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Isfahan University of Technology

Isfahan, Iran

Advanced
Digestive Physiology
(part 3)

By: A. Riasi
(PhD in Animal Nutrition & Physiology)
http://riasi.iut.ac.ir
Rumen and reticulum properties

The ruminoreticular fold partially separates the


reticulum and cranial sac of the rumen.

Pillars divide the rumen to dorsal and ventral


parts.
Rumen and reticulum properties

The dorsal part of rumen is divided to:


Cranial sac

Dorsal sac

Caudodorsal blind sac

The ventral part of the rumen is divided to:


The ventral sac

The caudoventral blind sac


Ruminoreticular wall structure

Highly vascularized connective tissue core is


in the mucosal layer of the finger-like papillae

Epithelium is involved in the absorption of


short-chain fatty acids.
Ruminoreticular wall structure

In the reticulum the mucosal layer contains


laminae.

Conical projections (papillae) are found on


the surface of laminae.

The stratified squamous epithelium of


reticulum also absorb short-chain fatty acids.
Ruminoreticular wall structure

The reticulum has a role in mechanical


digestion of food.

Myenteric plexus can be found between the


muscularis externa layers.
Blood circulation of forestomachs

Celiac artery supply the blood flow to


forestomach and the most part of abomasum.

The venous blood drains into the hepatic


portal vein.
Innervations and the receptors

The innervations of fore stomachs:


Vagal nerves (10:1 afferent/efferent ratio)

Splanchnic nerves (3:1 afferent/efferent ratio)


Innervations and the receptors

The vagal nerves transmit sensory information


from two known kinds of sensory receptors:

Tension receptors

Epithelial/mucosal receptors
Innervations and the receptors

The splanchnic nerves transmit sensory


information from:
Serosal receptors

Possibly tension receptors


Ruminoreticular motilities

The movements serve to:


Mix the ingesta

Aid in eructation of gas

Propel fluid and fermented foodstuffs into the


omasum.

A cycle of contractions occurs 1 to 3 times per


minute.
Ruminoreticular motilities

Two types of contractions are identified:


Primary contractions

Secondary contractions
Ruminoreticular motilities

Typical primary cycle consist of:


Biphasic (double) contraction of the reticulum

Caudally moving monophasic contraction of the


dorsal ruminal sac

A contraction of the ventral ruminal sac


Ruminoreticular motilities

Secondary cycle may occur and consist of


sequential contractions of:
The caudoventral ruminal blind sac

A cranially moving contraction of the caudodorsal


ruminal blind sac followed by the middorsal
ruminal sac

A contraction of the ventral sac.


Ruminoreticular motilities

When ingesta enter the foestomach, heavy


objects fall into the reticulum and lighter
material enters the rumen.

Added to this mixture are voluminous


quantities of gas produced during fermentation.
Ruminoreticular motilities
Ruminoreticular motilities

The forestomachs possess a rich enteric


nervous system.
Contractions coordination need the central input.

Motility centers in the brainstem control


both the rate and strength of contraction via
vagal efferents.
Ruminoreticular motilities

There are also vagal afferents from the rumen


to the motility centers which allow stretch
receptors and chemoreceptors in the rumen to
modulate contractility.
Ruminoreticular motilities

Feeble intrinsic contractions responsible for


the smooth muscle tone in the forestomach wall
arise from nervous activity in its intrinsic nerve
networks.
Ruminoreticular motilities

The gastric centers do not have spontaneous


activity and need to be driven by
Excitatory inputs

Inhibitory inputs
Ruminoreticular motilities

The principal inputs to the gastric centers are


from:
Forestomachs

Abomasum

Duodenum by way of vagus nerves


Stimulus Receptors Projections Effect
Stimulus Receptors Projections Effect
Stimulus Receptors Projections Effect
Ruminoreticular motilities

The known sensory receptor mechanisms are


responsible for the vagal inputs.
The tension receptors are located in the muscle
layer of different parts.

The epithelial receptors are located closed to the


basement membrane of the luminal epithelium of
the forestomachs.
Ruminoreticular motilities

Conditions inside the rumen can significantly


affect motility.
Acidic ruminal contents

High roughage diet


Rumination and its components
Ruminants are well known for "cud chewing
It provides effective mechanical breakdown of roughage
and increases substrate surface area.

