Ruminant Digestion

Integrative Veterinary Physiology Ruminant Digestion
Dr G W Reynolds Room 2.17 Riddett Building Institute of Food Nutrition & Human Health Extn. 7507 Email G.W.Reynolds@massey.ac.nz
GWR 227206 Integrative Veterinary Physiology
Herbivores

One of the most successful groups of terrestrial animals Their main energy source is plant structural carbohydrates (cellulose, lignin). Mammals do not produce cellulase Herbivores have established a symbiotic relationship with suitable microbes (produce cellulase) Specialised regions within the GI tract that serve as large fermentation chambers. In the odd toed ungulates e.g. the horse, this fermentation chamber is a greatly enlarged hindgut. In ruminants there is an enlarged region between the oesophagus and the gastric stomach the forestomach [Latin ruminare means to chew again]
Advantages of Ruminants
The forestomach allows ruminants to exploit as their food source the most abundant carbohydrate form on the planet. It allows ruminants to thrive in niches where the quality of the herbage is too low to support nonruminants For example above the Arctic Circle (musk oxen), in high mountains (llamas, yaks) and in the hot deserts (camels, goats).
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Advantages of Ruminants
Tree goats - Morocco
The microbes synthesis relatively high quality proteins from

low quality plant proteins non-protein nitrogen recycled nitrogen-containing end products of metabolism (e.g. urea)
The high quality microbial proteins become available to the ruminant when the microbes are digested The microbes supply the host animal with vitamin B complex, provided there is an adequate supply of the trace element cobalt for B12.
Disadvantages of a Ruminant
Ruminants must spend

~ 4-7 hours per day gathering food ~ 8 hours per day chewing the cud
Failure to effectively expel the large volumes of gas produced during fermentation can be life threatening Ruminants have no direct control over the digestive activities of the microbes Can indirectly influence the rate of fermentation
End products of fermentation are mainly the volatile fatty acids (VFAs)
Acetic acid
acetic, propionic and butyric acids,
Pathways for intermediary metabolism must be geared to their use. Propionic acid is the only VFA capable of being converted to glucose The ruminant has a high glucose requirement during the later stages of foetal growth and lactation.
Propionic acid
Butyric acid
In the wild state these disadvantages may be of little consequence. Under intensive farming systems easily fermented concentrates often replace the normal roughage diet Physiological strategies regulating fermentation may be unable to cope with the greatly increased rate of VFA production Can cause a variety of digestive and metabolic disorders.
Ruminant Stomach
Ruminants have a single compound stomach divide into 4 distinct divisions. Fermentation occurs in the first three, reticulum, rumen and omasum, which are non-secretory Collectively referred to as the forestomach. The 4th division, abomasum, is the true stomach and is analogous to the simpler form of stomach found in dogs, cats and humans It secretes hydrochloric acid, which kills the microbes and sterilises the digesta. Pepsinogen produced by the abomasum begins the breakdown of protein, mainly microbial cell wall protein.
Ruminant Stomach Right Side
ventral sac
abomasum omasum
reticulum
Ruminant Stomach left Side

atrium dorsal sac
Caudal Blind sac
abomasum reticulum Ventral rumen
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Forestomach
Reticular epithelium is thrown into folds that form polygonal cells that give it a honey-combed appearance. Numerous small papillae stud the interior floors of these cells
The interior surface of the rumen forms numerous papillae that vary in shape and size from short and pointed to long and foliate.
The inside of the omasum has broad longitudinal folds, reminiscent of the pages in a book. The omasal folds represent ~ 1/3rd of the total surface area of the forestomach
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Innervation of Ruminant Stomach
The innervation to the stomach

Vagus nerves splanchnic nerves, Both supply motor (efferent), and sensory (afferent) nerves.
The left and right vagi merge in the lower thoracic region and then divide to form dorsal and ventral branches Vagal (parasympathetic) efferent activity is essential for the orderly and sequential contractions of the forestomach Stimulation of splanchnic motor (sympathetic) nerves inhibit motility
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to celiacmesenteric ganglion
to liver & pylorus Rumen Dorsal vagus
Ventral vagus Omasum Abomasum Reticulum
From: Dyce, K. M., et al (1996) Textbook of Veterinary Anatomy. 2 nd Edition. W.B Saunders Company
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There is a predominance of sensory over motor fibres in the vagal (10:1) and splanchnic (3:1) nerves, hence both are predominantly sensory nerves The vagal sensory fibres innervating the stomach are associated with:
Tension receptors

slowly adapting mechanoreceptors located in the muscle layers activated by
passive distension of the stomach active contraction of the smooth muscle of the stomach wall
Epithelial/mucosal receptors.

