Digestion in Ruminants
• Buccle cavity
• Mestication
• Chewing
• Regurgitation/rumination
• Reticular groove
Ruminant Stomach:
• Compound stomach, four parts.
• Fore stomach acts as fermentation vat.
• Abomasum like non-ruminant stomach i
• In fore stomach ( rumen, reticulum, omasum)
microbial fermentation of ingesta, mainly by
hydrolysis and anaerobic oxidation.
• Microbes generate ATP for their own purpose
& VFA production, fermentation end products
used as energy source.
• Rumen is biggest of all, divided into dorsal
and ventral sacs & papillae.
Intestine Dorsal part
Ventral part Similar to stomach of
Non-Ruminants
Rumeno-reticulum occupies the entire left
side of the abdomen.
Reticulum is approx. spherical, honeycomb
like structure.
Ruminoreticulum operates as a combined
functional unit.
The main function of the reticulum is to
collect smaller digesta particles and move
them into the omasum, while the larger
particles remain in the rumen for
further digestion.
The reticulum also traps and collects heavy/dense
objects the animal consumes.
Reticular groove – By pass the rumen in young
ones.
Abomasum is almost like stomach of non-
ruminants.
Blood supply is rich, further increases when
fermentation products are being absorbed.
Rumen
Reticulum
• Omasum is kidney shaped, larger in cattle
than sheep & goat, body consists of
parallel leaves, attached at greater
curvature, lesser curvature forms omasal
orifice/canal, leaves bear small papillae to
increase surface area.
• The omasum is where food particles that
are small enough get transferred into the
abomasum for enzymatic digestion. The
large surface area of omasum allows it to
play a key role in the absorption of water,
electrolytes, volatile fatty acids, minerals,
and the fermentation of food.
Rumen Ecosystem or Environment
1. Anaerobic conditions
2. Watery environment- water intake, saliva,
urea in saliva, type of ration etc.,
3. pH is maintained for microbial survival,
buffering substances. pH range – 5.7 to 7.3.
Normal pH of rumen 6.8
4. Antifoaming agents, prevents bloat. (Trochar)
5. Digestion as well as absorption activities.
6. End products of digestion & metabolism-
VFAs (Acetate, 65-75%, Propionate 15-20%,
butyrate 5-15%, Valeric, isoval ) lactic acid,
NH3, HCo3, PO4, Na, K etc.
Mon
frolgenous roger
Uber
Micro-organisms in Rumen.
to /ml SRL lo to to meIRL
Bacteria: 109 to 1010
Protozoa: 105 to 106 /ml SRL
Fungi:
toot
104 to 105 /ml SRL its
Bulk (volume) of bacteria & protozoa same. c
Syntropic association.
Existance: i. Free in liquid medium - 30 %
ii. Attached to feed particles - 70 %
iii. Adhering to epithelium of rumen wall,
vi. Bacteria attached to protozoa
Bacteria Classification: On the basis of substrate they act.
1. Cellulose fermenting:
a. Bacteriodes succinogens
b. Ruminococcus albus
c. Ruminococcus flavifaciens
d. Butyrivibrio fibrisolvens
2. Starch & Sugar fermenting:
Bacteriodes amylophyllus
Selenomonas ruminantium
Streptococcus bovis
Succinovibrio dextrisolvens
3. Lactic acid fermenting: Veillonella alcolescence
4. Deaminating: Bacteriodes ruminocola
5. Methanogenic: Methanobacterium ruminantium
Protozoa
Classification: On the basis of body structure.
Two broad catagories:
1. Holotrichs or Trichostomatida:
Body covered with cilia, further of two types.
a. Isotricha: Large speciess.
i. Isotricha prostom
II. Isotricha intestinalis
b. Dasytricha: small species.
Entodiniomorphids:
Small species as well as large sps.
Entdenium: no skeletal plate, one tuft of
cilia.
Diplodenium: no sk plate, two tuft of cilia.
Eudiplodenium: one narrow sk plate.
Ostraconidium: broad sk plate.
Polyplastran: 2 or 3 sk plates
Metadenium: resemble polyplastran, 2
narrow sk plates.
Dietary substrates:
Cellulose: β-1,4 linkage, glucose polysaccharide
Hemicellulose: β-I,4 linkage, xylose pentose
branched molecules,side linkages.
Variety of other monosaccharides like arabinose,
mannose & galactose.
