The Professional Animal Scientist 30 (2014):129–139

©2014 American Registry of Professional Animal Scientists

Iof ruminants:
R
NVITEd : Applied nutrition
Fermentation and
EVIEW

digestive physiology1
C. R. Krehbiel,2 PAS
Department of Animal Science, Oklahoma State University, Stillwater 74078

ABSTRACT
Since the first observation of the 4
chambers and subsequent identification
of its microbial population, the rumen
has been investigated for its role in
nutrient digestion and to manipulate its
microbial ecosystem to increase animal
performance and efficiency. Ruminants
have the ability to digest plant poly-
saccharides through substrate-specific
enzyme activities of the highly specific
population of ruminal bacteria, proto-
zoa, and fungi. A synergistic relation-
ship provides these microorganisms with
nutrients, desirable ambient temperature,
and a buffered environment to enhance
microbial growth, and the microbes
provide the host animal with B vitamins,
VFA, and microbial protein. Microbial
populations are not static and fluctuate
with changes in the ruminal environment
and diet, resulting in changes in VFA
composition and concentration. Recent
research using terminal restriction
fragment length polymorphism analy-
sis, 16S ribosomal DNA libraries, and
metagenomic approaches has increased
1
Presented at the American Registry of
Professional Animal Scientists (ARPAS)
Symposium: Applied Nutrition of
Ruminants—Current Status and Future
Directions, July 10, 3013, ADSA/ASAS Joint
Annual Meeting, Indianapolis, Indiana.
2
Corresponding author: clint.krehbiel@
okstate.edu
our understanding of the importance of
microbial communities within the rumen
and the microbial diversity under dif-
ferent dietary environments. Enhanced
understanding of these changes in micro-
bial communities could result in methods
to enhance microbial growth rates and
enzyme secretions in the rumen, result-
ing in increased digestibility, nutrients
delivered to the small intestine, and im-
proved animal production and efficiency.
In addition, increased understanding of
molecular-level adaptation of ruminal
epithelia and gastrointestinal mucosa
could provide the physiological basis for
their role in regulation of ruminal pH
and nutrient transport. Recent advances
and continued research in the area of
ruminal fermentation and digestive
physiology have the potential to positively
affect animal production, health, and the
environment.
Key words: digestive physiology,
ruminal fermentation, ruminant nutri-
tion
INTRODUCTION
Ruminant production systems and
diets fed to ruminants are diverse.
Understanding form and function of
the rumen is important because ru-
men dynamics are ultimately respon-
sible for providing nutrients to the
host animal. In adaptive terms, the
ruminant animal has a digestive tract
that allows digestion of fibrous mate-
rials, allowing survival without energy
or protein for several days. In harvest-
ing forage, the ruminant uses the larg-
est carbohydrate source in the world
(i.e., cellulose and hemicellulose),
and produces food and other useful
products for man (Church, 1988). The
utilization of fiber by ruminants is a
result of enzymes generated by the
anaerobic (and facultative) bacteria,
protozoa, and fungi in the rumen.
Anaerobic fermentation, as occurs in
the rumen, is often viewed as being
inefficient because only 2 to 6 ATP
are generated per mole of glucose
fermented as opposed to 38 ATP
from aerobic metabolism. Nonethe-
less, because the system is anaerobic,
the microbes extract oxygen from
substrates and excrete energy-rich
compounds that provide the majority
of the metabolizable energy for the
host animal. Therefore, the microbes
extract enough energy to grow and
ferment and still leave energy for the
animal to use for physiological pro-
cesses. Pregastric fermentation offers
some unique advantages to the host
animal: 1) it provides energy from
fibrous materials; 2) the microbes
synthesized provide amino acids; and
3) microbial fermentation provides
B-complex vitamins. There are also
disadvantages associated with pregas-
tric fermentation: 1) readily available
substrates such as starch, sugar, and
130
proteins are used less efficiently than
in the small intestine; 2) biologi-
cal value of high-quality proteins is
decreased; and 3) fermentation is not
always well-controlled, resulting in the
potential for metabolic dysfunction
(e.g., ruminal acidosis).
It is interesting to note that similar
to our domestic production systems,
ruminants in nature can be divided
into 3 groups based on their feeding
preferences—concentrate selectors,
intermediate feeders, and bulk and
roughage eaters (Hofmann, 1988).
These 3 main categories depict dif-
ferences between general food types,
whereas divisions within the bulk and
roughage eaters are made according
to water availability in the forage.
Cattle are the most developed and
the most unselective grazers, perhaps
because the temperate plants cattle
consumed during their evolution are
less differentiated than tropical plants
grazed by more selective ruminants.
This paper provides a review of the
anatomy of the ruminant stomach
compartments and its components. A
discussion of animal, diet, and micro-
bial factors involved in rumen func-
tion and dysfunction follows.
COMPONENTS OF THE
RUMINANT STOMACH
Anatomy
The stomach of the ruminant
animal is composed of the reticulum,
rumen, omasum, and abomasum
(Hofmann, 1988). The reticulum and
rumen are joined by a fold of tissue
called the reticulo-ruminal fold. The
reticulo-rumen is a large fermentation
vat that houses 109 to 1011 bacteria,
104 to 107 protozoa, and 103 to 105
fungi per milliliter of ruminal fluid
(Qi et al., 2011; Krause et al., 2013).
The reticulo-rumen occupies most
(>3/4) of the left side of the ab-
dominal cavity, except a small area
occupied by the spleen and small
intestine. The left side resides dorsal
to the diaphragm, abdominal wall,
and spleen, whereas the right side is
Krehbiel
more irregular and resides dorsal to
the liver, left kidney, omasum, aboma-
sum, pancreas, aorta, and posterior
vena cava. Peritoneum and connective
tissue firmly attach this organ to the
diaphragm and sublumbar muscles.
