduced and inactivated the chloroform.
environment, the U.S. government has undertaken a
detailed examination of all facets of global warming. It the mid-1970s a veterinary pharmaceutical com-
Carbon dioxide is clearly the most abundant green- pany developed a product called monensin to treat an
house gas, but it is not the most potent one. Methane intestinal disease, coccidiosis, in pigs and chickens.
is 15 times more However, researchers noted that this antibiotic could
potent than carbon also improve feed efficiency of sheep. Monensin and
dioxide in trapping other ionophroes dont completely inhibit methane
the suns radiation. production, but reductions as great as 33% have been
reported. Monensin has a different mode of action
Methane gas is from chloroform. Methane-producing bacteria are not
derived from a sensitive to monensin, but it prevents carbohydrate-
variety of sources fermenting bacteria from producing hydrogen, a pre-
that include natural cursor of rumen
gas leaks from the methane.
petroleum industry The impact of American
as well as biological The rumen is nor-
sources. Biological
cattle on global biogas mally well buff-
methane, or bio- production and global ered by saliva, but
gas, is emitted from warming is still being rumen pH can de-
wetlands, rice pad- analyzed, but the effect is cline (acidosis) if
dies, termite guts, the rate of starch
and ruminants. Es- probably insignificant. fermentation is
timates of ruminant rapid. Low rumen
methane produc- pH causes a vari-
tion have changed ety of problems (discussed in forthcoming segments),
considerably. Early but it has at least one positive effect. Low rumen pH
estimates indicated inhibits methane-producing bacteria. Animals with
that ruminants might be producing as much as a third low rumen pH produce less methane and retain more
of the total, but more recent estimates are 15 to 20%. of their feed energy.

Methane is a natural end-product of rumen fermenta- The impact of American cattle on global biogas pro-
tion and feed digestion. Cattle produce as much as 50 duction and global warming is still being analyzed, but
liters of gas per hour and as much as 1/3 of this gas the effect is probably insignificant because American
can be methane. Because methane represents a sig- cattle consume very little feed per unit of production
nificant loss of feed energy, nutritionists sought ways (pounds of milk or meat), often have a rumen pH that
of reducing methane production in the early 1970s. inhibits methane production, and are sometimes fed
Rumen bacteria that produce methane are very sensi- additives that inhibit methane production.
Ruminations 6
 If a cow can digest cellulose,
Q: why cant I feed her sawdust?
Cellulose is the most abundant inside out. Woody plant cells are nearly completely

A: polymer in nature, but mammals

do not produce enzymes that can
digest this material. Ruminant
animals, by developing a symbiotic
relationship with cellulose-digesting bacteria, have
developed considerable capacity to digest cellulose,
filled with cellulose, and have much less surface area
for microbial attack.

Another feature that limits the digestion of cellulose

and other cellulosic materials is lignin. Lignin is a
highly branched compound that binds to hemicellu-
and ruminant animals can be fed diets that are rich in lose. Lignin cannot be degraded by ruminal bacteria
cellulose and other fibrous materials. and it acts to block the enzymes of rumen microor-
ganisms from ever using some of
Dr. Paul Weimer (U.S. Dairy the carbohydrate. Woody plants
Forage Research Center) esti- Rumen bacteria have the capacity (sawdust) typically have a much
mated that the global rumen vol- to digest virtually any type of higher lignin content than for-
ume of domesticated ruminants age crops and are inherently less
is approximately 100 billion cellulose, but the cellulose digestible.
liters (2 billion animals with 50 digestion rate of mature and highly
liters per animal). Based on this lignified plant cells can be Rumen bacteria have the capac-
estimate, the rumen is the largest ity to digest virtually any type
very slow.
commercial fermentation vat in of cellulose, but the cellulose
the world. digestion rate of mature and
highly lignified plant cells can be very slow. Sawdust
Cellulose and starch are both repeating glucose would provide some energy to a cow, but not enough
structures, but the two molecules have different to maintain her. Corn stalks and poor quality hay pro-
bonds and physical structures. Starch chains can vide enough energy for maintenance but not enough
bind to other starch chains, but the hydrogen bond- for rapid weight gain or lactation. Rapidly growing
ing is relatively weak and can be easily disrupted and heavily lactating animals can utilize high qual-
with mild treatments (e.g. heat). Any person who has ity forages, but they usually need at least some very
made gravy has watched the starch dissolve when the rapidly fermented carbohydrate (e.g. starch).
water was heated. By contrast, cellulose chains are
so strongly bonded to other cellulose chains that only
strong acid can cause solubilization.