rapidly adapting mechanoreceptors and chemoreceptors are most numerous in the reticulum, cranial sac of the rumen, abomasum (and duodenum) respond to tactile stimuli (light brushing), acids, alkali and hyper- and hypotonic solutions
The splanchnic sensory fibres transmit information from serosal receptors, which are especially concentrated at the attachment of the mesenteries.
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Blood Supply to Ruminant Stomach
The blood supply to the stomach is via branches of the celiac artery. Blood flow to the stomach increases markedly after feeding when fermentation end products are being absorbed. The nutrient rich blood drains into the portal vein and passes through the liver before being returned to the heart via the caudal vena cava.
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Blood Supply to Ruminant Stomach
From: Dyce, K. M., et al (1996) Textbook of Veterinary Anatomy. 2nd Edition. W.B Saunders Company
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Microbial Fermentation
The microbes in the forestomach are mostly anaerobic bacteria, but also yeast-like fungi and protozoa. They break down the ingesta by hydrolysis and anaerobic oxidation. The protozoa feed on ruminal bacteria, starch and polyunsaturated fatty acids in the ingesta. They are very sensitive to changes in intraruminal conditions and their presence in ruminal fluid is a good indicator of its normality. Very little is known about the importance of the ruminal fungi in fermentation.
Rumen bacteria
Rumen protozoa
Rumen fungi
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Microbial Fermentation
The plant structural carbohydrates in roughage are -1,4 linked glucose units (cellulose) and -1,4 linked xylose units (hemicellulose)
Cellulose
Slowly broken down by the ruminal cellulolytic bacteria to VFAs.
Carbohydrate in grain-based feed contains linked glucose units
rapidly degraded by amylolytic bacteria to VFAs and lactic acid.
Both types of bacteria are classified as primary bacteria Secondary bacteria convert the lactic acid to propionate and hydrogen to methane. This last reaction re-oxidises reduced coenzymes making them available again as hydrogen acceptors. A small amount of O2 is taken in with the food and water; some O2 diffuses from arterial blood across rumen wall
Starch
Quickly used up by facultative anaerobic bacteria
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Ruminant Digestion

The digesta in the forestomach do not form an homogenous mass Instead the most recently ingested food is added to a raft of fibrous material that floats on the underlying soupy fluid Above this raft is a layer of ruminal gas During clinical examination, palpation of the lumbar fossa should detect the gas layer above the textured fibrous raft. Failure to remove (by eructation) the gas causes distension of the stomach (bloat) Excessive hardness of the raft is a sign of ruminal impaction, and softness or absence of the raft indicates the animal has not recently consumed roughage
From: Dyce, K. M., et al (1996) Textbook of Veterinary Anatomy. 2nd Edition. W.B Saunders Company
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Digestion in the Rumen

Average composition of the mixed volatile fatty acids from the rumen and large intestine, given as percentages of total acid. (Adapted from AT. Phillipson, Nutr. Abstr. Rev., 17:12-18, 1947-48.)
Animal Sheep Ox Horse Dog
Organ Rumen Rumen Caecum Colon
Acetic 64 71 73 51
Propionic 20 15 20 36
Butyric 16 14 7 13
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Digestion in the Rumen
Most of the VFAs produced by the microbes are absorbed into the venous blood across the wall of the forestomach and used by the ruminant as substrates for metabolism. About half of the VFAs are absorbed in an undissociated state by passive diffusion the remainder is absorbed by facultative diffusion in exchange for bicarbonate During absorption, most of the butyrate is metabolised to -hydroxybutyrate
used as a substrate by most tissues, and especially the mammary gland during lactation.
About 1/3 of the propionate is metabolised to lactic acid during absorption. Most of the acetate is absorbed unchanged.
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Protein Digestion/Regeneration
About half the dietary protein is degraded in the forestomach
Dietary Nitrogen Salivary urea
Tissue proteins Amino acid synthesis
Overfeeding of protein can lead to excessive levels of ammonia production, ammonia toxicity and energy expenditure for conversion (detoxification) of ammonia to urea in the liver
GWR 227206 Integrative Veterinary Physiology 22
Rumen Gases
The production of gases by the microbes reaches a peak of up to 40 L/h in cattle 2-4 hours after a meal, when fermentation rate is at its maximum. The principal gases produced are CO2 (60%), CH4 (30-40%) and variable amounts of N2, H2S, H2 and 02. They are eliminated almost entirely by the process of eructation.
Composition of rumen gases in a dairy cow. From L.E. Washburn and S. Brody, Mo. Ag. Exp. Sta. Res. Bull.: 263, 1937.
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Control of Fermentation
Ruminants exert limited control over the rate at which fermentation proceeds by altering or satisfying several requirements. These include:

The regular supply of macerated substrate (chewed food) for microbial digestion. Removal of the end products of fermentation (VFAs, microbial products, gases). The movement of undigested particles to the abomasum, and thence to the small intestine. Mixing of the forestomach contents to prevent local accumulations of inhibitory end products, and aid absorption of VFAs. Providing a highly buffered fluid environment for the micro-organisms. Providing stable conditions of temperature, osmotic pressure, and pH.
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Salivary Secretion
Ruminants produce large volumes of saliva
Sheep: 6-16 L/day
cf blood volume ~ 3.5 litres cf blood volume ~ 40 litres
Cattle: 60-160 L/day
The secretions from the major glands are

isotonic with blood plasma have no significant amylase activity change their composition following salt depletion (K+ exchanged for Na+ due to the action of aldosterone), contain urea and alkali maintain a basal secretory rate even after denervation.
The main salivary glands of the sheep (from Kay, 1960). 1: parotid, 2: submaxillary (submandibular), 3: inferior molar, 4: sublingual, 5: buccal, 6: labial. Glands are paired on both sides of the head and mouth.
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Salivary Secretion
The reticulum, rumen and omasum produce no secretion of their own and depend for the provision of a liquid medium on:

water contained in food water in drink saliva
The parotid glands are serous glands

provide the major fraction of salivary secretion into the rumen. In sheep a single gland may secrete 1-4 L/day, In cattle, estimates for a single gland range from 20-80 L/day
Parotid saliva is approximately isotonic with tissue fluid
helps stabilise the osmotic environment for the microbes.
Has a high electrolyte content:

Electrolyte concentrations in ruminant parotid saliva
Na+ Saliva (mmol/l)

Plasma (mmol/l):
K+ 13
3.9-5.4
HCO3112
~27
HPO4-48
2-7
Cl11
95-105
170
140-180
The high levels of HCO3 and HPO4-- are important for their buffering action and maintaining a pH range of 5.5 - 7.0 despite VFA production. Urea in parotid saliva provides an energy source for the microbes
Control of Salivary Secretion
Salivary glands are innervated by parasympathetic and sympathetic divisions of ANS
Courtesy Dr J Patterson, Swinburne University, Melbourne
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As in other animals afferent stimuli arise in the mouth from taste and mechanical stimulation of the gums. In ruminants, important sensory inputs also arise from lower in the gut. Moderate stretch in several areas of the gut is an effective excitatory stimulus to parotid secretion.

Oesophagus - especially the thoracic part The opening of the oesophagus (the cardia) The reticulum- especially on its medial wall near the reticular groove and reticulo-omasal orifice Omasal canal
Vagal afferent fibres carry excitatory stimuli from these regions to salivary centres. Ensures a steady flow of saliva enters the rumen even when the animal is not eating or ruminating The stimuli mentioned so far all increase or excite parotid secretion
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The stimuli mentioned so far all stimulate parotid secretion Recordings in conscious animals suggest there are also inhibitory mechanisms operating When sheep and cattle eat
Initially there is a very high rate of secretion from the parotid glands After about 30 minutes of eating this declines and falls to below pre feeding levels This is despite the fact that feeding continues and excitatory stimuli from taste, buccal and oesophageal stimulation are present
Parotid and submandibular salivary secretion rates during eating. Submandibular flow is shown by the dotted line and parotid flow by the solid line. Feeding is marked by F and rumination by R. (From D.H. Carr, pers comm)
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Possible inhibitory stimuli:
High (excessive) levels of stretch of the oesophageal and gastric regions mentioned earlier will inhibit salivary secretion Mediated by vagal and splanchnic afferent nerves Important as digesta and secretions accumulate in the stomach Increases in blood osmotic pressure reduces parotid secretion. Plasma osmolality may increase by over 5% during feeding

Increased metabolite production in gut Movement of water into gut Drop in blood volume
Reduction in salivary flow helps maintaining stable conditions in the ECF during digestion.
Changes in the osmolality of jugular plasma, the rate of parotid salivary secretion and reticular contractions before and during feeding. (From D.H. Carr, pers comm).
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Sympathetic actions on the gland

