Digestion in ruminants

•Anaerobic

 pH between 6 and 7
 Each ml of rumen content contains…
 10 to 50 billion bacteria
 1 million protozoa
 Various numbers of yeast
and fungi

Image source: Lock Haven

University
Rumen Microbes
• What makes ruminant digestion of cellulose possible is the diverse
population of microbes that inhabit this structure.
• These microbes interact and support one another, passing along
partially digested forage.
• The waste of some microbes becomes the food other others
• These microbes are what actually
digest the forage
• They produce the cellulase enzyme
that breaks down the cellulose in
the plant cell walls.
Microbial Services
• Rumen microbes provide 4 key services to a cow:
• 1. Amino acid production: all animals must consume the amino acids their own bodies can’t
make in order to build proteins. Rumen microbes make these essential amino acids and
supply them to the cow’s body.
• 2. Protein production: some proteins cannot be made from plant sources. Microbes can
utilize sources of proteins that cows cannot (such as the urea created from protein
digestion) to produce more protein for the cow’s body.
• 3. Synthesis of B-vitamins: without
microbes, cattle would be deficient
in all but two of the B vitamins.
• 4. Break-down of cellulose – rumen
microbes produce the cellulase
enzyme needed to break down
cellulose into digestible glucose.
Microbial Fermentation
• Almost all feed ingested by the cow is actually used to feed the microbes in its
rumen.
• The cow itself gets the waste byproducts from the microbes after they ferment the forage.
• Fermentation occurs under anaerobic (w/o oxygen) conditions
• Because of this, the sugars freed when cellulose is broken apart become fermented into
Volatile Fatty Acids, or VFAs.
• If microbes were exposed to oxygen, they would completely break down the forage into
CO2
• VFA’s provide the majority of the energy needs of an herbivore.
• Carbohydrate Digestion
• Forages
• On high-forage diets ruminants often ruminate or regurgitate
ingested forage.
• This allows them to “chew their cud” to reduce particle size
and improve digestibility.
• As ruminants are transitioned to higher concentrate (grain-
based) diets, they ruminate less.
• Once inside the reticulorumen, forage is exposed to a unique
population of microbes that begin to ferment and digest the
plant cell wall components and break these components
down into carbohydrates and sugars.
• Rumen microbes use carbohydrates along with
ammonia and amino acids to grow.
• The microbes ferment sugars to produce VFAs
(acetate, propionate, butyrate), methane, hydrogen
sulfide, and carbon dioxide.
• The VFAs are then absorbed across the rumen wall,
where they go to the liver.
• The three major VFA absorbed from the rumen have somewhat
distinctive metabolic fates:
• Acetic acid is utilized minimally in the liver, and is oxidized throughout
most of the body to generate ATP. Another important use of acetate is
as the major source of acetyl CoA for synthesis of lipids.
• Proprionic acid is almost completely removed from portal blood by the
liver. Within the liver, proprionate serves as a major substrate for
gluconeogenesis, which is absolutely critical to the ruminant because
almost no glucose reaches the small intestine for absorption.
• Butyric acid, most of which comes out of the rumen as the ketone
beta-hydroxybutyric acid, is oxidized in many tissues for energy
production.
• Once at the liver, the VFAs are converted to glucose
via gluconeogenesis.
• Because plant cell walls are slow to digest, this acid
production is very slow.
• Coupled with routine rumination (chewing and
rechewing of the cud) that increases salivary flow,
this makes for a rather stable pH environment
(around 6).
• High-Concentrate Feedstuffs (Grains)
• When ruminants are fed high-grain or concentrate rations, the
digestion process is similar to forage digestion, 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 concentration 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 greatest 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 population by
decreasing the forage using microbial population and
potentially leading to a decrease in digestibility 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 ruminant’s ability to
buffer and absorb these acids and lead to metabolic
acidosis.
• The acidic environment leads to tissue damage within
the rumen and can lead to ulcerations of the rumen wall.
• Take care to provide adequate forage and avoid
situations that might lead to acidosis when feeding
ruminants high-concentrate diets.
• Protein Digestion
Two sources of protein are available for the ruminant to use:
protein from feed and microbial protein from the microbes that
inhabit its rumen.
A ruminant is unique in that it has a symbiotic relationship with
these microbes.
Like other living creatures, these microbes have requirements
for protein and energy to facilitate growth and reproduction.
• During digestive contractions, some of these
microorganisms are “washed” out of the rumen into the
abomasum where they are digested like other proteins,
thereby creating a source of protein for the animal.
• All crude protein (CP) the animal ingests is divided into
two fractions, degradable intake protein (DIP) and
undegradable intake protein (UIP, also called “rumen
bypass protein”).
• Each feedstuff (such as cottonseed meal, soybean hulls,
and annual ryegrass forage) has different proportions of
each protein type.
• Rumen microbes break down the DIP into ammonia
(NH3) amino acids, and peptides, which are used by the
microbes along with energy from carbohydrate digestion
for growth and reproduction.