The propensity of cellulose molecules to form

insoluble bundles reduces the surface area
exposed to enzymes, and this feature is the
rate limiting step in rumen cellulose digestion.
The animal rechews the cud (ruminates) on an
almost continual basis and increases the sur-
face area of the cellulose, but this mechanical
process of particle size reduction has clear cut
and obvious limitations. The plant cell walls of
forage crops can be easily disrupted and rumen
bacteria typically digest plant cells from the

Ruminations 7
 Why do cattle waste so
Q: much pasture protein?
Domestic ruminants have been digestion provides rumen bacteria with twice as

A: allowed to graze pasture since an-

cient times, but pasture utilization
declined when farmers had mecha-
nized methods of harvesting, stor-
ing, and feeding forages during the summer months.
In the 1980s, rotational grazing gained popularity,
much protein as they can efficiently use. The excess
protein is wastefully converted to rumen ammonia.

Ammonia is a highly volatile substance that is read-

ily absorbed from the rumen into blood. The animal
transports the ammonia to the liver and kidney and
and pasture management was purported to decrease converts it into urea, a less toxic substance. The
the costs of production. animal then excretes the urea as urine. Because graz-
ing ruminants stand
Grazing ruminants can still when they urinate,
do well on pasture, but the amount of nitrogen
it should be recognized applied to the soil can
that forages are inher- be extremely high on a
ently mismatched with g N/square foot basis. If
respect to carbohydrate the nitrogen percolates
and protein degrada- through the soil before
tion. Rumen bacteria the plant captures it, the
grow optimally when excess rumen protein
approximately 14% of ends up in the ground
the potentially digestible water as urea, nitrate or
organic matter is protein other types of nitrogen.
and 84% is carbohy- If the grazing density is
drate. Since lush grass high, ruminants can be
and other forage crops a very significant source
have a composition of environmental
similar to the amounts pollution.
preferred by rumen bacteria, This imbalance of carbohydrate
pasture would, in theory, be an and protein digestion provides Animals that are not allowed to
ideal substrate for rumen fer- graze pasture are typically fed
mentation. These simple com- rumen bacteria with twice as much total-mixed rations, and under
positions, however, ignore the protein as they can efficiently use. these conditions it is fairly easy
dynamics of rumen fermentation. to match the rates of rumen
protein and carbohydrate deg-
Lush forage contains some sugar and other readily radation. If the ration has an abundance of forage
fermented carbohydrate, but the bulk of the forage protein, one can simply increase the rate of carbohy-
carbohydrate is fiber (cellulose and hemicellulose). drate fermentation by adding high moisture corn or
The rumen degradation rate of fiber is so slow that some other rapidly fermented carbohydrate. Animals
only half of it is digested in the rumen. Forage pro- grazing on pasture could be supplemented in a simi-
tein is, by contrast, almost completely digested in the lar fashion, but this practice has not yet been widely
rumen. This imbalance of carbohydrate and protein adopted.

Ruminations 8
 Can too much starch in the
Q: diet kill my cow?
Wild ruminants do not consume tively, S. bovis numbers do not increase so dramati-

A: large amounts of starch, but cereal

grains are often added to domestic
rations. Starch increases the over-
all rate of fermentation, provides
a mechanism of balancing the rates of carbohydrate
and protein degradation, and usually causes an
cally and lactate does not accumulate.

