PHYSIOLOGY AND MICROBIOLOGY OF RUMEN

A. What is the process of rumination?

There are so many different species of ruminant animals. We have cattle, sheep, goats, buffalo,
deer, elk, giraffes and camels. These animals a a unique digestive systemfrom others and even from our
own. Instead of having once compartment in the stomach, these animals have four. And these four
compartments includes rumen, reticulum, omasum and abomasum. Rumen is the largest section among
them, this is the maindigestive centre. The rumen is filled with tiny microorganisms that can break
downgrassand other coarse vegetation that animals that only have one stomach can't digest.
Therumminant animals do not completely chew grass. The partially chewed grass goes intothe rumen
where it is stored and be broken down, these partially chewed grass will nowbe known as "cud." Then
the cud is pushed through the reticulumand then returnedtomouth, this process is called "rumination."
When the re-chewed food is swallowedagainit will now be sent to omasum. In the omasum, the cud is
broken down into evenmoresmaller pieces and then it will be sent in the abomasum which is the last
chamber of thestomach. Then the digested food goes to small intestine where absorption of
nutrientstakes place.
Dairy calves have a four-part stomach when they are born. However, theyfunctionprimarily as a
monogastric (simple-stomached) animal during the first part oftheir [Link] birth the first three
compartments of a calf’s stomach—rumen, reticulum, andomasum—are inactive and undeveloped. As
the calf grows and begins to eat a varietyoffeeds, its stomach compartments also begin to grow and
change. Theabomasumconstitutes nearly 60 percent of the young calf’s stomach, decreasing toabout
8percent in the mature cow. The rumen comprises about 25 percent of the youngcalf’sstomach,
increasing to 80 percent in the mature cow.

I. How are ruminants differ from monogastric animals?

The biggest difference between ruminant and monogastric animalsis
their stomach compartments. Ruminants have four compartments or chambers: the rumen,
reticulum, omasum, and abomasum. These chambers containmicrobes that deteriorate
cellulose. While monogastric animals have a simplesingle-chambered stomach, they only have
one compartment. Most monogastric
animals are unable to digest much cellulose-based food like grass. Theabomasum, which is
also called the true stomach, is the equivalent of amonogastric stomach chamber.
II. How are large ruminants differ from small ruminants?
Obviously, from the word itself, small and large ruminants differ inbody
build and body size. So, upon looking at this, we can say that they differ fromtheway of living
and the way of handling them. They differ, especially fromthepoint of view of the person who
nurtures these ruminants. Small ruminants areobviously less expensive to care for than large
ruminants because theyfit intosmaller holdings and eat fewer grains than large ruminants. They
alsoserveatriple purpose: they can be used for milk, meat, and [Link], some small ruminants,
like sheep and goats, have the ability to adapt to survive and producein difficult countries. In
 those countries with dry places that have a highaltitudeor maybe in countries where it is
extremely cold. So, even if other largeruminants can stand these countries or can give the three
purposes that small ruminants can, we can change the fact that taking care of small ruminantsis
cheaper than having large ruminants. Also, small ruminants have a muchsuperior capacity to
recycle urea than large ruminants. As for this reason, small ruminants digest more efficiently.