• Excess ammonia is absorbed via the rumen wall and

converted into urea in the liver, where it returns in the
blood to the saliva or is excreted by the body.
• Urea toxicity comes from overfeeding urea to ruminants.
• Ingested urea is immediately degraded to ammonia in
the rumen.
• When more ammonia than energy is available for
building protein from the nitrogen supplied by urea,
the excess ammonia is absorbed through the rumen
wall.
• Toxicity occurs when the excess ammonia
overwhelms the liver’s ability to detoxify it into urea.
• This can kill the animal.
• However, with sufficient energy, microbes use
ammonia and amino acids to grow and reproduce.
• The rumen does not degrade the UIP
component of feedstuffs.
• The UIP “bypasses” the rumen and makes
its way from the omasum to the abomasum.
• In the abomasum, the ruminant uses UIP
along with microorganisms washed out of
the rumen as a protein source.
• Digestion of Fat
• Fats are a source of energy for cow.
• Fats are either partially degraded in the rumen or
assume a bypass or protected form.
• When microbial fermentation of fat occurs in the rumen,
some vitamins required by cow are also produced.
• Fats are present in most of the more common dairy
feeds in relatively small amount.
• No more than 5% of the total diet dry matter ( or about
500gm/day) should consists of fat.
• Beyond this level fat will coat the dietary fibre in the digestive
tract, interfering with fibre digestion and decreasing the
palatability of the diet.
• Protected fat which escape microbial digestion in the rumen, can
be used to overcome digestive upsets caused by high levels of
rumen degradable fat.
• The protected fats are readily digested and absorbed across the
wall of the small intestine.
• Interest in feeding protected fats to lactating dairy cows is
growing.
• However, they are very expensive and only relevant to producing
herds (30 L/day or more), and therefore are not practically
relevant small holder farmers.
• Absorption of carbohydrate
• Carbohydrates are broken down into smaller
polysaccharide in the mouth by the action of salivary
amylase.
• Once in the small intestine, pancreatic amylase further
breaks the polysaccharide into disaccharides.
• The three most common disaccharide are maltose,
sucrose and lactose.
• These disaccharide are broken down into
monosaccharide by the digestive enzymes found at the
brush boarder of the enterocyte.
• Maltose is obtained via the digestion of starch by
amylase.
• It consists of two glucose monomers attached by an
alpha- 1,4 glycosidic bond which is broken down by
brush border enzyme called maltase.
• Sucrose consists of glucose and fructose and is broken
down by sucrase while lactose consist of glucose and
galactose and is broken down by lactase .
 mouth lumen of small intestine
Disaccharides ( Maltose,
Carbohydrate 1 Polysaccharide 2
Sucrose, Lactose)

