Understanding the Ruminant

Animal Digestive System

Ruminant livestock include cattle, sheep, slightly less than one-third of their time
and goats. Ruminants are hoofed mam- idling where they are, neither grazing nor
mals that have a unique digestive system ruminating.
that allows them to better use energy from The roof of the ruminant mouth is a
fibrous plant material than other herbi- hard/soft palate without incisors. The
vores. Unlike monogastrics such as swine lower jaw incisors work against this hard
and poultry, ruminants have a digestive dental pad. The incisors of
system designed to ferment feedstuffs and grass/roughage selectors are wide with a
provide precursors for energy for the ani- shovel-shaped crown, while those of con-
mal to use. By better understanding how centrate selectors are narrower and chisel-
the digestive system of the ruminant shaped. Premolars and molars match
works, livestock producers can better between upper and lower jaws. These
understand how to care for and feed rumi- teeth crush and grind plant material dur-
nant animals. ing initial chewing and rumination.
Saliva aids in chewing and swallow-
Ruminant Digestive Anatomy ing, contains enzymes for breakdown of
fat (salivary lipase) and starch (salivary
and Function amylase), and is involved in nitrogen recy-
The ruminant digestive system uniquely cling to the rumen. Saliva’s most impor-
qualifies ruminant animals such as cattle tant function is to buffer pH levels in the
to efficiently use high roughage feedstuffs, reticulum and rumen. A mature cow pro-
including forages. Anatomy of the rumi- duces up to 50 quarts of saliva per day,
nant digestive system includes the mouth, but this varies, depending on the amount
tongue, salivary glands (producing saliva of time spent chewing feed, because that
for buffering rumen pH), esophagus, four- stimulates saliva production.
compartment stomach (rumen, reticulum, Forage and feed mixes with saliva
omasum, and abomasum), pancreas, gall containing sodium, potassium, phosphate,
bladder, small intestine (duodenum, bicarbonate, and urea when consumed, to
jejunum, and ileum), and large intestine form a bolus. That bolus then moves from
(cecum, colon, and rectum). the mouth to the reticulum through a
A ruminant uses its mouth (oral cavi- tube-like passage called the esophagus.
ty) and tongue to harvest forages during Muscle contractions and pressure differ-
grazing or to consume harvested feed- ences carry these substances down the
stuffs. Cattle harvest forages during graz- esophagus to the reticulum.
ing by wrapping their tongues around the
Ruminants eat rapidly, swallowing
plants and then pulling to tear the forage
much of their feedstuffs without chewing
for consumption. On average, cattle take
it sufficiently (< 1.5 inches). The esopha-
from 25,000 to more than 40,000 prehensile
gus functions bidirectionally in ruminants,
bites to harvest forage while grazing each
allowing them to regurgitate their cud for
day. They typically spend more than one-
further chewing, if necessary. The process
third of their time grazing, one-third of
of rumination or “chewing the cud” is
their time ruminating (cud chewing), and
where forage and other feedstuffs are
forced back to the mouth for further chewing and mix- The reticulorumen is home to a population of
ing with saliva. This cud is then swallowed again and microorganisms (microbes or “rumen bugs”) that
passed into the reticulum. Then the solid portion slow- include bacteria, protozoa, and fungi. These microbes
ly moves into the rumen for fermentation, while most ferment and break down plant cell walls into their car-
of the liquid portion rapidly moves from the reticulo- bohydrate fractions and produce volatile fatty acids
rumen into the omasum and then abomasum. The (VFAs), such as acetate (used for fat synthesis), priopi-
solid portion left behind in the rumen typically onate (used for glucose synthesis), and butyrate from
remains for up to 48 hours and forms a dense mat in these carbohydrates. The animal later uses these VFAs
the rumen, where microbes can use the fibrous feed- for energy.
stuffs to make precursors for energy. The reticulum is called the “honeycomb” because
True ruminants, such as cattle, sheep, goats, deer, of the honeycomb appearance of its lining. It sits
and antelope, have one stomach with four compart- underneath and toward the front of the rumen, lying
ments: the rumen, reticulum, omasum, and aboma- against the diaphragm. Ingesta flow freely between
sums. The ruminant stomach occupies almost 75 per- the reticulum and rumen. The main function of the
cent of the abdominal cavity, filling nearly all of the reticulum is to collect smaller digesta particles and
left side and extending significantly into the right side. move them into the omasum, while the larger particles
The relative size of the four compartments is as fol- remain in the rumen for further digestion.
lows: the rumen and reticulum comprise 84 percent of
the volume of the total stomach, the omasum 12 per-
cent, and the abomasum 4 percent. The rumen is the
largest stomach compartment, holding up to 40 gal-
lons in a mature cow.
The reticulum holds approximately 5 gallons in
the mature cow. Typically, the rumen and reticulum
are considered one organ because they have similar
functions and are separated only by a small muscular
fold of tissue. They are collectively referred to as the
reticulorumen. The omasum and abomasum hold up
to 15 and 7 gallons, respectively, in the mature cow.