Lignification, silica require wider range of enzymes than
cellulase
Pectin: Polysaccharide β-1,4 galacturonan (galactose with
uronic acid). Enzymes for these found only in
microbes.
Grains/concentrates: Non structural, starch & fructosans
Polysaccharide of glucose, ά-1,4 linkage, branching
ά-1,6 linkage after 24 units.
Fermentation Pathways.
Takes place in four Stages:
i. Hydrolysis of plant polysaccharide to monosacc --
- Fructose-1,6 diphosphate (cellulose,
hemicelluloses)
ii. Anaerobic oxidation of Fructose-1,6 di PO4 to
pyruvate via PEP by glycolysis.
iii. Production of final metabolites of fermentation,
Pyruvate or PEP is main product leading to
further prodn of acetate, propionate, butyrate,
formate, H2 , CO2, etc.
iv. Synthesis of microbial compounds, particularly
amino acids using stage I to stage III
intermediates & ATP is derived.
Cellulose Fermentation:
PPT
β-1 linked compounds ( cellulose, hemicellulose,
5
fructosans & pectins) & four stage microbial activity,
except CH4 prodn by cellulolytic bacteria.
Cellulose fermentation is a slow process, cellulolytic
bacteria have low metabolic rate, take about 18 hrs to
double their numbers (doubling time)
For protein synthesis, cellulolytic bacteria do not need a.a.
supply, but need NH3. Stages 2&3 intermediates &
isoacids, (deamination of branched chain a.a.) of dietary
plant proteins.
pH optimum 6.2-6.8
Methanogens also require similar pH & supply of
formate, CO2, & H2.
Cellulolytic & methanogens produce VFAs, CO2, & CH4
Starch Fermentation
• α-linked starches (amylose, amylopectin) and
simple sugars (sucrose, maltose) by
amylolytic bacteria.
• Some of these are capable of all four stages of
process except methane production.
• Faster fermentation rate.
• Doubling time much shorter (0.25 – 4 hrs)
• pH optimum, 5.5 – 6.6,
• High concentrate diet (starch rich), high VFA
prod, propionate increases, ratio 70, 25, 05 %.
• Less production of 2H, hydrogen sink.
• Less methane production.
Starch fermentation contd.
• Amylolytic bacteria Need NH3 + some amino
acids for protein synthesis
• Secondary bacteria required for methane
production & for converting lactic acid to
propionate. These bacteria have long doubling
time (16 hr, pH- 6.2 – 6.8)
• Number of protozoa increase with concentrate
feed, engulph starch + bacteria and curb
bacterial amylolysis until pH falls below 5.5, at
this pH bacteria are quickly inactivated and later
die.
Fermentation of Proteins
Proteolytic bacteria, 12 – 38 %
About 50 % dietary proteins degraded in rumen,
rumen degradable proteins, rumen undegradable
proteins.
Bacterial proteolysis produce amino acid (a. a.) &
ammonia, isoacids like isobutyrate, a isovalerate
from leucine, isoleucine & valine.
Ammonia from endogenous & NPN compounds.
E.g. Urea in saliva.
Ammonia imp. substrate for microbial protein
synth, provided feed should provide sufficient
protein, NPN & readily fermentable CHO.
Feeding excess protein is wasteful practice.
Fermentation of Lipids
Fermentation is a continuous process.
Generally lipids stored in oil seeds(65- 80%), forage
plants, 3-10 %, in cell membrane.
50 % lipids as free FAs. Majority are phospholipids,
palmitic and linolenic acids.
Ruminal microbes also synthesize microbial lipids from
VFAs.
High levels of lipids have adverse effect on:
Food palatability
Cellulolytic activity
Appetite & fore stomach activity.
Protozoa play imp. role in lipid metabolism, they absorb
some PUFAs, protect them from saturation &
hydrogenation, make them available in intestine.
Fate of End Products of Fermentation
• Most of TVFAs absorbed across rumen wall.
• Fermentation of proteins also yield VFA with isoacids (5
% of total), more valuable to microbes for microbial
protein synthesis using NPN.
• At pH 6.6 most of them exist as their anions.
Absorption rates are higher : Butyrate Ketone Bodie
a. When ruminal pH is reduced Profinot Gilmore
b. As chain length increases e.g. butyrate more than
propionate & acetate.
About 50 % VFAs absorbed by passive diffusion.
Butyrate. Metabolized to ketone bodies which are
metabolized by most tissues of the body. In mammary
glands about half of short & medium chain FAs (C4-C16)
in ruminant milk.