Ruminants can also develop a subclinical form of

acidosis that does not involve S. bovis or lactate ac-
cumulation. Subclinical acidosis is caused by a very
high volatile fatty acid concentration. Researchers
increase in animal performance. Starch also causes have debated the cause of subclinical acidosis, but a
dramatic decreases in rumen pH, acute indigestion, decline in rumen motility and absorption is a likely
depressions in food intake, founder, and even death cause. When starch increases, forage decreases, and
of the animal. forage provides tactile stimulation needed to trigger
rumen movements. If the rumen does not mix, acids
Forage-fed cattle have low numbers of Streptococcus are absorbed more slowly. The role of tactile stimula-
bovis, a bacterium that grows very rapidly on starch; tion in acid absorption is illustrated by the work of
but its numbers can increase as Dr. Loerch at Ohio State Univer-
much as 1,000-fold if a large sity. He put nylon pot scrubbers
The best prevention for acute
amount of starch is added to the in the rumen and demonstrated an
diet. Because S. bovis produces acidosis is gradual adaptation increase in pH.
large amounts of lactic acid, the to starch and the timely feeding
buffering capacity of the rumen of carefully formulated rations. Hay is the typical remedy for
is overpowered and pH declines. acidosis, but the syndrome can be
When the pH is 5.6 or lower, the very difficult to reverse. Because
normal microorganisms are inhibited, and S. bovis normal rumen microorganisms are eliminated by low
has an even greater advantage. rumen pH, the fermentation tends to produce lactate
even after the starch is removed. Once the animals
Acute acidosis is a calamity. When the rumen pH de- have sore feet, they tend to lie down most of the
clines, the blood pH also decreases, time. When the animals get up, they
and this effect causes organ damage. eat their meals very quickly and
Because tissues above the horny this feeding pattern tends to bring
hoof are particularly sensitive to pH down. Chemical methods for
changes in blood pH, sore feet are preventing acidosis but have been
also a symptom. If the rumen pH incomplete. Rumensin, an additive
is less than 5.0, water rushes from that inhibits S. bovis only moderates
the blood into the rumen. This flux rumen pH somewhat. Sodium bicar-
of water causes hemoconcentration bonate is often marketed as a rumen
and death. The best prevention for buffer, but it really stimulates water
acute acidosis is gradual adaptation intake. When water intake increases,
to starch and the timely feeding of acid and unfermented starch can be
carefully formulated rations. Under washed out of the rumen.
these conditions the normal rumen
flora utilizes the starch more effec-

Ruminations 9
 What role do rumen bacteria
Q: play in grass tetany?
Lactating cattle that graze lush

A: grass sometimes develop a severe

form of magnesium deficiency
commonly called grass tetany.
Grasses causing tetany are low
in magnesium, but this deficiency alone could not
explain the problem. USDA researchers Dave Grunes
and Floyd Horn correlated the incidence of tetany
with the presence of an unusual acid in plants, trans-
aconitic acid (TA). These grasses accumulated as
much as 6% TA, and this acid can bind magnesium.

In the 1980s, we
examined the Grasses causing tetany
ability of rumen are low in magnesium, but
bacteria to me- this deficiency alone could
tabolize TA and
noted that TA was not explain the problem.
rapidly converted
to another acid, tricarballylic acid (TC). Animals
absorbed TC, but they were unable to degrade it.
TC bound blood magnesium, calcium and zinc and
increased urinary excretion. Selenomonas ruminan-
tium, a bacterium common in grass-fed cattle, con-
verted TA to TC.

The reason for TA accumulation in plants is not

entirely clear, but it may be linked to overfertiliza-
tion and high soil potassium. When grasses and small
grain forages are cultivated with high soil potas-
sium, they take up more potassium and this posi-
tively charged ion must be balanced by a negatively
charged ion. Some plants of the grass family use
trans-aconitate as a negatively charged ion.

Some years ago we isolated a bacterium from the ru-

men (Acidaminococcus fermentans) that was able to
convert TA into acetic acid, a normal end-product of
rumen fermentation. A. fermentans is found at very
low numbers in the rumen; it is a rapidly growing
organism when TA is available. A. Fermentans has
potential as a rumen inoculant. If A. fermentans num-
bers in the rumen could be increased, cattle would
have a mechanism to protect them from grass tetany.
Ruminations 10
 What do cattle and cigarette
Q: smokers have in common?
In the western United States, A variety of antibiotics (chlortetracycline, penicillin,