Decrease the blood supply through vasoconstriction Cause contraction of myoepithelial cells within the gland and expulsion of saliva
Followed by compensatory pause in flow as myoepithelial cells relax
Add protein to the saliva
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Salivary Secretion
In summary, the principal functions of saliva in ruminants are to:

Return urea to the rumen for microbial protein synthesis. Add fluid for proper microbial actions in the large reticulorumen fermentation vat. Supply bicarbonate and phosphate buffers to keep the pH of the reticulorumen within the normal limits (5.5 to 7.0)
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Motility of Reticulum & Rumen
Several different methods can been used to study motility of the reticulum and rumen in intact animals

Palpation Auscultation Intragastric pressure changes recorded using balloons or open tipped catheters positioned in the stomach via

Nasogastric tube Rumen fistula

Barium sulphate (insoluble) Radio-opaque markers surgically attached to stomach wall
Radiography

Electromyography Partial Exteriorisations
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Partial Exteriorisations
Partial exteriorisations are prepared by bringing small parts of the reticulum and rumen into a position immediately beneath the skin. Contractions of the stomach are then seen as skin movements.
Method of partially exteriorising the rumen. The circles represent stitches. (From C.S.W. Reid, Proc. N.Z. Soc. Anim. Prod., 23: 169188.)
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Reticulorumen A sequence

Two basic types of contraction sequence The first commences with a double contraction of the reticulum Contraction then spreads from the cranial to the caudal regions of the rumen Involves dorsal rumen first and then ventral rumen This is a backward moving contraction.
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Reticulorumen A sequence
This type of contraction has been variously termed:

Primary contraction. Mixing cycle A sequence
It recurs in sheep at about 30-90 sec - at the shorter interval if the animal has been recently fed, and toward the longer if fasted. The functions of the contraction include:

Mixing of digesta and distribution of microbes Mechanical breakdown of digesta Bringing the products of fermentation to absorptive surfaces Aiding the movement of digesta onward in the gut
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Reticulorumen B sequence

The second type of contraction does not involve the reticulum Starts as a contraction of caudal ventral blind sac Spreads to posterior then anterior regions of dorsal rumen Ends with a contraction of ventral rumen sac This is a forward moving contraction.
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Reticulorumen B sequence
This contraction has been variously termed:

Secondary contraction. Eructation (belching) cycle. B sequence.
B sequences follow a variable time after A sequences - not always in a 1:1 ratio. Animal will eructate (belch) with this contraction, depending on the rate of gas production.
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Control of RR Motility
Organised movements of the reticulum and rumen cease after

the vagus nerves have been cut after atropine has been administered
The evidence from anaesthetised animals indicates that both sensory and motor fibres controlling RR contractions are contained in the vagus nerves In anaesthetised animals reticulum contractions can be caused by

Stimulation of the intact vagus nerve. Stimulation of the peripheral end of the cut cervical vagus nerve. Stimulation of the central end of a cut cervical vagus nerve, provided the other vagus nerve is intact.
The vagi contain both motor (efferent) fibres to the stomach and sensory (afferent) fibres from it, both of which are normally involved in the regulation of gastric contractions
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Mechanical stimulation (touch or stretch) of the regions of the gut listed below is an effective stimulus to contractions of the reticulum and rumen.

The mouth (gums) Thoracic oesophagus Reticulum (especially medial wall) Reticulorumenal fold Reticulo-omasal orifice Slight stretch of the abomasum
All these are areas that would normally be stimulated during eating and digestion
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Vagus Indigestion
Vagus indigestion (abnormal motility patterns of RR) is due to damage to receptor areas rather than to vagi themselves - especially damage around medial wall of reticulum
Metal door spring removed from a cows reticulum
A nail has penetrated the reticulum, causing traumatic reticuloperitonitis (hardware disease) and the death of this cow
A nail is embedded in the epithelium of the reticulum
Images from Noahs Arkive, University of Georgia. GWR 227206 Integrative Veterinary Physiology
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Chemical stimuli delivered to the gut may also be involved in reflex stimulation of RR contractions. The addition of HCl at pH 1to the abomasum stimulates reticular contractions, provided the vagal branches to the abomasum remained intact. This observation has been criticised because abomasal pH rarely falls below pH 2.5. However acid is secreted by parietal cells at about pH 0.9 Provided the receptors are close to the site of acid secretion, stimulation would be possible.
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Co-ordination of the afferent information from chemo- and mechano-receptors occurs in reflex centres. These are located in the dorsal vagal nucleus of the medulla oblongata They regulate the efferent vagal discharge. Inhibitory as well as excitatory stimuli influence the activity of the medullary centres. Excess stretch is inhibitory.
Generally this applies to any of the areas where moderate stretch is excitatory, but especially to the abomasum.
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Detecting Contractions of Reticulum and Rumen in the field
Reticular contractions identified by a tinkling sound heard through stethoscope
Caused by rumen fluid flowing back into reticulum as it relaxes
Contraction of rumen detected by palpation in sub lumbar fossa region
A hardening and bulging caused by contraction of dorsal rumen
This allows A sequences and B sequences to be distinguished seperately