The reticulum rests against the dia-
phragm on the left side of the median
plane and is connected to the omasum
by a short narrow neck called the
reticulo-omasal orifice. The omasum
is spherical and lies on the right side
of the median plane. The abomasum
is an elongated sac that lies chiefly
on the abdominal floor, ventral to the
frontal plane.
The grooves on the outside of the
reticulo-rumen separate the organ into
sacs. These grooves correspond to pil-
lars internally. The most conspicuous
is the groove separating the reticulum
from the cranial portion of the ru-
men. Dorsally there is no separation
between them, and they join at the
entrance of the esophagus to form the
cardia. Although the reticulo-ruminal
groove is prominent, the opening be-
tween the 2 is large and makes the re-
ticulum resemble an anterior pouch of
the rumen. The reticulum and rumen
together form a large functionally in-
tegrated sac filled with microbes and
digesta. The interior of the reticulo-
ruminal compartments also contains
structures that aid in the functioning
of each part. These pillars, men-
tioned earlier, are muscles that aid
in movement and contraction of the
rumen, which promote mixing and
physical breakdown of food particles.
When contracted, the pillars are well
defined, but when relaxed they are
not visually prominent. The cardia is
the terminal end of the esophagus and
is also the beginning of the ventricu-
lar (or reticular) groove. The groove
begins at the cardia and extends 18 to
20 cm (7 to 8 in) in cattle, ending at
the reticulo-omasal orifice.
Structure of the 4
Compartments
The outer surface of all 4 stomach
compartments is coated by visceral
serosa with the exception of an area
of variable size on the dorsal ruminal
sac. There is a relatively large area of
fibrous attachment to the sublumbar
muscles and the diaphragm, which
allows for the contraction of the unat-
tached dorsal sac. There are 3 layers
to all compartments: 1) outer—serous
membrane; 2) middle—muscular tunic
containing thin superficial and thick
deep muscle (these 2 layers run in op-
posite directions); and 3) inner—epi-
thelial membrane.
The reticulum is easily distinguished
internally by its unique honeycomb
structure. The reticular epithelia are
raised into folds forming tissue of 4, 5,
or 6 sides. The epithelia are arranged
in intersecting crests that separate
cellulae reticuli. Small, vestigial papil-
lae are present in this honeycomb
structure. The dorsal portion of the
reticular epithelia blends through the
reticulo-rumen. The rumen does not
contain honeycomb structures but
rather numerous papillae of various
shapes and sizes. These ruminal papil-
lae can be defined as organs of ab-
sorption. The papillae are particularly
well developed in the ventral sac and
in the floor of the caudo-dorsal blind
sac, as well as on the cranial groove
and adjacent surfaces. In all rumi-
nants, the papillae are most numerous
and longest in the cranial sac followed
by the caudo-dorsal blind sac. Papil-
lae are typically 1 to 2 mm in length
in the dorsal sac and 1 cm in length
in the ventral sac. Size, number, and
distribution are related to feeding
behavior and diet selection. The great
absorptive surface provided by these
papillae permits ready movement of
solutes able to traverse the epithelial
cells separating the stomach contents
from the blood supply.
The omasum is also called the
manyplies because of the numer-
ous parallel sheets of tissue within it
(Becker et al., 1963). These omasal
leaves (Laminae omasi) are of various
sizes and bend to the wall of the oma-
sum. The free edges of the leaves ex-
tending to the channel lie against the
muscular pillar of the omasal groove,
an extension of the ventricular groove,
along the lesser curvature of the oma-
 Ruminal fermentation and digestive physiology
sum from the reticulo-omasal orifice
to the opening into the abomasum.
The free edges of the leaves contain
conical papillae. The omasum is much
larger in grazing animals than in con-
centrate selectors, and the omasum is
the bottleneck for food passage from
the reticulo-rumen. It lies slightly to
the right of the reticulo-rumen in the
abdominal cavity and extends caudal
from the 7th to the 11th rib. The
omasum connects the reticulo-omasal
orifice to the omasal-abomasal orifice.
The omasal-abomasal orifice is 3 to 4
times larger than the reticulo-omasal
orifice. The function of the omasum is
not well understood. It is known that
the Laminae omasi provide an absorp-
tive surface for H2O, Na+, K+, and
some VFA (Engelhardt and Hauffe,
1975). Fifteen percent of the water
that enters the omasum is absorbed
there. The omasum also catches and
separates particles, thereby regulat-
ing passage of particles entering this
compartment.
The tubular abomasum has an in-
ternal surface of folded spiraled ridges
that are flaps composed of epithelial
tissue. This organ is different from the
previous 3 in that it contains secre-
tory tissue. The glandular gastric
mucosa is supplied with specialized
secretory cells that produce mucus,
pepsinogin, and HCl. The fundus
is the initial portion that contains
gastric glands (HCl-producing pari-
etal cells). The pyloric region con-
tains thick glandular mucosa and the
pepsin-secreting chief cells. The histol-
ogy and physiology in the abomasum
is similar to that of the nonruminant
stomach.
Cell Types
The cells that line the reticulo-
rumen and the omasum are keratin-
ized, stratified, squamous epithelium.
They are not glandular nor do they
produce mucus. Four cell layers can
be distinguished in ruminal epithelia:
stratum basal, stratum spinosum,
stratum granulosum, and stratum
corneum (Steven and Marshall, 1970;
Graham and Simmons, 2005). These
layers seem to merge into each other
without distinct boundaries. At the
ultrastructural levels, however, only
the middle layers are indistinct with
clear definition of the stratum basal
and stratum corneum. Cells of the
stratum basal have large numbers of
ribosomes, mitochondria, and Golgi
vesicles, as well as ovoid nuclei. These
cells are perpendicular to the base-
ment membrane. Also evident in these
cells are free ribosomes and rough en-
doplasmic reticulum, which together
with the mitochondria and other cell
organelles are involved with assimila-
tion and metabolism of products (e.g.,
VFA) absorbed from the rumen.