A: cattle are often taken from the

range to irrigated pastures in the
fall, and this abrupt change in
diet can cause a condition known
as acute bovine emphysema. Researchers initially
suspected some type of mold or fungus, but none
neomycin) were able to inhibit skatole production in
the laboratory, but rumen bacteria quickly become
resistant to traditional antibiotics. Lactobacilli, how-
ever, can also be inhibited by the feed additives Ru-
mensin and Bovitec, and resistance does not seem to
be a problem. Rumensin and Bovitec have decreased
could be correlated with the incidence of this disease. the incidence of bovine emphysema, but these addi-
Subsequent work by J.R. Carlson and his colleagues tives can be difficult to administer to grazing cattle.
at Washington State University demonstrated that ru- Wise pasture management and gradual adaptation
men bacteria were converting the amino acid trypto- is still the most effective method of controlling this
phan into skatole, a highly selective lung toxin that is potentially fatal syndrome.
also found in cigarette smoke.

Skatole production in the rumen appears to be a . . . rumen bacteria were

two step process. When normal rumen bacteria are converting the amino acid
abruptly given large amounts of soluble protein tryptophan into skatole, a
(lush irrigated pasture), they ferment tryptophan and
convert it to indoleacetic acid. A highly specialized highly selective lung toxin
lactobacillus then takes up the indoleacetic acid and that is also found in
converts it to skatole. cigarette smoke.

ska
tole

Ruminations 11
 How much urea can cattle
Q: safely use?
Mammals never developed the Rumen bacteria are never able to utilize more than

A: capacity to synthesize all 20 of the

amino acids commonly found in
protein and require at least 8 amino
acids. Because most rumen bacteria
are able to synthesize all 20 amino acids, ruminant
animals can be maintained on diets that are protein-
7 g of urea per 100 g of rumen degraded carbohy-
drate. If less carbohydrate is being degraded in the
rumen or if the rumen bacteria have a source of ru-
minally degraded true protein, the animal will need
less urea. Dr. Larry Satter of the U.S. Dairy Forage
Research Center indicated that cattle receiving
free. If the rumen bacteria have sufficient non-protein 14% true protein receive no benefit from dietary
nitrogen (ammonia, urea, etc.), microbial protein urea supplements.
flow from the rumen provides a continuous supply of
all 20 amino acids. Adapted animals can protect themselves from
excess dietary urea, but excess urea may decrease
Virtually all of the rumen bacteria utilize carbohy- the energetic efficiency. The animal must expend
drates as an energy source to synthesize amino acids, energy to synthesize and excrete urea as urine. The
and urea utilization is directly proportional to the Cornell Net Carbohydrate and Protein System and
supply of ruminally degraded carbohydrate. Poor the 1996 Beef NRC have a rumen sub-model that
quality diets that are rich in fiber supply approxi- allows users to monitor ammonia accumulation in
mately half as much rumen available carbohydrate as the rumen as well as the cost of urea synthesis by
diets based primarily on cereal grains. the animal.

When cattle receive more urea than

the rumen bacteria use, ammonia When cattle receive more
accumulates in the rumen and urea than the rumen bacteria
eventually enters the blood. Am-
monia is a highly toxic compound,
use, ammonia accumulates
and the animal detoxifies the am- in the rumen and eventually
monia by converting it to urea in enters the blood.
the liver and kidney. The enzymes
needed to synthesize urea are
inducible. If the flux of ammonia into blood has been
low, the animal will have a low activity of urea cycle
enzymes. The animal can synthesize additional urea
cycle enzymes, but a period of adaptation is required.

If animals consuming poor quality diets are given a

large dose of urea, the rumen bacteria will not be able
to utilize all of the rumen ammonia. If the animal is
unadapted, ammonia flux into blood can be so great
that it kills the animal. The cause of death is still be-
ing debated, but the depletion of -ketoglutarate (a
metabolite in brain tissue) appears to be involved.