B. Rumen microorganisms and their role in Ruminal fermentation

I. Bacteria

Rumen microbes are the major source of protein n cow’s diet. They break
down Rumen Degradable Protein or RDP to amino acids, then ammonia. Ammoniais a major
sourceof nitrogen for microbial growth. The microbes also convert non-protein nitrogen to
ammonia. There are 25 billion bacteria inside the rumen, it includes protozoa, bacteria and
rumen. The rumen is a fermentation chamber inwhich bacteria digest cellulose in grasses and
feeds then convert themto fattyacidsand amino acids which are fundamental nutrients used by
the cowand basis for thecow’s production of milk.
Rumen bacteria can be divided into four essential sub populations according to their placeinto the
rumen:
1) those attached to the rumen epithelial cells which are less than 1%of the total rumen microbes,
2) those attached to feed particles (~ 70-80% of the total rumen bacteria) 3) those
suspended in the ruminal fluid (~ 20-25% of total rumen bacteria) 4) those attached to the
surface of protozoa or fungal sporangia.
Rumen bacteria are also classified according to their function, and these are:
● Cellulolytics- these bacteria are essential to the rumen environment. It secretes free enzymes depending
on the hydrolysis of lignocellulose into usable sugar enzymes with
specific substrate specificities. In order to perform cellulolytic activity, rumen cellulolytic bacteria
must adhere to cellulose. These species have a wide range of fibrolytic enzymes called glycoside
hydrolases (GHs) for cellulose degradation.
● Hemicellulolytics- these are non-cellulose fibers that degrade the rumen bacteria. They also have over a
hundred glycoside hydrolases. And the majority of cellulolytics can also break down hemicellulose.
● Pectinolytics- these are widely distributed among fungi and bacteria, they can be found in higher plants,
parasitic plants and some plant parasitic nematodes (Shrivastavaet al. 1994; Jayani et al. 2005). These
include the Treponema saccharophilumandLachnospira multiparus. These two bacteria break down the
pectin. Butyrivibrio
fibrisolvens, Bacteriodes ruminicola, Lachnospira multiparus, Succinivibriodextrinosolvens and
 Streptecoccus bovis. The three groups of bacteria mentioned above make up the majority of the
rumen’s bacterial population. These three species together with the amylolytics are called primary
rumen bacteria and the other bacterial groupsare called seconders due to using the end products
of the primary ones.
● Amylolytics- a microorganism can be considered as amylolytic when they can grow in the presence of
starch as the primary carbon source and form a degradation zone around the colony. Amylolytics grow
faster when ruminants are fed a higher proportionof grainfeed, fast-growing species with doubling
times ranging from 15 minutes to 4hourswhen compared to cellulolytics. As end products, these
species generate propionicacid, lactic acid, succinic acid, format, and CO2. They are involved in the
breakdownofstarch and soluble sugar. The Streptococcus bovis, Ruminobacter amylophilus, Prevotella
ruminicola, and Butyrivibrio fibrisolvens are mainly amylolytic bacteria. While, Succinivibrio
dextrinosolvens, Succnivibrio amylolytica, Selenomonasruminantium, Bifidobacterium ruminantium,
Lactobacillus acidophilus, Lactobacilluscasei and Lactobacillu fermentum are saccarolytic.
● Acetogens- these bacteria is described as the small proportion of rumen bacteria, because, theis
reductive acetogenesis capability from CO2 to H2. In the rumen, theseareacetitomaculum ruminis and
Eubacterium limosum. Although their reductiveacetogenesis capability can be proved in vitro,
acetogens have not functionedasalternative H2 sink in the rumen
● Proteolytics- these are the bacteria that can produce protease enzymes, the enzymes that can break down
peptide bonds in protein molecules. Most of these bacteria are foundinsoil, water, mud, and certain
strains of environmental conditions. The most commonproteolytics are Prevotella ruminicola,
Ruminobacter amylophilus, Clostridiumbifermentans and Colostridium proteoclasticum. Proteolytic
activity represents a majorloss of dietary amino acids for ruminant.
● Acid and urea utilizers- the urea that are transferred across the rumen wall fromtheblood and then it is
hydrolyzed to ammonia by resident a bacteria. Megasphaeraelsdeni and Anaerovibrio lipolytica have
a critical role when the rumen lactic acid productionishigh due to the high grain content feed.
Megasphaera elsdeni also utilizes ureatoammonia (NH3) and CO2.
● Lipolytic- these bacteria cause FFA to be released from esterified plant lipids andthenfollowed by
biohydrogenation, which reduces the number of double bonds. Glycerol isconverted to propionic acid
and succinic acid by Anaerovibrio lipolytica, while fructose, ribose, and lactic acid are converted to
acetic acid, propionic acid, and CO2. The critical importance of Anaerovibrio lipolytica came from its
ability to utilize lactic acidinruminants fed high grain feed. Every fermentation process generates a
small amount ofH2.

II. Methanogens
Methanogenesis is carried out by methanogenic archaea, it is a specialized groupofmicrobes found in a
variety of anaerobic environments such as the rumen. Methanogensinthe rumen primarily use H2 and
CO2 as substrates to produce methane, filling an important functional place in the ecosystem.
In a recent studies, they revealed that rumen methanogens constitute 2.8 to4%ofruminal
microorganisms and it belongs to the domain archaea. More than 90%of rumenarchaea are member of
genera; Methanobrevibacter (more than 60%), Methanomicrobium(up to 15%), and rest of the rumen
archaea referredtorumen cluster C approximately 16% or Thermoplasmatalesthat function in the
rumenisunknown. Although they are a small number of rumen microorganisms, the effectsof rumen
methanogen archaea on rumen fermentation are significant as theyarethe main CH4producers in the rumen.
Interestingly, a considerable proportionofmethane production in the rumen is attributed mainly not to the
MethanobrevibacterorMethanomicrobium, which constitutes roughly 75% of the rumen archaea. Instead, it
isattributedto the rumen cluster C archaea even though they are only 16%of rumenarchaea, and their
fundamental function and biochemical flow in the rumen has remainedunknown yet