Ruminal parakeratosis can be
defined as incomplete keratinization.
Parakeratosis results from increased
proliferation and outward migration
of the cells of the stratum basale and
has been associated with clumping of
the epithelial cells. This can ultimate-
ly decrease the surface area available
for nutrient absorption. Parakeratosis
is often attributed to increased VFA
or low ruminal pH.
Blood Supply and Motility
Blood supply to and from the ru-
men is extremely important physi-
ologically because of the tremendous
absorption that takes place in this
organ. Blood flow and surface area of
the rumen determine absorption rate.
Blood flow of nutrients is not unidi-
rectional. For example, urea diffuses
from the blood to the rumen when
ruminal concentrations of ammonia
are low. The ruminant stomach com-
plex receives its entire arterial supply
from the celiac-cranial mesenteric
trunk, which arises from the common
trunk of the abdominal aorta in sheep
or separate branches of the aorta in
cattle. The 4 major branches of the
celiac artery are 1) common hepatic
artery, which services the pancreas,
liver, gall bladder, and branches into
the gastroduodenal artery; 2) right
ruminal artery, which branches to
the pancreas and omentum; 3) left
ruminal artery, which branches to the
reticulum and esophagus; and 4) left
131
gastric artery, which branches to the
omasum and abomasum.
Blood flow from the reticulo-rumen
is by the right and left ruminal veins
that drain into the gastrosplenic and
gastroduodenal veins. The gastro-
splenic and gastroduodenal veins also
receive blood from the omasal-aboma-
sal and reticular veins. Volatile fatty
acids and other water-soluble nutri-
ents (e.g., ammonia) absorbed from
the rumen are transported directly
to the liver. The hepatic portal vein
contributes approximately 80% of the
blood flowing to the liver.
Nerve Supply
The vagus nerve supplies the gastro-
intestinal tract. Cattle have 12 cranial
nerves with the vagus being the 10th
one. The vagus nerve originates from
the medulla oblongata. The dorsal
trunk of the vagus serves the rumen,
whereas the ventral trunk serves the
omasum, abomasum, and reticulum.
The vagus aids in controlling gut
motility by regulating muscle contrac-
tion.
RUMINAL CONTENTS
As indicated, the microbial popu-
lation in the rumen consists of the
prokaryotic bacteria and archaebac-
teria (methane producers) and the
eukaryotic protozoa and fungi (Qi et
al., 2011; Krause et al., 2013). Bac-
teria are grouped according to their
cell-wall ultrastructure (gram negative
and gram positive), their 3 main mor-
phologies (cocci, rods, and spirilla),
and according to their size (generally
from 0.3 to 50 μm). They are also
grouped according to the type of sub-
strate fermented, including plant-cell-
wall degraders (cellulolytics, xylano-
lytics, pectinolytics), starch degraders
(amylolytics), protein degraders
(proteotylics), lipid degraders (lipolyt-
ic), lactate users, methanogens, and
detoxifiers. Most species of bacteria
are capable of fermenting more than
one substrate (Russell, 1984). Isolat-
ing and culturing ruminal microor-
ganisms has traditionally been used
132
to study the ruminal ecosystem (e.g.,
Bryant and Burkey, 1953; Hungate,
1966; Stewart and Bryant, 1988).
More recently, culture-independent
molecular methods (whole-genome se-
quencing, pyrosequencing, proteomics,
and transcriptomics) have been used
to study the rumen microbiome and
their functional genes (Krause et al.,
2013). Molecular procedures show
promise for characterizing the rumen
microbiome as influenced by ani-
mal, diet, and the environment and
thereby enhancing our understanding
of the complexity of this ecosystem
(Krause et al., 2013).
The methane-producing bacteria
(archaebacteria) are responsible for
regulating the overall fermentation
in the rumen. They remove hydrogen
gas by reducing carbon dioxide to
form methane. Producing methane
keeps the partial pressure of hydro-
gen in the rumen low, which allows
methanogenic bacteria to promote
the growth of other bacterial species
and provides for a more efficient fer-
mentation (Janssen and Kirs, 2008).
The effective removal of hydrogen by
these methanogenic species encour-
ages hydrogen-producing species to
produce more hydrogen and thus alter
their metabolism toward higher yield-
ing pathways. These higher yielding
pathways result in the synthesis of
more microbial cells, which increases
microbial cell protein available to the
ruminant.
The protozoa in the rumen are
influenced by feeding practices, and
the effects of defaunation have been
reviewed by Jouany et al. (1988).
Both ciliated and flagellated protozoa
exist in the rumen (Bohatier, 1991),
but the majority of the protozoa are
ciliates (104 to 106 cells/mL), with
over 100 species from about 25 genera
represented (Williams, 1986; Qi et al.,
2011). Different types of diets seem to
encourage different genera of proto-
zoa. For example, the population of
some protozoa are greater when diets
contain large amounts of soluble sug-
ars (e.g., holotrichs), whereas other
types predominate with high-starch
diets (e.g., entodiniomorphs). Pro-
Krehbiel
tozoa, similar to bacteria and fungi,
also contribute to fiber digestion. The
protozoa actively ingest bacteria as a
source of N, and thereby are respon-
sible for intraruminal N recycling
(Firkins and Yu, 2006). Protozoa
are an integral part of the microbial
population and have a marked effect
on the fermentation; however, their
benefit to the ruminant is still con-
troversial, especially when high-grain
diets are fed.