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New-Born Ruminant Stomach

Rumen
Weight as a percentage of birth weight (birth weight = 100) Dorsal sac Rumen Posterior Blind sacs Reticulum
Oesophagus
Ventral sac Omasum
Pylorus
Omasum
Abomasum
Small intestine, caecum & colon Abomasun
Age in weeks
The forestomach in the new-born ruminant is anatomically and functionally under-developed It contains no microbes (i.e. is sterile) and does not contribute significantly to the digestion milk
Rumen Development
Oesophagus Oesophagus Rumen Reticulum Abomasum Omasum Abomasum Rumen Omasum Reticulum
Birth to 2 weeks
8 weeks
Oesophagus Rumen Rumen Reticulum Abomasum Omasum Abomasum Omasum
Oesophagus
Reticulum
3-4 months
Mature
Rob Costello, Dairy Technical Specialist, Merrick's Inc.
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In the new-born ruminant, swallowed milk is directed from the oesophagus directly into the omasum and abomasum by the reticular groove. When contracted forms a conduit between the cardia and the reticulo-omasal orifice
From: D.A. Titchen and J.C. Newhook, 1975. In Digestion and Metabolism in the Ruminant. pp 15-29. Editors IW. McDonald and A.C.I. Warner. Australia, University of New England Publishing Unit.
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Within the abomasum, the newly ingested colostrum and milk is subjected to the clotting action of the enzyme rennin. Rennin has a pH optimum of near 6.5 and produces a hard clot or curd consisting of the butterfat and the curd proteins (caseinogens) precipitated as calcium caseinate) The whey fraction, which contains important immunoglobulins, passes into the small intestine The antibodies are absorbed, passively immunising the young animal In cattle the immunoglobulins constitutes about 70% of the whey protein of colostrum The immunoglobulins survive digestion in the gut because:

Neonates are achlorhydric, i.e acid, and pepsin, secretion is absent or low Colostrum contains an inhibitor to the proteolytic enzyme trypsin.
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Transmission of Passive immunity and time after birth for gut closure.
Species Ox, Goat, Sheep Pig Horse Dog Rat Rabbit Man
Prenatal 0 0 0 + + +++ +++
Postnatal +++ (36 hr) +++ (36 hr) +++ (36 hr) ++ (l0 days) ++ (20 days) 0 0
The immunoglobulins are absorbed by pinocytosis They coalesce to form a single large globule in the basal portion in the cell There are both specific, receptor mediated absorption mechanisms, and non-specific absorption mechanisms
egg albumin, gelatin and serum protein added to the colostrum is also absorbed
Adrenocortical hormones appear to cause premature closure of the gut Such hormones are produced under conditions of stress Lambs born under stressful conditions (climatic or nutritional) may have impaired globulin absorption.
Reticular Groove

Milk taken when young ruminants suck passes directly to the abomasum The milk flows directly to the abomasum where the enzyme rennin causes coagulation This delays its passage into the intestine Other swallowed fluids, e.g. saliva, are directed into the reticulum The reticular groove forms a channel from the oesophagus, through the reticulum to the reticuloomasal orifice. It has two well defined lips which flank a floor. The lips make a functional extension of the oesophagus directly to the reticulo-omasal orifice.
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Reticular Groove Reflex Control
Contraction of the reticular groove occurs as a reflex response. When the vagi are cut, suckled milk enters the reticulorumen.
In decerebrate preparations the reticular groove can be made to contract by
introduction of water into the posterior region of the mouth cavity touching the posterior mouth cavity with a probe Cranial laryngeal nerve stimulation (sensory)
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Reticular Groove Reflex Control
The responses of the reticular groove illustrate many of the classical properties of visceral reflexes. Latency:
2 -4 seconds may elapse between delivery of stimulus and contraction of the groove. .
Summation:

One afferent stimulus alone is often insufficient to excite a visceral reflex. Introduction of water alone into the mouth, or stimulation of afferent fibres in the cranial laryngeal nerve, is ineffective.
Inhibition:

Groove closure inhibited when the abomasum is full/stretched (vago-vagal reflex). Causes spillage of milk into the reticulum and rumen.
The rapid ingestion of a large volume of milk (e.g. once a day feeding) may cause ineffective clotting of the curd and rapid emptying of the abomasum. This in turn results in overloading of the small intestine with protein, bacterial overgrowth in the intestine and diarrhoea (scours).
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Reticular Groove Adult Animals
In about 70% of adult sheep the reticular groove can be made to contract by giving copper salts orally This has been used to direct drenches against nematodes directly to the abomasum Nicotine and arsenic were the active components of the drenches To achieve a lethal concentration for the worms (not the sheep) they had to be delivered to the site of parasite action (abomasum) In adult cattle sodium salts have the effect of causing groove contraction From time to time there is a resurgence of interest in using the reticular groove mechanism to aid animal production
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Reticulo-Omasal Orifice (ROO)
Passage of material from the reticulum to the omasum occurs via a relaxed ROO, which in sheep is about 1 cm in diameter. In the cow, for the greater part of the A sequence, the ROO is loosely open (some 60-70% of the time). During the first phase of the diphasic reticular contraction the orifice closes, it opens at the peak of the second phase of contraction of the reticulum Relaxation (opening) of the ROO is controlled by the vagus The neurotransmitter is most likely VIP
Motility of the Reticulo-Omasal Orifice R.O.O.
Reticulum
R.O.O.
Pressure recordings (balloon)
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Reticulo-omasal Orifice (ROO)
When the ROO opens, digesta flow into the omasum and the omasum relaxes Relaxation of the omasum combined with contraction of the reticulum contributes to the flow of digesta into the omasum Contractions of the omasum can be inhibited by distension of the abomasum This may provide a means for the abomasum to control the volume of ingesta entering it
Pressure records
Reticulum
R.O.O.
Omasum
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Omasum

The function of the omasum is unclear. May help regulate flow of digesta from the reticulorumen to the abomasum Fluid and electrolyte and VFAs are absorbed by the omasum. The surface area of the leaves is large - about 1/3 of the total epithelial lining of the reticulorumen The leaves undergo movement, especially near their bases. Contractions stimulated by infusions of VFAs into the omasal lumen. May not be essential for life Some ruminants, e.g. primitive deer, (lesser mouse deer) have no omasum. This is a browsing type of ruminant
Omasal folds
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Eructation

Removal of gas produced by micro-organisms during fermentative digestion Gas produced in the first 3 divisions of the ruminant stomach Rate of production increases after eating and decreases slowly until the next feeding. Most of the gas is produced in the reticulorumen Estimated that 1.2 - 2 litres of gas are formed per minute in the rumen of a 500 kg cow. Most of the gas is the result of bacterial action and CO2 liberation from salivary bicarbonate
CO2 65%
CH4 27%
N2 7%
O2
H2
H 2S
NH3 Trace
0.60% 0.20% Trace
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Eructation
Dorsal Rumen
Eructation usually begins with a B sequence contraction. Oesophagus This sweeps gas toward the oesophagus at the cardia. Whether eructation occurs at this stage depends on clearing the cardia of ingesta The sphincter at the diaphragm Reticulum relaxes, and the oesophagus fills with gas
Caudal Ventral Blind Sac Ventral Rumen
When the oesophagus has filled with gas the diaphragmatic sphincter closes and the pharyngo-oesophageal sphincter relaxes Gas moves up the oesophagus - in cattle aided by a rapid anti-peristaltic wave of contraction in the oesophagus - perhaps passively in sheep
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From: Van Soest, Peter J (1994) Nutritional Ecology of the Ruminant, Edition: 2nd. Publisher: Comstock Publishing
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Ruminant Methane Production
Since 1999 atmospheric methane concentrations have levelled off World population of ruminants has increased at an accelerated rate Prior to 1999 there was a strong relationship between change in atmospheric methane concentrations and the world ruminant populations Since 1999 this strong relation has disappeared. This change in relationship suggests that the role of ruminants in greenhouse gases may be less significant than originally thought
Global large ruminant equivalence and atmospheric methane concentrations

1850 1800 1750 1700 1650 1600 1550 1500
Large ruminant equivalence (*1,000,000) Methane concentration (ppb)
1979 1981 1983 1985 1987 1989 1991 1993 1995 1997 1999 2001 2005 Year
Figure 1. Global atmospheric methane concentrations from NOAA (2007) and cattle equivalents from FAO (2007). Large ruminant equivalences are calculated using 8 sheep or goats as being equivalent to a large animal FAO: Food and Agriculture Organisation IAEA: International Atomic Energy Agency NOAA: National Oceanic and Atmospheric Administration
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Ruminant Methane Production