Propionate: About 30 % metabolized in free form to lactic
acid. Lactic & propionic acids completely removed by liver,
converted to glucose, gucogenic, gluconeogenesis.
Acetate: a. Acetyl-COA , TCA cycle
b. In mammary glands- similar to butyric acid
c. A small amount- CO2 Acetate to FCAcych
Got
d. Mostly, acetyl-coA- TCA/Krab’s cycle
e. FAs synthesis
f. Glucose by gluconeogenesis
Lactale: Converted to propionate by secondary bacteria.
At low pH (3.8-4.6) absorption about 10 % to VFAs
mobilized by liver to pyruvate – glucose
(gluconeogenesis)/glycogen
Gases: Co2 60%, CH4 30-40 %, N2, traces of H2S, H2, O2
Other Important Ruminal Reactions.
• Synthesis of Vitamin B-Complex by microbes
• Adequate supply of sulpher, s-contg. a.a.
synthesis, wool production is affected.
• Mb excess (in some areas) affects availability
0f S & Cu. so
• Excess S, forms insoluble cupric sulphide,
render Cu unavailable
• Cu deficiency reduce melanin synthesis, loss
of crympiness in wool.
• Due to microbial action some toxic dietary
constituents rendered harmless e.g. Oxalates
are converted to formates & CO2.
Woofforoduction M
Modification of Toxic Substances
Some plants/substances are more toxic to non-
ruminants than ruminants because the rumen microbes
degrade them to non toxic. Microbes act as 1st line of
defense.
Important role of microbes:
1. Lactate acidosis: High conc./starch diet – lactate
production may drop rumen pH from 6 to 5 or less, normal
working of microbes affected if pH is less then 5.5. Rumen
microbes convert lactate to propionate.
2. Nitrate- nitrite toxicity: Drought stress, over fertilization of
cereal & new crops production of feed stuffs with toxic
levels of nitrate. A number of rumen bacteria & protozoa
reduce nitrate to nitrite. Nitrite is further converted to
ammonia
3. Glucosides: Some organic nitro compounds, e.g.
glycosides are rapidly hydrolyzed by rumen
microbes.
4. Oxalic acid toxicosis: Oxalic acid is present at low
levels in plants, when exceeds more than 10 % of
DM, causes toxicity, bind to circulating Ca. Rumen
microbes degrade oxalates to formate & carbon
dioxide. Animals should be switched over to
oxalate diets gradually.
5. Mimosine: Leucaena leucocephala- leguminous
tropical shrub, rich in protein (about 20%), causes
toxicity if supplemented more than 30 % in diet.
Toxicity due to mimosine which is degraded by
microbes to non-toxic products.
Stomach Motility
Stomach can be divided into 3 zones:
Dorsal portion or fundus- reception and storage of
contents & adaptation to the volume of contents,
excessive pressure is not developed.
The body or corpus- a vat for mixing saliva and gastric
juice with food.
The antrum is the gastric pump, regulates the
propulsion of food past the pyloric sphincter into
duodenum.
The pacesetter potential originate on the greater
curvature of the stomach and spread circumferentially
and distally, moving as a ring.
The peristaltic contractions of the distal part of
stomach mix the gastric juice with food, grind the solids
present in stomach & effect propulsion of contents
through antrum towards pylorus. Liquids pass & solids
retained & forced back into the body for grinding.
• Rate of emptying different in different species.
• Distention of stomach increase gastric motility
• Mechanical receptors activated
• Peristaltic contractions, 4-5/min
• Antral contractions by slow waves, which are
initiated on greater curvature & spread distally
with increasing velocity & voltage.
• These are governed by neural & hormonal
influences.
Ruminant Stomach
ppr 6
Format for Internal Assessment (B V Sc & AH)
Q 1. Fill in the blanks 10 x .5 = 5
2. Multiple choice 10 x .5 = 5
3. Matching question 10 x .5 = 5
4. Define/ justification/ 10 x 1 = 10
exlanation/diagram/
equation/enumerate.
5. Difference/short note 4 x 2.5 = 10
6. Detailed subjective 1x5=5
Total marks = 40 Weightage = 10
Matching: Match the answer of column A from column B &
write in answer column.
Column-A Answer Column-B
Q. No. 3 Match the Column-A with the proper answer from
Column-B & write the answer In answer column.