III. Protozoa
There are two types of rumen protozoa. Entodinomorphs and holotrichsThese protozoa are unique
in terms of phenotypic and behavioral adaptations that allow them to survive in this harsh
anaerobic environment. Protozoa engulf and digest a wide range of bacteria (Williams and
Coleman, 1992) and can reduce the shedding of potential pathogens from the animal, although the
effect is highlydependent on the composition of the protozoal population present (Stanfordet al.,
2010). In addition to the bacteria, the obligate or facultative anaerobic protozoawith various
species of ciliates and flagellates comprise another group of rumenmicroorganisms. Ciliata makes
up nearly half of rumen biomass, while flagellatesmake up much less. Some researchers believe
protozoa are important for digestivefunctions in the rumen. Other researchers, on the other hand,
reported themasvaluable as a nitrogen source to compensate for nitrogen-poor feed out of season
only for wild ruminants fed primarily with poor quality forage. Protozoa consumenutrients,
bacteria, fungi, and other protozoa. They play an important roleinpreventing rumen acidosis by
rapidly removing carbohydrates that are easilyfermented. Furthermore, they remove toxic
compounds from plants and reducetherisk of heavy metal poisoning. Protozoa, on the other hand,
are not requiredfornormal digestion.

IV. Viruses
Bacteriophages are viruses that infect rumen bacteria. In fact, the word"bacteriophage"
literally means "bacteria eater," because bacteriophages destroytheir host cells. All bacteriophages
are composed of a nucleic acid molecule that issurrounded by a protein structure. Their number
range from 2x107 to 1x108per ml of rumen fluid. Although more than 100 different
bacteriophages have beenidentified to date, this number can be expected to expand to a large
number ofbacteriophage species, given that bacteriophages are unique to all bacterial species.
Biotechnological methods can develop bacteriophages specific to the typeofbacteria, and there
may be a potential for suppression of bacteria whose rateofreproduction in the rumen is to be
decreased
 V. Parasites
The first study that empirically demonstrates disease-driven increasesinmethane (CH4) yield in
livestock reveals interesting results. Gastrointestinal (GI)parasitism is a disease caused by
different genera of parasites that inhabit thedigestive tract of cattle and sheep, causing
inappetence, anemia, diarrhea, poorgrowth, and economic losses in the herds. Basically, GI
parasitismin cattleand
sheep is caused by helminths and protozoa. Gastrointestinal parasite infestationsincrease
methane yield (g CH4/kg of DMI) by up to 33%

C. Manipulation of ruminal fermentation

A significant amount of greenhouse gases around the world is released due to agricultural
activities that play a critical role in food production and the economy. However, global warming and loss
of productivity due to greenhouse gas emissions arising fromtheseactivities have still been debated. While
animal products account for 40%of the world’sagricultural products, a significant proportion of this
production comes fromruminants. Ruminants produce meat and milk, which are precious nutrients for
humans, by digestingtheplant’s structural components that humans cannot digest. Humans provide life
safetyandhealthy feed resources for the ruminants the whole year and ruminants provide essential
nutrients for humans. This mutualistic cooperation with ruminants raised the world’sruminant population
enormously while humans supply essential nutrients such as milkandmeat. However, due to fermentation,
ruminants release greenhouse gases such as methane(CH4) and nitrous oxide (N2O) which are essential
contributors to global warming. Inaddition, the formation of nitrous oxide is significant during the
processing of agricultural land used in ruminant nutrition. Since the 2000s, scientists have performed
numerous studiesto develop new methods for reducing methane gas emissions in livestock. In those
studies, scientists have proposed various methods such as the addition of ionophores, organicacidsand
plant essential oils to feed, immunization, modification of feed composition, rumendefaunation,
alternative hydrogen (H2) sinks, modification of the microbial rumendistribution, and animal breed
 replacement with the minimal methane-producing breeds