The anaerobic fungi are the most
recently recognized group of rumen
microbes (Orpine, 1993; Stewart et
al., 1995). Six genera have been iden-
tified, accounting for 8 to 20% of the
total rumen microbial biomass (103
to 105 zoospores/mL) when animals
are fed a high-forage diet. Anaerobic
fungi have been shown to degrade
cellulose and xylans (Krause et al.,
2013) and contribute substantially to
fiber degradation in the rumen. They
colonize plant material by penetrat-
ing their mycelia deep inside particles
and they possess a range of hydrolytic
polysaccharidase and complementary
glycosidase activities (Stewart et al.,
1995). They colonize lignified regions
of the plant cell wall, which facilitates
bacterial colonization.
There are 4 interactive compart-
ments in which the microbes are lo-
cated in the rumen (Cheng and McAl-
lister, 1997). The first is defined as
the adherent compartment, where the
microbial groups are attached to feed
and degrade insoluble polysaccharides,
as well as the less-soluble proteins.
The second compartment consists of
the microorganisms that are associat-
ed with feed particles. These microor-
ganisms degrade oligopolymers of car-
bohydrate and protein. Compartments
1 and 2 compose approximately 75%
of the microbial mass. The third com-
partment is the fluid compartment,
where free-floating microbial groups
in ruminal fluid feed on monomers of
soluble carbohydrates (e.g., glucose)
and protein (e.g., ammonia and amino
acids). This compartment makes up
25% of the microbial mass. The final
compartment (<1%) consists of the
microbes that are attached to the
rumen epithelial cells or to protozoa.
These microorganisms include facul-
tative microbes that digest urea or
protein from sloughed epithelial cells.
Ruminal contents are highly vari-
able in particle size and composition.
Different particle lengths and densi-
ties are found in different areas in the
rumen. The dorsal rumen consists of a
gas dome. Typical gases found in the
rumen include hydrogen, 0.2%; oxy-
gen, 0.5%; nitrogen, 7.0%; methane,
26.8%; and carbon dioxide, 65.5%
(Sniffen and Herdt, 1991). Long hay
particles, which are typically lower in
density than water, tend to float and
form a mat or raft layer above the
liquid layer and under the gas dome.
Smaller particles will be suspended in
the liquid layer and can be entrapped
in the mat. Heavier particles, such
as grain, tend to sink to the ventral
rumen.
RUMEN FUNCTION
The importance of ruminal fermen-
tation for supplying nutrients to the
host animal has long been recognized.
Many attempts have been made to
biologically or mathematically model
these processes (Baldwin et al., 1987;
Dijkstra et al., 1992; Sauvant et al.,
1996; Bannink et al., 1997; Huhtanen
et al., 2006). Although none of these
models are perfect, they allow for crit-
ical evaluation of the current concepts
and available data and identification
of areas for needed research. Models
are built using inputs and outputs
from data involving rumen physiol-
ogy. For example, rumen metabolism
depends on the input (i.e., intake) of
nutrients, such as carbohydrates, pro-
teins, minerals, and vitamins. Carbo-
hydrate sources include soluble sugars,
starch, pectin, cellulose, and hemicel-
lulose, whereas protein sources consist
of NPN, soluble protein, degradable
intake protein, and undegradable in-
take protein. In the rumen, microbial
pools representing the 3 major types
of microorganisms can be distin-
guished by their substrate preference,
as previously discussed. Outputs from
the rumen include absorption of end
 Ruminal fermentation and digestive physiology 133

products and passage through the
reticulo-omasal orifice. The follow-
ing paragraphs briefly discuss these
factors and other components that are
involved in rumen function.
Diet Factors
Diet characteristics affecting ru-
minal function include chemical and
physical characteristics that ultimate-
ly influence intake, digestion, and rate
of passage. In the model of Dijkstra et
al. (1992), the amount of dietary nu-
trients entering the ruminal pool was
derived from the chemical description
of the diet and the amount consumed
by the animal. In addition to the car-
bohydrate and N fractions, the model
also included lipids and organic acids
(lactate, acetate, propionate, butyr-
ate, and valerate). Organic acids were
included to accommodate fermented
feeds such as silage (Dijkstra et al.,
1992). The soluble and insoluble,
degradable and undegradable compo-
nents of dietary protein and carbohy-
drates can be determined by the ny-
lon-bag, in situ technique (Nocek and
Grant, 1987; Tamminga et al., 1990).
During the in situ method, feed is
incubated in small porous bags in the
rumen of ruminally cannulated cattle,
and DM and component disappear-
ance from the bags is measured over
time (Vanzant et al., 1998; Ørskov,
2000). The time course usually spans
from a zero-time measurement to
determine initial solubility to a final
measurement at 48 or 72 h for rapidly
digested components (e.g., sugar and
soluble N) and 72 or 96 h for slowly
digested components (e.g., fiber) to
estimate the indigestible fraction
(Nocek and English, 1986). Rate of
passage (kp) is determined in vivo by
calculating recovery of an external
or internal marker, and is reported
as the outflow from the rumen (g/h,
mL/h) divided by pool size (total
mL or g) in the rumen. Passage rates
and ruminal volume as influenced by
diet can be estimated using single- or
multicompartmental models. These
models have been discussed in detail
by others (Owens and Goetsch, 1986;
Mertens, 1993; Ellis et al., 1994).