3 Brush boarder

Glucose Fructose galactose

• The cells of small intestine called enterocytes can only
absorb sugars in their monomeric from.
• Therefore they absorb glucose, galactose and fructose.
• Glucose and galactose enter enterocyte via sodium-
linked secondary active transport.
• ATP is used to establish and electrochemical gradient in
which there is a higher concentration of sodium on the
lumen side.
• As the sodium travels into the cell via a membrane
protein, it pulls glucose with it.
• Fructose however enter the cell via passive transport.
• 1. Enzymes on brush border (apical side) break down
diasaccharides.
• 2. Na+/ K+ ATPase found on the basolateral side use ATP
to pump three Na + out of cell and two K+ into the cell.
This establishes an electrochemical gradient a lower Na +
concentration inside.
• 3. The hexose cotransport protein found on apical side
moves sodium down its electrochemical gradient into
the cell, pulling glucose and galactose with it.
• Fructose moves via passive transport by using a different
protein transporter.
4. While inside majority of the fructose is
converted into glucose.
5. Fructose, glucose and galactose then leave via
passive transport on the basolateral side and enter
the blood and travel to tissue.
• Absorption of protein in small intestine
• Although protein undergo mechanical digestion in the mouth,
they do not begin chemical digestion until they are in the
stomach.
• In the stomach pepsin cleaves protein and breaks them down
into polypeptide.
• Once these polypeptides reach the lumen of small intestine,
pancreatic peptidase (trypsin, chymotrypsin, caboxypeptidase)
break down these polypeptide into small peptides.
• These small peptides are ultimately broken into tripeptide
dipeptides and aminoacids by brush border enzymes.
Protein stomach polypeptides