“Honeycomb” interior lining of the reticulum in an 8-week old calf

The reticulum also traps and collects heavy/dense

objects the animal consumes. When a ruminant con-
sumes a nail, wire, or other sharp heavy object, it is
very likely the object will be caught in the reticulum.
During normal digestive tract contractions, this object
can penetrate the reticulum wall and make its way to
the heart, where it can lead to hardware disease. The
reticulum is sometimes referred to as the “hardware
Right-sided view of ruminant digestive tract stomach.” Hardware disease is discussed in detail in
Mississippi State University Extension Service
Publication 2519, “Beef Cattle Nutritional Disorders.”
The rumen is sometimes called the “paunch.” It is
lined with papillae for nutrient absorption and divided
by muscular pillars into the dorsal, ventral, caudodor-
sal, and caudoventral sacs. The rumen acts as a fermen-
tation vat by hosting microbial fermentation. About 50
to 65 percent of starch and soluble sugar consumed is
digested in the rumen. Rumen microorganisms (pri-
marily bacteria) digest cellulose from plant cell walls,
digest complex starch, synthesize protein from nonpro-
Left-sided view of ruminant digestive tract tein nitrogen, and synthesize B vitamins and vitamin
K. Rumen pH typically ranges from 6.5 to 6.8. The enzymes secreted from the pancreas, such as pancreat-
rumen environment is anaerobic (without oxygen). ic lipase (breaks down fats). These secretions help pre-
Gases produced in the rumen include carbon dioxide, pare proteins for absorption in the intestines. The pH
methane, and hydrogen sulfide. The gas fraction rises in the abomasum generally ranges from 3.5 to 4.0. The
to the top of the rumen above the liquid fraction. chief cells in the abomasum secrete mucous to protect
the abomasal wall from acid damage.

Interior lining of the rumen, revealing papillae in an 8-week old calf Interior lining of the abomasum, the “true stomach,” in an 8-week
old calf
The omasum is spherical and connected to the
reticulum by a short tunnel. It is called the “many The small and large intestines follow the aboma-
piles” or the “butcher’s bible” in reference to the many sum as further sites of nutrient absorption. The small
folds or leaves that resemble pages of a book. These intestine is a tube up to 150 feet long with a 20-gallon
folds increase the surface area, which increases the capacity in a mature cow. Digesta entering the small
area that absorbs nutrients from feed and water. Water intestine mix with secretions from the pancreas and
absorption occurs in the omasum. Cattle have a highly liver, which elevate the pH from 2.5 to between 7 and
developed, large omasum. 8. This higher pH is needed for enzymes in the small
intestine to work properly. Bile from the gall bladder is
secreted into the first section of the small intestine, the
duodenum, to aid in digestion. Active nutrient absorp-
tion occurs throughout the small intestine, including
rumen bypass protein absorption. The intestinal wall
contains numerous “finger-like” projections called villi
that increase intestinal surface area to aid in nutrient
absorption. Muscular contractions aid in mixing diges-
ta and moving it to the next section.
The large intestine absorbs water from material
passing through it and then excretes the remaining
material as feces from the rectum. The cecum is a large
blind pouch at the beginning of the large intestine,
approximately 3 feet long with a 2-gallon capacity in
the mature cow. The cecum serves little function in a
ruminant, unlike its role in horses. The colon is the site
of most of the water absorption in the large intestine.
Interior lining of the omasum, revealing the “many piles” tissue
folds in an 8-week old calf Ruminant Digestive Development
Immature ruminants, such as young, growing calves
The abomasum is the “true stomach” of a rumi- from birth to about 2 to 3 months of age, are function-
nant. It is the compartment that is most similar to a ally nonruminants. The reticular groove (sometimes
stomach in a nonruminant. The abomasum produces referred to as esophageal groove) in these young ani-
hydrochloric acid and digestive enzymes, such as mals is formed by muscular folds of the reticulum. It
pepsin (breaks down proteins), and receives digestive shunts milk directly to the omasum and then aboma-
sum, bypassing the reticulorumen. The rumen in these and omasum account for 35 percent and 14 percent of
animals must be inoculated with rumen microorgan- the total stomach area in the newborn calf. As rumi-
isms, including bacteria, fungi, and protozoa. This is nants develop, the reticulorumen and omasum grow
thought to be accomplished through mature rumi- rapidly and account for increasing proportions of the
nants licking calves and environmental contact with total stomach area. In mature cattle, the abomasum
these microorganisms. encompasses only 21 percent of the total stomach
Immature ruminants must undergo reticulorumen- capacity, whereas the reticulorumen and omasum
omasal growth, including increases in volume and make up 62 and 24 percent, respectively, of the total
muscle. In a calf at birth, the abomasum is the largest stomach area. Rumen papillae (sites of nutrient
compartment of the stomach, making up more than 50 absorption) lengthen and decrease in numbers as part
percent of the total stomach area. The reticulorumen of rumen development.