Animal Factors
Several experiments (Grenet, 1989;
Sauvant et al., 1996) have demon-
strated a positive relationship be-
tween rate of intake within meals
and daily voluntary DMI. Sauvant et
al. (1996) estimated potential intake
rate (PIR) as PIR = [16.8 − 0.14 ×
NDF × proportion of large particles
in feed (PLPF)] × palatability. The
product of NDF and PLPF results
from a theoretical consideration of the
interaction between cell-wall content
and particle size. Data summarized by
Sauvant et al. (1996) showed a posi-
tive curvilinear relationship between
the percentage of large particles in
the diet and the percentage of large
particles in the masticate. In other
words, as particle length in the diet
increased, particle length in the mas-
ticate increased at a diminishing rate,
suggesting more comminution during
chewing. This might be important for
subsequent rumination and the rate
at which particles exit the reticulo-
rumen.
It is generally believed that only
enough chewing is done initially to
mix the food with saliva. The food
and saliva mixture form a bolus,
which is projected down the esopha-
gus and deposited in the reticulum.
The form, weight, and consistency of
the bolus depend on the type of food
being eaten. Boluses made from for-
age material will float in water, which
aids in their regurgitation. Boluses
from grain or concentrate feeds are
heavier and have a tendency to sink
in the ruminal fluid. During initial
chewing, mixed saliva (contains both
serous and mucous types of saliva) is
secreted, which aids in lubrication and
protection of the mouth and esopha-
gus. During rumination, more serous
saliva is secreted (parotid gland).
This saliva serves as a bicarbonate,
phosphate buffer with the 2 anions
comprising more than 90% of total
anion content and is strongly buffered
against acid production; however,
amounts of bicarbonate and phos-
phate secreted are often not sufficient
to buffer acids produced during grain
fermentation. Physical and chemical
characteristics of the feeds, such as
moisture content, NDF content, and
particle size determine the amount of
saliva produced daily. The amount of
saliva added to concentrates dur-
ing eating is much lower (0.76 to
1.12 mL of saliva/g of DM) than for
forages (3.40 to 7.23 mL of saliva/g
of DM; Bailey, 1961; Beauchemin et
al., 2008). In addition, among for-
ages there is about a 3-fold difference
in saliva production between silage
and hay on an as-is but not on a DM
basis (Beauchemin et al., 2008). Feed
characteristics influence total daily
salivary secretion mainly by affecting
time spent eating and ruminating,
rather than the salivation rate (mL/
min) directly. Greater ensalivation of
feed occurs as eating time and rumi-
nation increase (Beauchemin et al.,
2008). Considerably less saliva is pro-
duced when cattle are fed grain diets
compared with forage-based diets.
Mixing Cycles. Ruminant animals
depend on several mixing cycles for
normal rumen function. The cycles
are important for mixing contents
(primary cycle), eructation of fermen-
tation gases (secondary contractions),
rumination, and passage of contents
through the reticulo-omasal orifice.
Normal and abnormal motor function
associated with these cycles has been
extensively reviewed (Ruckebusch,
1988; Constable et al., 1990a,b).
Primary contractions are denoted by
2 contractions of the reticulo-ruminal
fold followed by a wave of contrac-
tions that pass caudally over the
rumen. Different environmental and
physiological situations affect ruminal
motility (Ruckebusch, 1988). Rate of
ruminal contractions during sleeping
are lower than resting states but are
still apparent and readily distinguish-
able. Eating causes an increase in the
rate of ruminal contraction. Cattle
that were fed increased the rate of
contraction from 60 to 105 cycles/h,
whereas sheep showed increases to 3
cycles/min after feeding.
Secondary contractions are usually
associated with eructation and may or
may not occur after primary con-
tractions (Ruckebusch, 1988). They
generally involve contraction of the
134
dorsal coronary pillar, contraction of
the caudo-dorsal blind sac and dorsal
sac, and relaxation of the caudo-
ventral blind sac. The longitudinal
and cranial pillars are probably also
involved. The contraction lasts about
30 s. The normal ratio of primary to
secondary contractions in domesticate
ruminants is 1:1. Variations of 2:1 to
1:2 can be found within a population
of animals. Receptors for these con-
tractions are located in the reticulo-
rumen complex. The tension receptors
are located in the contractile elements
of the muscle layers, particularly in
the reticulum and the cranial sac of
the rumen. Their densest distribution
is in the area of the reticular groove.
The reticular tension receptors have
an overall excitatory effect on the
rate, amplitude, and form of primary
contractions. They are associated
with nerve fibers having a mean con-
duction velocity of 12.5 m/s. These
receptors might also be concerned
with reflex-induced salivation and
with monitoring ruminal fill. Other
receptors have been identified in the
reticulum, cranial sac, around the
cranial and longitudinal pillars, and
other parts of the rumen. The recep-
tors are located near the basement
membrane of the ruminal epithelia.
The epithelial receptors are excited by
acids, alkalis, hyper- and hypoosmotic
solutions, and distension. The effi-
ciency of stimulation by acids seems
related to its strength (dissociation
constant), molecular, and titratable
acidity but not to pH. The lower-mo-
lecular-weight organic acids are more
effective.
Rumination and Eructation.
Rumination or regurgitation is initi-
ated by an extrareticular contraction
that precedes a primary and second-
ary contraction (Ruckebusch, 1988).
The contraction produces measurable
pressure changes in the reticulum.
These contractions are thought to
respond to pressure changes in the
rumen. Rumination is composed
of regurgitation, swallowing regur-
gitated liquids, remastication of a
bolus, reinsalivation, and reswallow-
ing the bolus. The reticular contrac-
tion causes a marked increase in fluid
Krehbiel
pressure in the cardial area of the
rumen. Then a sharp contraction of
the diaphragm causes a negative pres-
sure that aspirates reticular contents.