[Link] Column A Answer Column B
1 Haustra Pseudopodia
2 Camel Large intestine
3 Rumen Somach
4 Chief cells Microbes
Role of Protozoa
Feed upon bacteria, plants, starch granules &
other readily digestible materials including
polyunsaturated fatty acids
Protozoa are very sensitive to rumen environment,
their presence in rumen is indication of normality.
Most of the protozoa are in the raft of fibrous
digesta in dorsal rumen sac.
Reservoir of microbial protein
Help to prevent over proliferation of bacteria at
time of starch loading by engulfing starch
particles thus prevent high rate of starch
degradation by amylolytic bacteria.
When passed to abomasum, they provide:
High biological value microbial protein.
Unfermented starch.
Some PUFA, which otherwise might have been
saturated.
Protozoa, primary cellulolytic bacteria & most
secondary bacteria require pH 6.2 or above.
Other Important Ruminal Reactions.
• Synthesis of Vitamin B-Complex by microbes
• Adequate supply of sulpher, s-contg. a.a. synthesis,
wool production is affected.
• Mb excess(In some areas) affects availability 0f S & Cu.
• Excess S, forms insol cupric sulphide, render Cu
unavailable
• Cu deficiency Reduce melanin synth, loss of
crympiness in wool
• Due to microbial action some toxic dietary constituents
rendered harmless. Oxalates are converted to formate
& CO2.
Modification of Toxic Substances
Some plants/substances are more toxic to non-
ruminants than ruminants because the rumen
microbes degrade them & make non toxic
substances act as 1st line of defense---Imp. role
of microbes.
1. Lactate acidosis: High concentrate/starch diet
lead to high lactate production may drop rumen
pH from 6 to 5 or less, normal working of microbes
affected, if pH is less then 5.5, rumen microbes
convert lactate to propionate.
2. Nitrate- nitrite toxicity: Drought stress, over
fertilization of cereal & new crops, production of
feed stuffs with toxic levels of nitrate. A number of
rumen bacteria & protozoa reduce nitrate to nitrite.
Nitrite is further converted to ammonia
3. Glucosides: Some organic nitro compounds, e.g.
glycosides are rapidly hydrolyzed by rumen
microbes.
4. Oxalic acid toxicosis: Oxalic acid are present at
low levels in plants, when exceeds more than 10 %
of DM, causes toxicity, bind to circulating Ca.
Rumen microbes degrade oxalates to formate &
carbon dioxide. Animals should be switched over
to oxalate diets gradually.
5. Mimosine: Leucaena leucocephala- leguminous
tropical shrub, rich in protein, causes toxicity if
supplemented more than 30 % in diet. Toxicity due
to mimosine, which is degraded by microbes to
non toxic products (more in sheep & goat).
Stomach Motility
Stomach can be divided into 3 zones:
Dorsal portion or fundus- reception and
storage of contents & adaptation to the volume
of contents, excessive pressure is not
developed
The body or corpus- a vat for mixing saliva and
gastric juice with food.
The antrum is the gastric pump, regulates the
propulsion of food past the pyloric sphincter
into duodenum.
The pacesetter potential originate on the
greater curvature of the stomach and spread
circumferentially and distally, moving as a ring.
Stomach Motility
• The peristaltic contractions of the distal
part of stomach mix the gastric juice with
food, grind the gastric solids & effect
propulsion of contents through antrum
towards pylorus, liquids pass & solids
retained & forced back into the body for
grinding.
Rate of Gastric Emptying
Rate of emptying different in different species.
Liquid diet move faster than solid.
Distention of stomach increase gastric motility.
Antral contractions by slow waves, which are
initiated on greater curvature & spread distally
with increasing velocity & voltage.
These are governed by neural & hormonal
influences.
Distention of stomach with food.
Mechanical receptors activated.
Peristaltic contractions, 4-5/min.
Small Intestine motility.
Function- Digestion & absorption
Flow of contents regulated for mixing with
secretions of pancreas & bile and
maximum exposure of undigested &
digested nutrients to the mucosa of SI for
digestion & absorption.
Primary pacemaker located in duodenum
near the entrance of bile duct.
Slow waves generated by pacemaker @
14-18/min in different species. Waves move
aborally & slows down in lower SI (10-
11/min)
Small intestine motility
Ileocecal junction provide some delay in
transit, contents are retained more longer in
ileum than jejunum.
Peristaltic waves induce propulsion-
contraction & relaxation of circular &
longitudinal muscles
Segmentation, important for mixing by
inducing back & forth mixing.