Ruminations 12
 If cows utilize non-protein nitrogen,
Q: why do we feed true protein?
Rumen bacteria can utilize non-

A:
Rumen microorganisms can utilize non-protein nitro-
protein nitrogen sources like urea gen, but they prefer to utilize true protein. Because
and supply the animal with amino rumen microorganisms grow faster and more effi-
acids, but animals fed only non- ciently when they have true protein, the net return of
protein nitrogen never attain high amino acids to the animal can be greater if ruminally
levels of production. Highly productive animals have degraded protein is added to the diet. The optimal
very high amino acid requirements, and the flow ratio of non-protein nitrogen, ruminally degraded
of microbial protein from the rumen is inadequate. true protein, and rumen undegraded true protein is
By supplying protein that is protected from rumen not easy to calculate. Cornell Net Carbohydrate and
degradation, producers have in many cases increased Protein System and the 1996 Beef NRC have calcu-
amino acid supply and im- lations for determining poten-
proved animal productivity. Rumen microorganisms can utilize tial benefit of rumen degraded
protein to the microorganisms
When animals are fed rations non-protein nitrogen, but they and this relationship is based
containing high quality forages prefer to utilize true protein. on efficiency of rumen degrad-
or soybean meal, rumen micro- able carbohydrate as well as pH.
organisms often have more true protein than they can Users are able to monitor the protein or energy allow-
utilize, and the remainder is converted to excess ru- able production, the overall and rumen amino acid
men ammonia and eventually urinary urea. Because balance, as well as total nitrogen excretion.
rumen bacteria can utilize non-protein nitrogen,
many ruminant nutritionists thought that virtually all Silages often contain large amounts of non-protein
of the true protein should be protected from rumen nitrogen, but this fraction can have a large amount
degradation. of peptides and amino acids. Dr. Rich Muck of the
U.S. Dairy Forage Research Center indicated that
the non-protein nitrogen of high
quality silages can be as much as
2/3 amino nitrogen. Because rumen
microorganisms use amino acids as
efficiently as other forms of rumen
degraded protein, the potential ben-
efit of silage crude protein can be
underestimated.

We prefer the
real stuff,
true protein.

Ruminations 13
 Are rumen protozoa
Q: bad for my cow?
The rumen is one of the most

A: constant and carefully controlled

habitats in all of nature. The cow
consumes food almost continuous-
ly, the temperature is maintained
at 39 C, the pH is controlled with the bicarbonate of
saliva, acids are absorbed directly across the rumen
wall, gases escape by eructation, and mixing motions
mix the feed with microorganisms. This constancy
has selected for a highly diverse microbial popula-
tion in the same way that the tropical rain forest has
selected for a broad range of plant life.

The rumen has at least 50 species of bacteria, 11 gen- The rumen protozoa are predators
era of protozoa and at least 5 types of fungi. Bacteria
are clearly the most important microorganisms in that graze on bacteria as well as
the rumen, and they are responsible for most aspects plant materials. When protozoa
of the fermentation. Protozoa are much larger than ingest bacteria, they digest the
bacteria, and they grow much more slowly. Nonethe-
less, the protozoa can be a very significant part of the
microbial protein.
population. In many cases, protozoa can account for
as much as half of the microbial protein in the rumen.

The rumen protozoa are predators that graze on

bacteria as well as plant materials. When protozoa
ingest bacteria, they digest the microbial protein.
This turnover of bacteria increases the ammonia con-
centration and decreases the overall flow of microbial
protein from the rumen. The protozoa are a relatively
unstable population, and the spontaneous breakdown
of protozoa also causes an increase in ammonia.

Protozoa can be removed from the rumen in at least

two ways. If calves are taken from their mothers
immediately after birth or by Caesarean section and
raised in isolation, the rumen is not inoculated. The
protozoa can also be removed by chemicals (dime-
tridazole), but these treatments can kill the animal.

Ruminations 14
 What do rumens have in
Q: common with athletes foot?
The mental association of fungi possessed by bacte-