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Relative proportions of stomach compartments in cattle and sheep at various ages

Because immature ruminants do not have a functional species and also for multiple species grazed together
rumen, feeding recommendations differ for developing or on the same acreage.
ruminants compared with adult ruminants. For Concentrate selectors have a small reticulorumen
instance, it is recommended immature ruminants are in relation to body size and selectively browse trees
not allowed access to feeds containing non-protein and shrubs. Deer and giraffes are examples of concen-
nitrogen such as urea. Developing ruminants are also trate selectors. Animals in this group of ruminants
more sensitive to gossypol and dietary fat levels than select plants and plant parts high in easily digestible,
mature ruminants. Design nutritional programs for nutrient dense substances such as plant starch, protein,
ruminants considering animal age. and fat. For example, deer prefer legumes over grasses.
Concentrate selectors are very limited in their ability to
Ruminant Feeding Types digest the fibers and cellulose in plant cell walls.
Based on the diets they prefer, ruminants can be classi- Grass/roughage eaters (bulk and roughage eaters)
fied into distinct feeding types: concentrate selectors, include cattle and sheep. These ruminants depend on
grass/roughage eaters, and intermediate types. The diets of grasses and other fibrous plant material. They
relative sizes of various digestive system organs differ prefer diets of fresh grasses over legumes but can ade-
by ruminant feeding type, creating differences in feed- quately manage rapidly fermenting feedstuffs.
ing adaptations. Knowledge of grazing preferences Grass/roughage eaters have much longer intestines
and adaptations amongst ruminant livestock species relative to body length and a shorter proportion of
helps in planning grazing systems for each individual large intestine to small intestine as compared with con-
centrate selectors.
Goats are classified as intermediate types and prefer tissue damage within the rumen and can lead to ulcer-
forbs and browse such as woody, shrubby type plants. ations of the rumen wall. Take care to provide ade-
This group of ruminants has adaptations of both con- quate forage and avoid situations that might lead to
centrate selectors and grass/roughage eaters. They acidosis when feeding ruminants high-concentrate
have a fair though limited capacity to digest cellulose diets. Acidosis is discussed in detail in Mississippi
in plant cell walls. State University Extension Service Publication 2519,
“Beef Cattle Nutritional Disorders.” In addition, ener-
Carbohydrate Digestion gy as a nutrient in ruminant diets is discussed in detail
in Mississippi State University Extension Service
Forages Publication 2504, “Energy in Beef Cattle Diets.”
On high-forage diets ruminants often ruminate or
regurgitate ingested forage. This allows them to “chew
their cud” to reduce particle size and improve
Protein Digestion
digestibility. As ruminants are transitioned to higher Two sources of protein are available for the ruminant
concentrate (grain-based) diets, they ruminate less. to use: protein from feed and microbial protein from
Once inside the reticulorumen, forage is exposed the microbes that inhabit its rumen. A ruminant is
to a unique population of microbes that begin to fer- unique in that it has a symbiotic relationship with
ment and digest the plant cell wall components and these microbes. Like other living creatures, these
break these components down into carbohydrates and microbes have requirements for protein and energy to
sugars. Rumen microbes use carbohydrates along with facilitate growth and reproduction. During digestive
ammonia and amino acids to grow. The microbes fer- contractions, some of these microorganisms are
ment sugars to produce VFAs (acetate, propionate, “washed” out of the rumen into the abomasum where
butyrate), methane, hydrogen sulfide, and carbon they are digested like other proteins, thereby creating a
dioxide. The VFAs are then absorbed across the rumen source of protein for the animal.
wall, where they go to the liver. All crude protein (CP) the animal ingests is divid-
Once at the liver, the VFAs are converted to glu- ed into two fractions, degradable intake protein (DIP)
cose via gluconeogenesis. Because plant cell walls are and undegradable intake protein (UIP, also called
slow to digest, this acid production is very slow. “rumen bypass protein”). Each feedstuff (such as cot-
Coupled with routine rumination (chewing and tonseed meal, soybean hulls, and annual ryegrass for-
rechewing of the cud) that increases salivary flow, this age) has different proportions of each protein type.
makes for a rather stable pH environment (around 6.0). Rumen microbes break down the DIP into ammonia
(NH3) amino acids, and peptides, which are used by
High-Concentrate Feedstuffs (Grains) the microbes along with energy from carbohydrate
When ruminants are fed high-grain or concentrate digestion for growth and reproduction.
rations, the digestion process is similar to forage diges-