small lumen of small

peptides intestine-
brush border

tripeptide, dipeptides and

aminoacids
• All proteins are digested by proteolytic enzymes into
either their aminoacids constituents or dipeptide
and tripeptides before being absorbed by the cells of
the small intestine (enterocytes).
• Individual aminoacids are absorbed by enterocyte
using sodium dependent cotransport (secondary
transport).
• ATP is used to establish and electrochemical
gradients and then the sodium moves down it
electrochemical gradient and bring the aminoacids
with it.
•Dipeptide and tripeptide use the hydrogen
ion dependent cotransporter system.
•Here, a sodium- hydrogen exchange protein
is used to set up a hydrogen gradient.
•Then the hydrogen ion enters the cell and
brings the di/tri peptide with it.
• 1. Hydrolysis of small peptide at brush border on
apical side.
• 2. Creation of Na gradient by Na /K ATPase on
basolateral side.
• 3. Na+/ H+ exchange on apical side creates H+
electrochemical gradients.
• 4. Co- transport of aminoacid using Na+ movement
down gradients.
• 5. Co- transport of di/tri peptide using H+
movement down gradient.
• 6. Majority of di/tri peptide are broken down
into aminoacids.
• 7. Aminoacids and di/tri peptide exit via
basolateral side and enters the blood system
where they travel to the cells of the body.
• Absorption of lipid
• The bulk of dietary lipid is neutral fat or triglyceride,
composed of a glycerol backbone with each carbon
linked to a fattyacid.
• Food stuffs typically also contain phospholipid, sterols
like cholesterol and many minor lipids, including fat
soluble vitamins.
• Finally small intestinal contents contain lipid from
sloughed epithelial cells and considerable cholesterol
derived in bile.
•In order for the triglyceride to be
absorbed two process must occur.
•Large aggregates of dietary triglyceride
which are virtually insoluble in an aquous
environment, must be broken down
physically and held in suspension- a
process called emulsification.
• Triglyceride molecule must be enzymatically
digested to yield monoglyceride and fatty acids,
both of which can efficiently diffuse or be
transported into the enterocyte.
• The key role in these two transformation are bile
acids and pancreatic lipase, both of which are
mixed with chyme and act in lumen of the small
intestine.
• Bile acids also necessary to solubilize other lipids,
including cholesterol.
• Emulsification, hydrolysis and micelle formation
• Bile acids play their first critical role in lipid assimilation by
promoting emulsification.
• As derivatives of cholesterol, bile acids have both hydrophilic
and hydrophobic domains (i e. they are amphipathic).
• On exposure to large aggregates of triglyceride, the
hydrophobic portion of bile acids intercalate into the lipid, with
the hydrophilic domains remaining at the surface.
• Such coating with bile acids in breakdown of large aggregates
or droplets into smaller droplets.
• Hydrolysis of triglyceride into mono glyceride and
free fatty acids is accomplished predominantly by
pancreatic lipase.
• The activity of this enzyme is to clip the fattyacids
at position 1 and 3 of triglyceride, leaving two free
fatty acids and a 2- monoglyceride.
• Lipase is a water soluble enzyme, and with a little
imagination, it is easy to understand why
emulsification is a necessary prelude to efficient
activity.
• Shortly after a meal lipase is present within the
small intestine in rather huge quantities, but can
act only on the surface of triglyceride droplets.
• For a given volume of lipid, the smaller the droplet
size, the greater the surface area, which means
more lipase molecule can get to work.
• The drug orlistat (Xenical) that is promoted for
treatment of obesity works by inhibiting pancreatic
lipase thereby reducing digestion and absorption of
in small intestine.
• As monoglyceride and fattyacids are liberated
through the action of lipase, they retain their
association with bile acids and complex with other
lipids to form structure called micelles.
•Micelles are essentially small aggregates (4-8
nm in diameter) of mixed lipid and bile acids
suspended within ingesta.
•As the ingesta is mixed, micells bump into the
brush boarder of small intestinal enterocytes,
and the lipids, including monoglyceride and
fattyacids are taken up into the epithelial cells.
• Absorption and transportation into blood
• The major products of lipid digestion – fattyacids
and 2-monoglyceride- enters the enterocyte by
simple diffusion across the plasma membrane.
• A considerable fraction of the fattyacids also enter
the enterocyte via specific fattyacid transporter
protein in the membrane.
• Lipids are transported from enterocyte into blood
by a mechanism distinctly different from what we
have seen for monosaccharide and aminoacids.
•Once inside the enterocyte, fattyacids and
monoglyceride are transpoted into the
endoplasmic reticulum, where they are used to
synthesize triglyceride.
•Bigining in the endoplasmic reticulum and
continuing in golgi, triglyceride is packed with
cholesterol, lipoprotein and other lipids into
particles called chylomicrons.
• Chylomicrons are extruded from the golgi into exocytic
vesicles, which are transported to basolateral aspect of
the enterocyte.
• The vesicle fuse with plasma membrane and undergo
exocytosis, dumping the chylomicron into the space
outside the cells.
• Instead of being absorbed directly into capillary blood,
chylomicrones are transported first into the lymphatic