The aspirated slurry is moved up the
esophagus by antiperistaltic contrac-
tions (100 cm/s). The rapid contrac-
tions take place because unlike many
other mammals, the esophagus of the
ruminant contains striated muscle.
Rumination does not seem to be
triggered by VFA, urea, or pH in
ruminal fluid but is more closely re-
lated to the bulk of the diet. Grinding
forages effectively mimics rumination
in that it decreases the mean reten-
tion time spent in the rumen, and
hence there is a decrease in DM, OM,
and NDF digestibility. This does not
mean that rumination is a negative
effect, but rather a positive effect on
rumen emptying, rate of passage, and
increasing intake. This phenomenon
is very critical on low-quality forage
diets. Increasing removal of materi-
als from the rumen allows greater
intake of low-quality roughages that
enables an animal to maintain itself
on a poor-quality diet. This could be
a mechanism of preservation (Welch,
1982). Rumination could be a preser-
vation mechanism in another sense.
Ruminants can eat rapidly during
the daylight hours and chew leisurely
at night. This means the animal will
spend less time grazing out in the
open where predators have an advan-
tage and can chew its food at a later
time at a safe place.
Rumination is the only way particle
size is decreased in the rumen. Mas-
tication during eating and rumina-
tion differs. Chewing during eating
is primarily responsible for preparing
food for swallowing, releasing soluble
constituents, and damaging plant tis-
sues for microbial digestion, whereas
rumination decreases particles so that
material can pass from the reticulo-
rumen (Ulyatt et al., 1986). There-
fore, rumination, and not chewing,
is the principle means of decreasing
food particle size. Although microbial
digestion in the rumen contributes to
decreasing the dry weight of particles,
its direct contribution to decreasing
particle length is small (<20%). It
mainly serves to weaken the cell-wall
structure so that particle breakdown
during rumination is facilitated.
Therefore, digestion of the fiber
does not decrease particle size per
se but does make the cell wall more
fragile and break down more easily
when chewed. Rumination also helps
maintain a more uniform amount of
ingesta in the gastrointestinal tract
over a 24-h period. It might also help
maintain a stable microbial popu-
lation because of a more constant
source of substrates.
Passage Through the Reticulo-
Omasal Orifice. The proportion of
feed digested is hypothesized to be a
balance between digestion rate and
the passage of undigested feed from
the reticulo-rumen (Waldo and Smith,
1972). This implies that both micro-
bial and animal factors are involved.
Particulate passage rate has a nega-
tive relationship with extent of diges-
tion and in many cases total-tract
digestibility (i.e., the faster the mate-
rial passes, the less thoroughly it is
digested). In contrast, there is a posi-
tive relationship between particulate
passage rate and fluid dilution rate
(i.e., as passage rate increases, fluid
dilution rate increases). Forage intake
by ruminants has been reported to be
affected by the proportion of indigest-
ible fiber and rate of passage from
the reticulo-rumen and resistance of
large particles to breakdown to small
particles within the reticulo-rumen
(Hungate, 1966). Rate of passage of
digesta through the digestive tract in-
creases with feed intake, which allows
the animal to increase its energy in-
take, even though ruminal and total-
tract digestibility might be decreased
(Firkins et al., 1998). Retention of
undigested fiber allows ruminants to
increase ruminal fiber digestion, but
extended ruminal retention times of
the retained fiber can decrease feed
intake because of ruminal distention
(i.e., filling effect; Allen, 1996).
Okine et al. (1998) reviewed the
literature and concluded that in
addition to physical and chemical
properties of the feed, physiological
and metabolic state of the animal
influence digesta passage and that
 Ruminal fermentation and digestive physiology
ruminants have some control over
digesta passage from the reticulo-
rumen. Regulation of the relaxation of
the reticulo-omasal orifice coordinated
with muscular contractions of the
reticulo-rumen might be important
for controlling digesta passage from
the reticulo-rumen, and thus rumen
fill and voluntary intake (Okine and
Mathison, 1991).
A theory of control of digesta pas-
sage through the reticulo-omasal
orifice was outlined by Deswysen
(1987). Briefly, immediately after the
second phase of reticular contraction,
synchronized contractions of the oma-
sal canal and omasal body (between
the leaves) occur. Contractions of the
omasal body are very slow and de-
liberate. Reticular contractions come
in 2 phases. The first phase serves to
distribute forage in the rumen and
push the hay mat into the rumen.
The lips of the ventricular groove
close the reticular-omasal orifice to
prevent large particles from enter-
ing the omasum. During the second
phase, fine feed particles are pushed
up and enter the omasum. Several
factors contribute to decreased escape
of digesta from the rumen. Particles
sequestered within the hay mat have
a decreased probability of leaving the
rumen. As particles are decreased
in size through digestion and rumi-
nation, fermentation gases are re-
leased, functional-specific gravity is
increased, and particles sink to the
ventral rumen. This allows particles
to be moved by ruminal contractions
toward the reticulo-ruminal orifice
where they are more likely to pass
from the rumen (Sutherland, 1987;
Faichney, 1993).
Microbial Factors
Several factors influence the relative
proportions of microbial species with-
in the rumen. For example, Jouany et
al. (1988) suggested that recycling of
microbial matter via protozoa pre-
dation can significantly affect both
energy and protein utilization in the
rumen and subsequent nutrient supply
to the animal. The factors involved
have been reviewed (Jouany et al.,
1988). Engulfed microbial protein can
be incorporated into protozoal cells,
fermented to VFA and ammonia,
or released into ruminal fluid. The
amount of engulfed bacterial pro-
tein incorporated into protozoal cells
depends on the availability of energy.