tion, with a few exceptions. Typically, on a high-grain
diet, there is less chewing and ruminating, which leads
to less salivary production and buffering agents’ being
produced. Additionally, most grains have a high con-
centration of readily digestible carbohydrates, unlike
the more structural carbohydrates found in plant cell
walls. This readily digestible carbohydrate is rapidly
digested, resulting in an increase in VFA production.
The relative concentrations of the VFAs are also
changed, with propionate being produced in the great-
est quantity, followed by acetate and butyrate. Less
methane and heat are produced as well. The increase
in VFA production leads to a more acidic environment
(pH 5.5). It also causes a shift in the microbial popula-
tion by decreasing the forage using microbial popula-
tion and potentially leading to a decrease in digestibili-
ty of forages.
Lactic acid, a strong acid, is a byproduct of starch
fermentation. Lactic acid production, coupled with the
increased VFA production, can overwhelm the rumi-
Protein digestion in the ruminant
nant’s ability to buffer and absorb these acids and lead
to metabolic acidosis. The acidic environment leads to
Excess ammonia is absorbed via the rumen wall and
converted into urea in the liver, where it returns in the that cannot be used by or are not available to other
blood to the saliva or is excreted by the body. Urea tox- animals. Ruminants are in a unique position of being
icity comes from overfeeding urea to ruminants. able to use such resources that are not in demand by
Ingested urea is immediately degraded to ammonia in humans but in turn provide man with a vital food
the rumen. source. Ruminants are also useful in converting vast
When more ammonia than energy is available for renewable resources from pasture into other products
building protein from the nitrogen supplied by urea, for human use such as hides, fertilizer, and other inedi-
the excess ammonia is absorbed through the rumen ble products (such as horns and bone).
wall. Toxicity occurs when the excess ammonia over- One of the best ways to improve agricultural sus-
whelms the liver’s ability to detoxify it into urea. This tainability is by developing and using effective rumi-
can kill the animal. However, with sufficient energy, nant livestock grazing systems. More than 60 percent
microbes use ammonia and amino acids to grow and of the land area in the world is too poor or erodible for
reproduce. cultivation but can become productive when used for
The rumen does not degrade the UIP component ruminant grazing. Ruminant livestock can use land for
of feedstuffs. The UIP “bypasses” the rumen and grazing that would otherwise not be suitable for crop
makes its way from the omasum to the abomasum. In production. Ruminant livestock production also com-
the abomasum, the ruminant uses UIP along with plements crop production, because ruminants can use
microorganisms washed out of the rumen as a protein the byproducts of these crop systems that are not in
source. Protein as a nutrient in ruminant diets is dis- demand for human use or consumption. Developing a
cussed in detail in Mississippi State University good understanding of ruminant digestive anatomy
Extension Service Publication 2499, “Protein in Beef and function can help livestock producers better plan
Cattle Diets.” appropriate nutritional programs and properly man-
age ruminant animals in various production systems.
Importance of Ruminant Livestock
The digestive system of ruminants optimizes use of
rumen microbe fermentation products. This adaptation
lets ruminants use resources (such as high-fiber forage) References
Church, D. C. ed. 1993. The Ruminant Animal Digestive Physiology and Nutrition. Waveland Press, Inc. Prospect Heights, IL.
Oltjen, J. W., and J. L. Beckett. 1996. Role of ruminant livestock in sustainable agricultural systems. J. Anim. Sci. 74:1406-1409.
Parish, J. A., M. A. McCann, R. H. Watson, N. N. Paiva, C. S. Hoveland, A. H. Parks, B. L. Upchurch, N. S. Hill, and J. H. Bouton. 2003. Use
of non-ergot alkaloid-producing endophytes for alleviating tall fescue toxicosis in stocker cattle. J. Anim. Sci. 81:2856-2868.
Van Soest, P. J. 1987. Nutritional Ecology of the Ruminant. Cornell University Press. Ithaca, NY.

Photographs of the ruminant digestive system are courtesy of Dr. Stephanie R. Hill, assistant research professor, Animal and Dairy Sciences,
Mississippi State University.
Copyright 2009 by Mississippi State University. All rights reserved. This publication may be copied and dis-
tributed without alteration for nonprofit educational purposes provided that credit is given to the
Mississippi State University Extension Service.

By Dr. Jane A. Parish, Associate Extension/Research Professor; Dr. J. Daniel Rivera, Assistant Extension
Professor; and Dr. Holly T. Boland, Assistant Research/Extension Professor

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Publication 2503
Extension Service of Mississippi State University, cooperating with U.S. Department of Agriculture.
Published in furtherance of Acts of Congress, May 8 and June 30, 1914. MELISSA J. MIXON, Interim Director
POD 08-09