A: and athletes foot is a strong one,

but it should be realized that fungi
are capable of occupying many
habitats. Anyone who has walked
along the floor of an old growth forest has observed
the activity of cellulose-digesting fungi. White and
ria or protozoa, and
they observed that
fungi could burrow
directly into feed
particles. On the
other hand, fungi
brown rot fungi attack dead and fallen trees and are seem to grow slow-
able to digest even the most crystalline and highly ly in the rumen and
lignified forms of cellulose. are usually present
at low numbers.
Rumen microorganisms also digest cellulose, but the
role of fungi in rumen fermentation was only recently Animals consum-
recognized. Cellulose digestion could be explained ing cool season
by the activity of cellulose-digesting bacteria and grasses (e.g. North-
protozoa, and the involvement of fungi seemed to be ern United States)
precluded by the observation that most fungi need generally have very low numbers of fungi, but fungi
oxygen, a nutrient that is not present in the rumen. numbers increase as the climate becomes more tropi-
cal. Warm season grasses have thick, highly lignified
Scientists had observed large swimming microorgan- cell walls and fungi are better suited to digest this
isms with tails (flagella) in rumen fluid , but these material.
cells were classified as protozoa rather than fungal
zoospores. In the 1970s, Dr. Colin Orpin became
convinced that fungi were living and growing in Rumen microorganisms also digest
the rumen. Feed particles were sometimes covered
with a dense mat of fungal rhizoids, the flagellated
cellulose, but the role of fungi in
cells arose from fungal sacks called a sporangia; and rumen fermentation was only
chitin, a carbohydrate unique to fungi, was detected. recently recognized.
Since Dr. Orpins pioneering work, many species of
anaerobic fungi have been isolated from the rumen
and gastrointestinal tract.

The importance of fungi in rumen fermentation has

been at times hotly debated. Proponents cited the fact
that fungal enzymes had more activity than the ones

Ruminations 15
 Do rumen bacteria cause
Q: polio in cattle?
Rumen fermentation produces

A: gases (chiefly carbon dioxide and

methane) that are normally belched
away, but some gas always passes
into the lungs. If the rumen gas
contains hydrogen sulfide, there can be an immediate
and acute effect on the animal that is very similar to
polio in humans. In ruminants, excess rumen hydro-
gen sulfide causes a type of polioencephalomalacia
(PEM) and brain lesions virtually identical to the
viral PEM that was common in humans before a vac-
cine was developed.

Hydrogen
If the rumen gas contains sulfide is the
hydrogen sulfide, there can be gas respon-
an immediate and acute effect sible for the
stench of
on the animal. rotten eggs,
and in the
rumen this gas is produced by a specialized group of
bacteria that use dietary sulfate as an energy source.
How does this gas affect the animal? Hydrogen
sulfide prevents oxygen from binding to mitochon-
drial cytochromes. When these structures cannot bind
oxygen, the animal dies of metabolic asphyxiation.
At lower levels of exposure, chronic hydrogen sulfate
toxicity causes nerve damage and a gradual loss of
motor activity.

Because rumen bacteria and ruminant animals need

dietary sulfate to grow, PEM is not always easy to
prevent. Nutritionists usually recommend sulfur to
nitrogen ratios less than 1 to 10, but this arbitrary
guideline does not differentiate whether the sulfur is
coming from sulfate or other sources. If the nitrogen
is derived from urea, higher ratios can be required.

Bacteria-producing hydrogen sulfide can be inhib-

ited by a class of chemicals called anthraquinones,
but these compounds have not yet been adequately
screened for use in cattle diets. Because anthraqui-
nones are fat soluble compounds, they could very
well accumulate in meat or milk.
Ruminations 16
 What do rumen bacteria
Q: do with nitrate?
The overuse of chemical fertilizers

A: or manure has in many cases in-

creased the nitrate concentration of
soil and ground water, and nitrates
can be a problem for cattle. Forage
crops tend to accumulate nitrate, and it has long been
recognized that oat hay can be poisonous. Green oat
hay is a particular problem, but virtually any forage
crop can accumulate nitrate if the soil concentration
is high and the rate of plant growth is rapid.

