17/6/23, 10:58 Rumen Physiology and Rumination

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of Herbivores

Rumen Physiology and Rumination

The rumen is a fermentation vat par excellance, providing an anaerobic
environment, constant temperature and pH, and good mixing. Well-masticated
substrates are delivered through the esophagus on a regular schedule, and
fermentation products are either absorbed in the rumen itself or flow out for
further digestion and absorption downstream.

Ruminants evolved to consume and subsist on roughage - grasses and shrubs

built predominantly of cellulose. Despite the fact that some ruminants, feedlot
steers for example, are feed large quantities of grain, this section will focus on a
ruminant's "natural diet".

Dynamics of Cranial Digestion

Feed, water and saliva are delivered to the reticulorumen through the esophageal
orifice. Heavy objects (grain, rocks, nails) fall into the reticulum, while lighter
material (grass, hay) enters the rumen proper. Added to this mixture are
voluminous quantities of gas produced during fermentation.

Ruminants produce prodigious quantities of saliva. Published estimates for adult

cows are in the range of 100 to 150 liters of saliva per day! Aside from its normal
lubricating qualities, saliva serves at least two very important functions in the
ruminant:

provision of fluid for the fermentation vat

alkaline buffering - saliva is rich in bicarbonate, which buffers the large
quanitity of acid produced in the rumen and is probably critical for
maintainance of rumen pH.

All these materials within the rumen partition into three primary zones based on
their specific gravity. Gas rises to fill the upper regions, grain and fluid-saturated
roughage ("yesterday's hay") sink to the bottom, and newly arrived roughage
floats in a middle layer.

The rate of flow of solid material through the rumen is quite slow and dependent
on its size and density. Water flows through the rumen rapidly and appears to be
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critical in flushing particulate matter downstream.

As fermentation proceeds, feedstuffs are reduced to smaller and smaller sizes

and microbes constantly proliferate. Ruminal contractions constantly flush lighter
solids back into the rumen. The smaller and more dense material tends to be
pushed into the reticulum and cranial sac of the rumen, from which it is ejected
with microbe-laden liquid through the reticulo-omasal orifice into the omasum.

The function of the omasum is rather poorly understood. It may function to absorb
residual volatile fatty acids and bicarbonate. The tendency is for fluid to pass
rapidly through the omasal canal, but for particulate matter to be retained
between omasal leaves. Periodic contractions of the omasum knocks flakes of
material out of the leaves for passage into the abomasum.

The abomasum is a true, glandular stomach which secretes acid and otherwise
functions very similarly to the stomach of a monogastric. One fascinating
specialization of this organ relates to its need to process large masses of
bacteria. In contrast to the stomach of non-ruminants, the abomasum secretes
lysozyme, an enzyme that efficiently breaks down bacterial cell walls.

The processes described above apply to adult ruminants. For the first month or
so of life, the ruminant is functionally a monogastric. The forestomachs are
formed, but are not yet fully developed. If milk is introduced into such a rumen, it
basically rots rather than being fermented. To avoid this problem in such young
ruminants, suckling causes a reflex closure of muscular folds that form a channel
from the esophageal orifice toward the omasum (the esophageal groove),
shunting milk away from the rumen and straight toward the stomach where it can
be curdled by rennin and eventually digested enzymatically.

Reticuloruminal Motility

An orderly pattern of ruminal motility is initiated early in life and, except for
temporary periods of disruption, persists for the lifetime of the animal. These
movements serve to mix the ingesta, aid in eructation of gas, and propel fluid and
fermented foodstuffs into the omasum. If motility is suppressed for a significant
length of time, ruminal impaction may result.

A cycle of contractions occurs 1 to 3 times per minute. The highest frequency is

seen during feeding, and the lowest when the animal is resting. Two types of
contractions are identified:

Primary contractions originate in the reticulum and pass caudally around the
rumen. This process involves a wave of contraction followed by a wave of
relaxation, so as parts of the rumen are contracting, other sacs are dilating.
Secondary contractions occur in only parts of the rumen and are usually
associated with eructation.

The animation below is based on data collected by radiographing sheep (Wyburn,

1980) and should impart at least some appreciation of the complexity of ruminal
motility. Although shown much faster than in life, the major reticuloruminal
contractions are timed appropriately. Note the movements which bring the gas
bubble (stippled area) forward to the esophagus for eructation.

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What about control of rumen motility? The forestomachs possess a rich enteric
nervous system, but coordinated contractions require central input. Motility
centers in the brainstem control both the rate and strength of contraction via vagal
efferents. Cutting the vagus nerve in a ruminate abolishes coordinated
reticuloruminal motility. There are also vagal afferents from the rumen to the
motility centers which allow stretch receptors and chemoreceptors in the rumen to
modulate contractility.

Conditions inside the rumen can significantly affect motility. If, for example,
ruminal contents become very acidic (as occurs in grain engorgement), motility
will essentially cease. Also, the type of diet influences motility: animals on a high
roughage diet have a higher frequency of contractions than those on a diet rich in
concentrates.

Rumination and Eructation

Ruminants are well known for "cud chewing". Rumination is regurgitation of

ingesta from the reticulum, followed by remastication and reswallowing. It
provides for effective mechanical breakdown of roughage and thereby increases
substrate surface area to fermentative microbes.

Regurgitation is initiated with a reticular contraction distinct from the primary

contraction. This contraction, in conjunction with relaxation of the distal
esophageal sphincter, allows a bolus of ingesta to enter the esophagus. The
bolus is carried into the mouth by reverse peristalsis. The fluid in the bolus is
squeezed out with the tongue and reswallowed, and the bolus itself is
remasticated, then reswallowed.

Rumination occurs predominantly when the

animal is resting and not eating, but that is
a considerable fraction of the animal's
lifespan. The chart shown here (adapted
from Lofgreen et al., J Animal Sci 16:773,
1957) depicts how steers spend their day
on an alfalfa pasture relative to time spent
grazing and ruminating.

Fermentation in the rumen generates

enormous, even frightening, quantities of
gas. We're talking about 30-50 liters per
hour in adult cattle and about 5 liters per hour in a sheep or goat. Eructation or
belching is how ruminants continually get rid of fermentation gases. As mentioned
above, an eructation is associated with almost every secondary ruminal
contraction. Eructated gas travels up the esophagus at 160 to 225 cm per second
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(Stevens and Sellers, Am J Physiol 199:598, 1960) and, interestingly, a majority is

actually first inspired into the lungs, then expired.

Anything that interferes with eructation is lifethreatening to the ruminant because

the expanding rumen rapidly interferes with breathing. Animals suffering ruminal
tympany (bloat) die from asphyxiation.

Rumen gases, particularly methane, are increasingly in the news because of their
contribution to greenhouse gas and climate change. As with most topics that
attract the attention of activists and politicians, it is somewhat difficult to obtain
accurate estimates of the contribution of ruminant digestive processes to global
greenhouse gas. However, data from the Food and Agriculture Organization of
the United Nations indicates that ruminants are responsible for roughly 20% of
global methane emmisions, which equates to approximately 3-5% of total
greenhouse gas production.

Digestive Anatomy of Ruminants Nutrient Absorption and Utilization in

Ruminants

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