Other factors influencing growth of
microorganisms in the rumen include
substrate preference (Russell and
Baldwin, 1978), maximum growth
rates (Russell and Baldwin, 1979a),
growth factors (branched-chain VFA),
substrate affinities (Russell and
Baldwin, 1979b), maintenance energy
requirements of the microbes, and
uncoupled fermentation.
Ruminal pH is one of the most vari-
able factors, and it can influence the
microbial population and the levels
of VFA produced. The ruminal pH at
which certain functions are optimized
can differ. Fibrolytic microorgan-
isms are most active at a pH of 6.2 to
6.8. Cellulolytic bacteria and metha-
nogenic bacteria can be decreased
when the pH begins to fall below 6.0.
Amylolytic microorganisms prefer a
more acidic environment, a pH of 5.2
to 6.0. Certain species of protozoa can
be greatly decreased when pH falls
below 5.5. To accommodate all these
needs, normal feeding practices should
maintain a pH range between 5.6 and
6.5.
The diet fed to ruminants influences
the number and relative proportions
of the different microbial species
in the rumen. The final composi-
tion of the ruminal microflora is a
direct reflection of the physiology
of the microorganisms present. No
single organism is responsible for the
complete degradation of complex feed
substrates in the rumen. Digestion
of feed is accomplished by a myriad
of complementary organisms that
produces an array of enzymes needed.
Complex feedstuffs are colonized and
broken into oligomers and eventually
tri-, di-, or monomers of each compo-
nent, which are in turn used by other
members of the microbial population
to produce VFA, B vitamins, or cofac-
tors needed by other microorganisms
135
(Krause et al., 2013). This symbiotic
relationship prevents the accumula-
tion of digestive end products and
accelerates the digestive process. If a
microorganism is to be stably main-
tained in the rumen, the growth rate
of that microorganism must exceed
0.69 (i.e., ln = 2) of the passage rate
of rumen digesta. One of the most
common problems encountered by
nutritionists involves sudden changes
in the diet of the ruminant to include
large amounts of readily fermentable
carbohydrates. Feeding diets of this
type results in a succession of changes
in the rumen microbial population
during the adaptation period, specifi-
cally in those bacteria that produce
and use lactate. During this process,
acid-sensitive lactate users can be re-
placed by acid-tolerant lactate users.
Lactic acidosis can result from too
abrupt of a shift to a high-concentrate
diet (see below).
DIGESTION KINETICS
The digestive process in ruminants
is complex, with the animal, dietary,
and management factors discussed
interacting to affect rate and extent of
feed digestion in the rumen and total
digestive tract. Nutrient supply to
the animal is influenced by nutrient
digestibility, and therefore the ability
to measure site and extent of diges-
tion is important. Feed components
vary in the rate and extent they are
digested in the rumen and digestive
tract. For example, sugar and soluble
cell-wall carbohydrates (e.g., betaglu-
cans, pectins) are almost completely
degraded in the rumen (Van Soest,
1994), whereas digestion of starch in
the rumen is variable depending on
feed source and processing method.
Nonetheless, total-tract digestion of
starch is typically greater than 80%
as a result of extensive postruminal
digestion (Firkins et al., 2001). In
contrast to starch, digestion of fiber
in the rumen depends on the physical
and chemical attributes of the fiber
consumed and the microbial popu-
lation, with fiber escaping ruminal
digestion only partially degraded in
136
the hind-gut (Huhtanen et al., 2006).
Typically less than 10% of NDF di-
gestion occurs in the hindgut of cattle
(Huhtanen et al., 2006).
The balance of rate of microbial di-
gestion of feed and the rate at which
feed passes from the rumen deter-
mines the digestion of fiber in rumi-
nants. For example, a faster passage
rate or a slower digestion rate will
decrease ruminal digestion, whereas a
slower passage rate or a faster diges-
tion rate will increase ruminal diges-
tion. Most diet components in feeds
that are not part of the cell wall are
rapidly digested, with rates that are
3 to 10 times faster than rates of pas-
sage (Mertens, 1993). In contrast, rate
of plant-cell-wall digestion is generally
similar to rate of passage, although
rate of plant-cell-wall digestion is
highly variable depending on plant
species (e.g., grasses vs. legumes)
and maturity of the forage (Mertens,
1993). Retention time in the rumen
increases extent of fiber digestion but
lowers DMI because of rumen fill.
Total-tract DM digestibility ranges
from 40 to 50% for low-quality forages
(Waiss et al., 1972; Horton, 1978)
to 75 to 90% for high-starch grain
diets fed to feedlot cattle (Zinn, 1990;
Beauchemin et al., 2001; Corona et
al., 2006).
RUMINAL FERMENTATION
Volatile fatty acids and microbial
protein are produced as a result of ru-
minal digestion. The VFA provide ap-
proximately 80% of the metabolizable
energy needs of the animal. Propor-
tions of ruminal VFA usually range
from 50 to 70% of the total VFA for
acetate, 15 to 35% for propionate,
and 10 to 12% for butyrate. Butyrate
is largely converted to ketones during
absorption through the rumen epithe-
lium, and β-hydroxy-butyrate ac-
counts for more than 80% of ketones.
The proportion of VFA is greatly
influenced by diet (forage:concentrate
ratio) and the status of the methano-
genic microbial population in the ru-
men. Although the main end products
of fermentation, VFA, branched-chain
VFA, formate, ethanol, lactate, and
Krehbiel
succinate, are also formed. In addi-
tion, ammonia, CO2, and H2 gas are
produced. Long-chain fatty acids are
released from lipids, and many other
end products result from microbial
degradation of minor components of
the feed. Hydrogen gas produced dur-
ing microbial fermentation of feed is
used by Archaea (i.e., methanogens)
in a syntrophic process known as
interspecies H2 transfer. The methano-
gens reduce CO2 with H2 to produce
CH4, which is removed from the ru-
men by eructation and respiration.