Nitrate itself is not a toxic substance, but it can be re- When cattle consume nitrates rapidly (lush forage
duced to nitrite by rumen bacteria. Nitrite is a potent or hay), step 1 is much faster than step 2, and nitrite
poison. Nitrite combines with blood hemoglobin and accumulates in the rumen. Step 2 is somewhat adapt-
prevents it from transporting oxygen to the tissues. able. If animals are gradually adapted to increasing
Symptoms of nitrate poisoning include trembling, amounts of nitrate, step 2 increases, and the animal
staggering, rapid breathing and death. Nitrate con- can tolerate more nitrate in the diet. The potential
centrations in plants greater than 0.02% can be toxic, impact of nitrates on cattle can also be decreased by
but this value is highly dependent on other ingre- making sure that the diet has an abundance of rumi-
dients in the diet as well as adaptation and feeding nally degradable carbohydrates (starch and sugars).
behavior. When rumen bacteria have an ample amount of car-
bohydrates, they can increase their rate of growth and
Nitrate-reducing bacteria are always found at high amino acid synthesis.
numbers in the rumen, and many of these bacteria
can use nitrate as a electron acceptor to increase
Nitrate itself is not a toxic
energy production. Nitrate can be used as a nitrogen
source to synthesize amino acids, but this process is a substance, but it can be
three step conversion. Nitrate must first be converted reduced to nitrite by rumen
to nitrite, the nitrite is converted to ammonia, and the bacteria.
ammonia is then used to synthesize amino acids:

Ruminations 17
 Can ruminants be inoculated
Q: with better bacteria?
It is generally agreed that tents from a variety of wild ruminants. Dr. Hungate

A: bacteria were the first forms of

life to inhabit the earth, and this
highly adaptable group of organ-
isms can grow in even harsh
habitats (for example, hot springs with tempera-
tures near boiling and the gastric stomach of man).
found the same types of bacteria in wild ruminants
as had been obtained from domestic ruminants in the
United States, and no new species were obtained.
Later surveys of other geographic locations supported
the conclusion that wild and domestic ruminants had
the same types of bacteria.
Because bacteria are ubiquitous in nature (present
everywhere at all times), microbiologists had gener- Since the early 1980s, there has been considerable
ally assumed that inoculation of natural environ- interest in improving rumen bacteria via genetic en-
ments would be a futile exercise. If the gineering, but this has been a difficult
environment would permit growth, the task. Rumen bacteria are very good
bacteria would already be there. at degrading the various substrates in
feed, and in most cases, surface area
Leucaena is a leguminous plant that on the feed particles is the rate limit-
it commonly found in the tropics. ing step in the fermentation. There is,
Because the leaves are rich in protein however, at least one case when rumen
and highly digestible, Leucaena has bacteria are not up to the task.
potential as a forage crop for ruminant
animals. Leucaena, however, had one When cattle are fed large amounts of
serious drawback. Its leaves and seeds cereal grain, rumen pH can decrease to
had a toxic amino acid called mimo- the point where cellulose is no longer
sine. Mimosine caused low weight digested. The enzymes needed for
gain, hair loss, goiter and ulcers in the cellulose digestion are still active, but
esophagus. Dr. Raymond Jones from Australia noted the bacteria producing these enzymes (cellulases) are
that Hawaiian goats were able to consume Leucaena unable to maintain a favorable intracellular pH and
without the toxic side effects, and he hypothesized grow. Some years ago we identified a rumen bacte-
that these goats might have mimosine degrading rium that was able to grow at low pH, and this bacte-
bacteria in their rumens. In vitro studies indicated rium produces a very weak cellulase.
that this hypothesis was correct, and in vivo inocula-
tion of Australian cattle with rumen bacteria from We have been attempting to improve the acid-resis-
Hawaiian goats corrected the tant rumen bacterium to increase
toxic symptoms of Leucaena its rate of cellulose digestion.
consumption.
Since the early 1980s, there has We were able to genetically
been considerable interest in modify the cellulase so it would
The success of rumen inocula- improving rumen bacteria via digest cellulose at a faster rate,
tion as a means for correcting genetic engineering, but this has but transfer of the reconstructed
mimosine toxicity in Australia gene back into the rumen bacte-
indicated that bacteria were not been a difficult task. rium has been a tedious exercise.
always ubiquitous in nature, Rumen bacteria do not readily
but this example is probably rare. In the 1960s, Rob- take up genetic material, and many of them have dif-
ert Hungate, the father of rumen microbiology trav- ferent mechanisms for expressing genes than E. coli,
eled extensively in Africa and sampled rumen con- the work-horse of genetic engineering.