Rumination is a unique characteristic of:


True ruminants (deer, giraffes, and bovidae)

Pseudoruminants (camels and llamas).


Rumination and its components

Rumination occurs in resting.

The highest incidences of rumination occur


during afternoon and middle of the night.
Many lactating ruminants ruminate while they are
suckling their young or are being milked.
Rumination and its components

The time spent ruminating by a given animal


depends on:
The texture of the food

The amount of food ingested

Cattle may ruminate from 35 to 80 minutes per


kilogram of roughage consumed.
Rumination and its components

Pharmacological agents have been used to


examine the physiological mechanisms
involved in evoking rumination.
Volatile fatty acids

Catecholamines

Gastric hormones

Opioids

Autacoids
Rumination and its components

Rumination is centrally mediated by the


"gastric centers located at:
Medulla oblongata

Ventral hypothalamic area

Tactile stimulation of the reticular and ruminal


epithelia is a powerful stimulus for rumination.
Rumination and its components

Sensory information from digesta that is


perceived in pillars:
Digesta texture

Digesta consistency

Rumen fill
Rumination and its components
Rumination and its components

o Four phases of a rumination cycle:


o Regurgitation

o Remastication

o Reinsalivation

o Reswallowing

o Regurgitation starts with an inspiration effort.


Rumination and its components

o The bolus is carried into the mouth by reverse


peristalsis.
o Antiperistaltic waves of regurgitation passes over the
esophagus at a velocity of 0.2 m per second.
Rumination and its components

o In about two seconds, the "retained" bolus has the


fluid squeezed out of it.
o This fluid is immediately swallowed, and the remainder is
chewed, reinsalivated, and subsequently swallowed.
Rumination and its components

o Slower and more regular rate in remastication


phase
Rumination and its components
Rumination and its components

o The rate and duration of rumination chewing are


controlled:
o The texture (coarseness)

o Quantity of the food


Rumination and its components

o The reinsalivation phase of rumination coincides


with the remastication phase.

o Rumination saliva and ingestion saliva are secreted


at about the same rate and about 2.5 times the resting
rate.
Eructation

oEructation is the physiological process of expelling


ruminoreticular gases:
o Carbon dioxide, 65%;

o Methan, 25%;

o Nitrogen, 7%;

o Oxygen, 0.5%;

o Hydrogen, 0.2%;

o Hydrogen sulfide, 0.01%.


Eructation

o The eructation is associated with almost every


secondary ruminal contraction.

o Eructated gas travels up the esophagus at 160 to 225


cm per second
o Interestingly, a majority is actually first inspired into the
lungs, then expired.
Eructation

o A large portion of the eructated gases is inspired and


recycled into the organism by absorption into the
lungs.
Eructation

o The rumen raft is the site of origin of most of the


gases of fermentation.
o When dorsal surface of raft splits, releasing of free gas into
the gas layer occur.

o Not much of the gas is absorbed, and most must be


eliminated by eructation.
Eructation

o If the gas layer reaches the cardia and clear it of


fluid, the eructation mechanism is evoked.

o Eructation is a vagovagal reflex, with centers in the


medulla ablongata.
o Mechanical receptors to detect distention are present in
o Rumen dorsal sac

o Reticular groove

o Around the cardia and esophagus


Eructation

o The primary type of rumen contraction always


follows the biphasic reticular contraction and spread
caudally.
o In cattle

o 66% with the secondary type of rumen contraction,

o 20% with the primary type.


Eructation

o In sheep

o 3% no dependent rumen motility

o 37% with secondary-type rumen contractions

o 60% with the time with the primary type


Strategies for lower methane emission

o Increasing the efficiency in which animals use


nutrients to produce milk or meat.

o Rumen modifiers such as ionophores improve dry


matter intake efficiency and suppress acetate
production, which results in reducing the amount of
hydrogen released.
Strategies for lower methane emission

o The grinding and pelleting of forages can reduce


emissions by 40% .

o Dietary fats have the potential to reduce CH4 up to


37%.

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