R. A. Leng Department of Biochemistry, Microbiology and Nutrition, University of New England, Armidale, N.S.W. 2351
Methane production (kg/ton Lwt. Gain)
Large ruminants produce some 1520% of the global production of methane Ruminants on low quality feeds possibly produce over 75% of the methane from the world's population of ruminants Supplementation can reduce methane production as

(A)
% digestible energy Fermented to methane
20
No supplements
(B) 1200
No supplements
15
Urea/mineral supplements
800
10
400
Urea/mineral + bypass protein supplements
% digestible energy consumed per kg live-weight gain
The effects of supplements on methane emissions for ruminants on low feeds. (A) The effects of improving the efficiency of rumen fermentative activity on methane production/kg of digestible energy consumed. (B) the production of methane/kg gain in supplemented cattle (feed conversion efficiency (FCR) (9:1) or un supplemented cattle (FCR = 40:1) fed straw-based diets (after Saadullah 1984)
FCR is the mass of the food eaten divided by the body mass gain over a specified time
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Control of Eructation

Gaseous distension of the rumen stimulates eructation and secondary contractions of the rumen Occurs only if the animal is in an upright or nearly upright position Does not occur under general anaesthesia Caused by stimulation of receptors in the caudal region of the dorsal sac of the rumen, Is a vago-vagal reflex. Receptors in the cranial part of the rumen and reticulum are important in determining whether or not eructation occurs with the secondary contractions Eructation does not occur when the area around the cardia is covered with ingesta, water, foam This is important because any liquid that was regurgitated into the mouth with the eructated gas can enter the airways and lungs (glottis open during eructation).
Cardiac Opening Fluid Distension Distension Secondary cycles
No cardiac opening
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Eructation
Other stimuli, apart from gaseous distension of the rumen, are important in causing eructation Feedlot cattle in USA are often subject to a chronic and appetitedepressing bloat when fed a concentrate food. This can be corrected by ensuring the incorporation of some scabrous material (roughage) in the diet. Apparently tactile stimulation of the RR is important
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Rumination
Rumination is the process by which ingesta are regurgitated, rechewed, reensalivated and reswallowed. This second period of mastication appears to have two main functions:

Cut transversely Unaltered after 40 hrs
To reduce food particles size. To facilitate microbial attack on the plant material
Comparisons have been made of the rate of digestion of lucerne stalks placed in nylon bags in the rumens of sheep. The lucerne was cut either transversely or longitudinally, placed in a nylon bag and then examined at intervals with scanning EM.
Cut longitudinally Unrecognisable After 6-7 hrs
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Rumination
Rumination has been shown to be very effective in reducing particle size of food Observations were made on sheep with oesophageal fistulae Boluses were collected on their way to or on their way from the mouth (after re-chewing)
Particle size Coarse Medium
To mouth 30% 37%
From mouth 10% 51%
Fine
33%
39%
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Rumination

Involves both the digestive and respiratory systems. Rumination is characterised by:

An extra reticular contraction before the normal diphasic contraction Respiration momentarily stops and an inspiratory effort made against a glottis at the peak of the extra phase of reticular contraction The bolus moves rapidly up the oesophagus to the mouth. Excess fluid from the bolus is swallowed Bolus is remasticated
The process is completed in 45-60 seconds by re-swallowing the bolus. The whole sequence is repeated after the previous bolus is reswallowing. Rumination commonly continues for an hour or more.
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Electromyogram (EMG) Recordings During Rumination in a Sheep

A. EMG of reticulum showing normal biphasic contraction B. A normal biphasic contraction preceded by an extra contraction C. EMGs of the oesophagus near the glottis (1), upper thorax (2) and close to the cardia (3), and the reticulum (Re) D. Jaw and respiratory movements The regurgitation phase (AP) of rumination is closely followed by swallowing of the excess liquid on two occasions (P1 and P2) and later of the bolus (P3).
A.
B.
C.
D.
68
Rumination
69
Rumination
Periods of rumination and feeding in sheep and cattle on legume pasture Although diurnal patterns differ, both species tend to graze in the morning and evening and rest during the hotter part of the day
70
Rumination