Volatile fatty acids are passively
absorbed through the ruminal wall.
This continuous removal of VFA by
absorption from the reticulo-rumen
is important for maintaining a stable
ruminal pH. Removal of acid products
is also important for the continued
growth of cellulolytic organisms. Vola-
tile fatty acids in digesta can flow to
the omasum and are absorbed. Rate
of VFA absorption from the rumen
is influenced by the chain length of
individual acids (butyrate > propio-
nate > acetate) and by ruminal pH.
Lower pH and the resultant increases
in the proportion of the acids in the
rumen favor more rapid absorption.
Amino acids, oligopeptides, and am-
monia released by microbial degrada-
tion of proteins and NPN compounds
are used by microorganisms and
converted to microbial cell protein.
The microbial protein and ingested
plant proteins supply the animal with
dietary amino acids. The microbes,
undigested feed particles, and com-
pounds dissolved in the rumen fluid
can be used by the animal when they
exit the rumen and enter the aboma-
sum and lower digestive tract.
Lactic acid does not normally ac-
cumulate in the rumen. If gradual
introduction of grains is practiced,
the lactate-using bacteria will develop
and permit only a transient increase
in lactic-acid accumulation follow-
ing ingestion of a diet high in readily
fermentable carbohydrates. Problems
arise when large amounts of starch
or cereal concentrates are fed. Total
lactate in acute acidosis can compose
50 to 90% of the rumen acids. The
absorption of large amounts of lactic
acid across the rumen wall to the
blood produces systemic acidosis and
associated metabolic problems (see
below).
RUMEN DYSFUNCTION
Ruminal Acidosis
Owens et al. (1998) reviewed the
literature pertaining to the effect of
the overconsumption of high-grain
diets by ruminants. Acute acidosis
is an overt illness caused by cattle
consuming an excessive quantity of
readily fermentable carbohydrate
(e.g., starch), which increases total
acid concentration (including lactic
acid) in the rumen. This results in
ruminal pH falling below 5.2. Absorp-
tion of acids into portal blood leads
to systemic acidosis, stiffness of legs,
laminitis and founder, severe rumi-
nal lesions, and sometimes death.
Other signs of acute acidosis include
abdominal kicking, decreased rumen
motility, anorexia, shallow breathing,
increased heart rate, profuse diar-
rhea, gurgling gas in the rumen, and
visual observation that cattle overate
high-concentrate feed. In addition
to affecting the animal, low rumi-
nal pH has been shown to alter the
rumen microbiome (Nagaraja and
Titgemeyer, 2007) in a manner that
decreases the rate and extent of fiber
digestion. Furthermore, low pH (<5.6)
can damage the ruminal epithelia
leading to decreased barrier function
(Penner and Beauchemin, 2010) and
associated animal health issues (e.g.,
lameness, rumenitis, liver abscesses,
acute-phase inflammatory response).
Subacute acidosis is more difficult to
detect, but it is usually associated
with decreased or erratic feed intake
and lower animal performance. In
subacute acidosis, increased acid load
results in a ruminal pH of approxi-
mately 5.6.
The increase in VFA and lactate
production and decreased ruminal pH
associated with acidosis imposes a
challenge to the metabolism and the
regulation of intracellular pH homeo-
stasis of ruminal epithelia (Penner
et al., 2011). The ruminal epithelia
 Ruminal fermentation and digestive physiology
respond in part by enlarging the
absorptive surface area. In addition,
recent evidence suggests that changes
in epithelial cell function may be the
initial response. Gene expression anal-
ysis indicates that pathways involved
in fatty-acid metabolism, ion trans-
port, and intracellular homeostasis are
affected during adaptation and after
adaptation to a highly fermentable
diet (Penner et al., 2011). This is im-
portant because metabolism of VFA,
particularly butyrate, by ruminal
epithelium helps maintain a concen-
tration gradient across the rumen wall
and into portal blood circulation.
Ruminal Bloat
Eructation is a normal and neces-
sary process for gas removal from
the rumen. As discussed earlier, gas
production in the rumen is a by-prod-
uct of ruminal fermentation. Bloat is
defined as the excess build-up of gas
that if unresolved ultimately results in
asphyxiation of the animal, resulting
in death. In specific conditions, the
fluid layer of the rumen may develop
frothy foam (feedlot bloat). Eructa-
tion of gas through the esophagus is
inhibited when the cardia is covered
with foam (Cheng et al., 1998). Foam
has an increased surface tension and
is able to trap gases, thereby prevent-
ing their release. Similar to acidosis,
frothy bloat often occurs during adap-
tation to a high-grain diet; however,
sporadic outbreaks are often reported
in cattle that are well adapted to
high-grain diets.
IMPLICATIONS
The unique features of the complex
stomach of the ruminant animal allow
for use of food resources not other-
wise useful to humans. In addition,
the ruminant animal is adaptable to
feedstuffs other than fibrous mate-
rial, which when fed to ruminants
can increase efficiency of feed use.
Many factors involving diet, intake,
ruminal fermentation, and removal
of end products are associated with
supplying nutrients to the host ani-
mal. Increasing our understanding
of the interactions that exist among
the diet, rumen and gastrointestinal
microbiome, and animal physiology
is essential for improving efficiency of
ruminant production systems. Under-
standing adaptation of ruminal epi-
thelia and gastrointestinal mucosa at
the molecular level could also increase
our ability to manage cattle to opti-
mize ruminal pH, increase nutrient
transport, and mitigate adverse meta-
bolic disorders. Recent advances and
continued study in the area of ruminal
fermentation and digestive physiology
have the potential to positively affect
animal production and efficiency, ani-
mal health, and the environment.
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