Spiral movements.
If intestine is distended/irritated, local reflex
initiated, movement is fast.
Pendular movements.
Motility of Large Intestine
•Microbial digestion,
• Absorption of water & electrolytes
•Synthesis of vit. B-comp. & vit.-K
•Slow movement as above functions require
transit be delayed.
•Cecal contractions results in mass
movementof ingesta to colon
•Contraction for gas transport (cecum+colon)
•Mixing movements
•Aboral peristaltic movements
•Mass movements
Defecation
Faeces are discharged from rectum.
Stimulus- distention of terminal colon and rectum
Defecation reflexes are cholinergic,
parasympathetic drugs make it more vigorous.
Strong peristaltic movements.
Contraction of longitudinal muscles of rectum.
Relaxation of internal and external sphincters.
Frightened animal frequently defecate-
stimulation of reflex center in brain.
Vomiting or Emesis
Complex reflex activity, can be initiated by many
stimuli
Emotion 2. Pain
Migraine 4. Chemical comp. of food
5. Drug ingestion 6. Alcohol ingestion
7. Rotation 8. Pharyngeal irritation
9. Peritonitis 10. Renal failure
11. Liver failure..
Carnivore & omnivore vomit easily (dog, cat,
swine) In ruminants vomiting does not occur,
some times ejection of abomasal contents in fore
stomach occurs. In horse vomiting is extremely
rare.
Electrical slow waves amplitude decreases in
duodenum and superseded by long burst of
intense spikes activity spreading towards mouth.
Contents of small gut to stomach, to esophagus,
to mouth.
Mechanoreceptors in pharynx.
Tension & chemo receptors in gastric and
duodenal mucosa.
Vomiting center (in chemoreceptor trigger zone)
area in brain stem close to 3rd ventricle, drugs &
toxins in blood may stimulate vomiting.
Hunger Contractions
Intense contractions when stomach is empty for a
long time. Rhythmic peristaltic contractions,
mixing waves in stomach becomes extremely
strong, cause a continuous tetanic contractions
lasting 2-3 min.
Sensation of pain in stomach, hunger pangs.
Don't begin before 12-24 hrs after the last meal.
Starvation- reach greatest intensity at 3-4 days &
then gradually weakens.
Appetite , Hunger Control
Permeability of intestine, absorption
regulation, transport of water & electrolytes.
Epithelium contains absorptive, goblet, endocrine
& other cells in lamina properia, immune cells,
lymphocytes, phagocytes, mast cells.
Water & electrolytes transport in gut and kidney
affected by hormones – adrenal epinephrine,
aldosterone.
Angiotensin-II increase salt & water absorption in
intestine
Hexose's & amino acids absorbed by special
active transport mechanism.
Large intestine does not posses active transport
system for hexose's & amino acids.
Absorption in neonates: Intestine has ability to uptake
large amt. of intact proteins. In adults it is in trace amt.
primary cause of food allergy.
Carnivores & omnivores, proximal intestine- about 60%of
total fluid, remainder in ileum & Large Intestine
In post gastric herbivores, proximal intestine Very little
absorption, major in distal SI & LI
Salt & Water absorption
Water & electrolytes across the epithelium.
Na enters via epical membrane Na pump.
Lipid soluble substances against conc. Gradient.
Osmotic, hydrostatic or electrochemical gradients are
present across the epithelium.
Water absorption across the epithelium is strictly a
consequence of osmotic and hydrostatic gradient setup by
baso-lateral Na-K pump.
Absorption 0f CHO
Glucose, fructose & galactose all are completely
absorbed, speed differs
Speed - glucose – 100 %, galactose – 110 %
Fructose -- 40 %
Amino acids by Na dependent co-transport
mechanism
Enteric epithelium has highest turnover of any
tissue, highest regenerative capacity. Turn over
rate differs with age e.g. new born pig - 7 -10 days.
3 wk old pig - 2 - 4 days.
VFAs: Short chain FA readily absorbed from
rumen & colonic mucosa.
Active Transport- Energy supplied by Na, K, ATPase pumps
Uniport: Transport by a carrier for a single compound e.g. Na+
called uniport.
Symport: Simultaneous transport of two compounds in same
direction e.g. Na & Glucose or Na & a.a.
Antiport: Movement of a compound in one direction, driven by a
movement of second compound in opposite direction
e.g. Na & H.