D. Modification of the feed composition

The composition of the feed given to ruminants significantly affects the productionof methane.
Methane production can be reduced by up to 90% with the modificationof thefeed composition.
Fermentation performance decreases with poor-quality roughage, lackingin vitamins, minerals, proteins,
and energy, thus increases methane production. However, supplementation of minerals and nitrogen
sources improves fermentation performanceandconsequently decreases methane production. Fresh
roughage reduces methane production. Fresh alfalfa, oats, sorghum instead of hay and replacing the
30%wheat strawwithfreshsorghum reduces methane production by 33%. Feeding with roughage rich in
tannins, lowfiber, a high rate of dry matter, and a shorter digestion time in rumen reduces methane
production. Feeding with fast degradable roughage, slowly digestible starch instead of fasterone, legumes
instead of meadow, silage instead of fresh or dry grass, and even optingforcorn silage with relatively slow
degradable starch instead of meadow silage reduces methaneproduction by up to 28%. Fermentation of
starch promotes the production of propionicacidcompared to feeds with low starch content. Propionic acid
production decreases methaneproduction by allowing the greater use of metabolic H2 and suppressing the
protozoathat
are important H2 suppliers for methanogens, by reducing rumen pH. Sugar digestion, ontheother hand,
leads to more methane production than starch. Since sugar can be dissolvedinwater, it is quickly
fermented in the rumen and is mainly used to produce butyricacid. Butyric acid increases methane
production when rumen pH is high and adequate metabolicH2 is present. Methane production can be
reduced by up to 90% when the concentratefeedsrate increases to 90%. On the other hand, in this case,
the risk of subacute ruminal acidosis(SARA) should be considered. Subacute ruminal acidosis (SARA)
is a metabolic diseaseinhigh-producing dairy cattle. This disease is caused by feeding high concentrate
diets andisdefined as a depression of ruminal pH below 5.6 at least 3 h/day. Feed with a ratioof
90%concentrates is not sustainable in ruminants.

E. Effects of supplementing organic acid to the feed

Organic acids are not recommended for livestock since they are expensiveanddifficult to apply in grazing
ruminants. However, researchers reported that organic acids(fumarate and malate) reduce the synthesis of
ruminal methane. Organic acids increase the synthesis of propionic acid by using H2 as an alternative
biochemical pathway instead of methane synthesis. Thus, methane synthesis is reduced due to the lack of
H2 in the rumen. It has been reported that methane gas production decreases to a negligible level of
1-2%with an 80-90% concentrate proportion of feed. However, in this case, the risk of subacuteruminal
acidosis (SARA) arises. Further reduction in rumen pH may be preventedbyanalternative H2 sink created
via organic acid supplementation. In this case, feeding withhighconcentrates, which radically reduces the
methane synthesis, may become sustainable
F. Effects of supplementing antibiotics to the feed
Supplementing the feed with the ionophore group of antibiotics to increasetheruminant's yield also
significantly decreased the greenhouse gas synthesis unto the 2000s. Ionophore antibiotics are fermentation
products of different Streptomyces species andotherfungi. They are polyether antibiotics, which exert their
antibiotic action by disruptingthetransport of ions in the cell membranes. Ionophores reduce the H2
synthesis by upto30%via suppressing H2 producers such as Gram-positive bacteria and ciliates rather
thanmethanogen archaea. Although ionophores increase ruminant’s yield, they are not valuablefor reducing
methane production in advanced enterprises where ruminants feedwithconcentrates with high protein and
energy to meet the high nutrition needs. In addition, theinhibition effect of the ionophore antibiotics on
methane production is not constant. Although ionophores generally significantly reduce greenhouse gas
emissions that arisefrom enteric fermentation, they have been banned in European Union Countries
andTurkeysince 2006 due to various concerns such as developing resistant microorganisms
andfoodresidues. Although it is not prohibited in other countries, the livestock and agriculture sectoris
forced to take alternative and even more effective measures against ionophores due tothereaction
worldwide against the antibiotic additives
G. Effects of probiotic addition to feed
Oeztuerk reported a decrease in acetic acid/propionic acid (A/P) rate and improvedfermentation
performance in an in-vitro study 0.7% alive Saccharomyces cerevisiaesupplemented to the ruminant feed.
Saccharomyces cerevisiae is a species of yeast (single-celled fungus microorganisms). It is reported that
the addition of yeast to the diet reducesmethane production by increasing the synthesis of propionic acid,
reducing the number ofprotozoa, and increasing animal yield reported that yeast supplementation
accelerates thesynthesis of acetic acid by acetogens and consequently suppresses methane synthesisthrough
the consumption of metabolic H2 in the rumen. Adding yeast to the feedalsocontributes to the stabilization
of the rumen pH, thus ruminants fed with concentratesbecome more resistant to subacute rumen acidosis
(SARA).
Probiotics are typically used to improve gastrointestinal health, regulate, andbalancegut microbes,
promote growth and development of animals, and improve the host resistanceto diseases. In ruminants,
probiotics are administered to target the rumen (main site of feeddigestion) where they influence rumen
fermentation especially on feed digestibilityanddegradability and rumen microbiota. Probiotic positively
affect cellulolysis and synthesis of microbial protein during digestion and stabilizes rumen pH and lactate
levels. In addition, probiotics can enhance nutrient absorption. Direct-fed probiotic have been shown
toreduceruminal acidosis.

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