vessels that penetrates into each villus.
• Chylomicron rich lymph then drains into the system
lymphatic system which rapidly flows into blood.
Digestion in the chicken
• The digestive mechanism in the chicken is considerably simpler
than for omnivorous and especially herbivorous mammals.
• The overall length of the digestive tract is relatively short
(expressed as a multiple of body length) when compared to the
mammalian forms.
• The beak encloses a beak cavity that communicates with the
sublingual cavity, analogous to the buccal cavity.
• Birds lack teeth but certain species are capable of mechanically
reducing ingesta by virtue of sharp edges on the beak.
• In other species, the beak is merely prehensile.
• The tongue of the chicken is essentially inflexible and
incapable of propagated contraction from front or rear as
the first step toward swallowing as in mammals.
• In addition, the beak and nasal cavities are incompletely
separated, there is no hard palate nor are there cheek
muscles.
• Salivary gland occur both in paired and single form and
secrete an acid mucous from sublingual cavity.
• Recall that mammalian saliva is either mucous or serous,
the latter being a dilute buffer solution of slightly alkaline
reaction.
• The oesophagus is a multilayered, striated muscle tube
possessing a mucus- secreting epithelium.
• Distally in the thoracic region and slightly offset to the
right of the median plane, the esophagus is dilated into
the crop, a storage and secretory organ that is filled
with ingesta after the stomach is loaded.
• The stomach has two distinct compartment.
• Anteriorly, the proventriculus, which is thin – walled and
highly glandular, secretes gastric juice consisting of
hydrochloric and enzymes.
• The ventriculus (gizzard) is thick- walled, muscular and
keratinized.
• Its limited secretory function is mainly of mucus and is
presumably protective to the epithelium.
• In its most distal area called the pylorus as in mammals, the
lining possesses villous folds with pyloric glands.
• There is no pyloric sphincter between gizzard and the
duodenum.
• The intestine differs from that of mammals in the virtual lack of
a colon (or large intestine).
• The duodenum loops to enclose multiple lobes of pancreatic
tissue on the mesentery, and two pancreatic tissue on the
mesentery, and two pancreatic ducts and one bile duct enter
the duodenum near the jejunum.
• The ileum terminates in a very short segment of colon from
which two ceca arise, partially isolated by cecal valves.
• Many birds possess, in the distal portion of the small intestine,
a lymphoid structure called Meckel’s diverticulum, a remnant
of the stalk of the yolk sac from embryonic life.
• The short rectum passes to the cloacal structure, consisting of
three parts, each separated by annular folds.
• The proximal portion is a dilated ampulla, and the
middle portion is glandular with the uretal openings, the
vasa deferentia of the male reproductive tract and the
slit-like opening of reproductive tract in females.
• The distal portion is noteworthy because a lymphoid
tissue unique to birds (the Bursa of Fabricius) is located
dorsally.
• Finally, the cloaca leads to the anus (vent), consisting of
two sphincters, an inner smooth muscle, and outer
striated muscle.
• The liver exists as two lobes connected by an isthmus.
• The right lobe possesses a gall bladder supplied by a bile
duct from the hepatic tissue.
• On the left side, the bile duct bypass the gallbladder and
directly supplies the intestine.
• Chicken bile is weakly acidic and possesses some amylolytic
activity.
• Pancreatic juice is similar to that of mammals but is less
basic.
• It is was noted earlier that digestion in chicken occurs
without distinctly alkaline intestinal phase characteristic of
mammals.
• Digestion mechanism
• Ingesta entering the beak, the prehensile structure, and
sublingual cavities is, in effect, crammed back toward the
pharynx because there is little opportunity for any propagated
muscular effort within these most proximal cavities.
• Passage down the esophagus is by slow peristalsis.
• Water intake poses special problems because the nasal
passages are always open and there are no cheek muscles to
create a vacuum for sucking.
• Drinking is achieved by scooping up water in the beak then
elevating the head so the liquid simply flows by gravity into the
esophagus.
• During feeding the proventriculus and gizzard are first
filled, then the crop is used to hold additional ingesta.
• Vagomotor controls regulate emptying of the crop
contents into the stomach.
• Secretory function has both a cephalic and gastric
component.
• The most important specialization stems from the powerful
muscular contractions of the gizzard, which provide a
crushing and grinding action on the gastric contents.
• When the tract is dissected, gritty mineral material is often
found in the gizzard.
• Domestic birds, especially layers, are supplied
with grit such as crushed shells and limestone
as part of diet.
• The strongly acidic conditions favour
solubilization of calcium so much of the mineral
residue is material other than calcium salts.
• The ceca provide for microbial digestion of fibre
and serve as a secondary site of absorption.
• The major location of absorption of digested nutrients is
the epithelium of the ileum.
• The rectum has some capacity for water absorption.
Fecal material is brownish in color, but is voided as a
mixture with whitish uric acid crystals from urinary
excretion.