Ruminations 18
 Can bacteria prevent kidney
Q: stones and oxalate poisoning?
Some forages (halogeton, tur- Oxalobacter formigenes, and this bacterium grows

A: nip tops, mangolds) and human

foods (tea, spinach, rhubarb) have
relatively high concentrations of
oxalate, an organic acid that can
be toxic to animals and man. High oxalate intake
causes diarrhea, distress and even death. In humans,
slowly and cannot utilize other substrates. The role of
O. formigenes in oxalate degradation is supported by
the observation that inoculation increased the degra-
dation rate of oxalate in unadapted animals.

Enteric diseases such as Crohns disease or ileal

oxalate forms a insoluble complex with calcium, and bypass surgery increases a persons risk of kidney
this material can ac- stones and other
cumulate in the kidney aspects of oxalate
as stones. Kidney In humans, oxalate forms a insoluble complex with poisoning. Physicians
stones are more than calcium, and this material can accumulate in the explained the increased
70% calcium oxalate. uptake of oxalate by
kidney as stones. . . . The role of O. formigenes in
a depression of fat
In the early 1980s, oxalate degradation is supported by the observation absorption. Undigested
Dr. Milton Allison that inoculation increased the degradation rate of fats bind calcium and
(USDA, Ames, IA) oxalate in unadapted animals. increase the amount of
adapted cattle and free oxalate, but there
sheep gradually to may also be a complete
increasing amounts of oxalate and demonstrated an loss of oxalate-degrading bacteria. When the colon
enhanced degradation by rumen bacteria. Later work is removed, bile salts are not absorbed as well, and
with rabbits, guinea pigs, and horses indicated that these salts seem to kill O. formigenes. Recent work
oxalate-degrading bacteria were also found in the has also indicated that antibiotic therapy may inhibit
intestines, but oxalate-degrading bacteria are not al- O. formigenes and increase the risk of kidney stones.
ways present. Wild rats degrade oxalate, but at least
some strains of laboratory rats cannot.

Oxalate is a highly oxidized acid that cannot be

degraded by most rumen or intestinal bacteria. The
only gut bacterium that appears to degrade oxalate is

Kidney stones. Source: www.sdm.buffalo.edu/scic/gallery (South Campus Instruemntation Center, University at Buffalo, State Universtiy of New York.

Ruminations 19
 Do rumen bacteria cause
Q: liver abscesses?
Cattle that are fed large amounts of racycline, Tylosin, etc.). Good feeding management

A: cereal grain (starch) have a higher

incidence of liver abscesses than
cattle that are fed forages, and
in the United States over 3 mil-
lion livers are condemned each year. The economic
impact of the abscesses is much greater than the loss
can also be an effective preventative measure. Liver
abscesses are most common when cattle are abruptly
shifted to hot rations after a period of starvation
(e.g. trucking).

of the liver itself. Cattle with liver abscesses have

lower weight gain, feed intake, and feed efficiency
than normal animals; and the dressing pecentage and
carcass grade are often reduced.

Diets causing liver abscesses alter end-products of

rumen fermentation. Rumen fermentation usually
produces volatile fatty acids (acetic, propionic and
butyric), but starch-digesting ruminal bacteria can
also produce large amounts of lactic acid. Lactic acid
is a very strong acid, and it can cause a pronounced
decrease in rumen pH. When pH is very low, the ru-
men wall is irritated. In severe cases, the rumen wall
deteriorates to the point of chronic bleeding.

Lactate accumulation in the rumen also promotes

the growth of Fusobacterium necrophorum, a lactic
acid-utilizing species. F. necrophorum is able to at-
tack the rumen wall and pass into the blood. Blood
supply from the rumen carries F. necrophorum to
the liver where it produces abscesses. The genetic
technique, restriction fragment length polymor-
phism (RFLP), was a key part of the O. J. Simpson
trial, and rumen microbiologists have used RFLP to
identify bacteria. Dr. T. G. Nagaraja of Kansas State
University has shown that F. necrophorum isolates
from the rumen wall are similar to those from the
liver but unrelated to those in rumen fluid. These
findings indicate that only certain F. necrophorum
strains cause liver abscesses, but the exact nature of
this difference has yet to be defined.

The study of liver abscess disease is complicated by

the fact that other bacteria are often involved in the
infection, but the incidence can be substantially re-
duced by a variety of antibiotics (bacitracin, chlortet-

Ruminations 20