The physical nature of the diet influences the age at which rumination commences It may begin in lambs and calves as soon as two weeks after birth A diet containing roughage causes rumination to start earlier than a diet of solely milk.
The drive for roughage is strong - milk fed calves will eat the wooden rails of pens and lambs pull wool from other lambs and chew and swallow it
Tactile stimulation (touch) or light stretch particularly of the area around the cardia, reticular groove, reticulo-ruminal fold and reticulo-omasal orifice is very effective. Thus the desire to ruminate seems to be related in large part to the volume of contents in the reticulorumen, and to the tactile stimulation by coarse material in the rumen Rumination is associated with characteristic changes in the demeanour of animals and often occurs at night. A reduction in external stimuli (visual and auditory) may be a pre-requisite.
71
Rumination
Current theories on the basis of rumination suggest that the initial reticular contraction at the time of regurgitation has two functions:
To flood the area of the cardia with ingesta which is then available for regurgitation. To raise the pressure in the stomach - there may be an increase of 5-6 mm Hg during this contraction. The inspiratory effort against a closed glottis produces a large reduction in pressure within the thorax, and hence within the thoracic oesophagus. The steep gastric-oesophageal pressure gradient - it may be 40-60 mm Hg in cattle drives the movement of digesta into the oesophagus when it relaxes. The bolus is then swept to the mouth by a reverse peristaltic wave.
72
Abomasum

The abomasum is structurally and histologically comparable to the simple stomach of monogastric animals. There are three regions which are defined by the type of gland:

A very small cardiac region encircling the omasoabomasal orifice which has cardiac glands The body or fundus which contains gastric or fundic glands The pyloric region which contains pyloric or antral glands.
The body has large spiral mucosal folds and this results in about 90% of the total surface area being formed by the body region. The pyloric region consists of the antrum, which has no folds but only surface rugae, the pyloric canal and sphincter.
Pyloric sphincter
Omasoabomasal orifice
73
Abomasum Gastric Glands
In the fundic region, the glands contain
parietal (or oxyntic) cells which produce HCl chief (or peptic) cells which produce pepsinogen mucous neck cells
The chief cells are present in the lower third of the gland The parietal cells occupy much of the middle third Endocrine cells are found mainly in the lower part of the gland The gastric pits are lined with surface epithelial cells.
74
Abomasum
The cardiac glands contain the same cell types as the fundic glands. There are is larger number of mucous cells which produce a very viscous mucus. The pyloric (antral) glands are coiled and produce both mucus and pepsinogen, There are virtually no parietal cells. The gastrin-secreting G cells are present in antral glands.
75
Abomasum
The composition of gastric juice in the ruminant is similar to that in other mammals pH averages about pH 2.8 contains pepsin and intrinsic factor The principal functions of the abomasum are:
Digestion of protein through action of pepsin Rumen microorganisms entering the abomasum are killed by the acid and digested to provide nutrients for the host Secretes intrinsic factor which is essential for absorption of Vitamin B12 in the ileum In the immature ruminant the abomasum secretes the enzyme rennin which causes rapid clotting of milk. Pepsin is also produced in the preruminant animal and is involved in proteolytic digestion of the milk
76
Abomasal Secretions
Both abomasal secretion and the inflow of digesta from the omasum are continuous. Sheep secrete about 5 litres of gastric juice per day. The volume, acidity and pepsin content increase with feeding and rumination, decreases with starvation Experimental sheep may also develop a conditioned secretory response to the sight and smell of food. Abomasum secretions
stimulated by vagal nerve stimulation, cholinomimetics Inhibited by atropine i.e. parasympathetic blockers
77
Abomasal Secretions
Control of abomasal secretions can be studied using surgically prepared pouches
Secretion from pouch increases when animal feeds
78
Abomasal Secretions
The increase in abomasal secretion with feeding is associated with an increased inflow of digesta into the abomasum. Part of the response due to distension of the stomach which is one of the known stimuli to abomasal secretion in ruminants
79
Abomasal Secretions

Distension also stimulates the release of the antral hormone gastrin. Gastrin secretion is important in ruminants (as well as monogastrics) in mediating food-stimulated gastric secretion. The composition of the digesta coming into the abomasaum appears to be a key factor. Gastrin is released by ammonia and peptone in the abomasal fluid and when the pH of the abomasal contents rises. The high pH of incoming digesta appears to be a powerful stimulant to gastrin release in the sheep
80
Abomasal Parasites
Abomasal nematodes

Ostertagia circumcincta (sheep) 0stertagia ostertagi (cattle) Trichostrongylus axei Heamonchus contortus
All impair the secretory activity of the abomasum. The abomasal pH rises since acid secretion is inhibited, serum gastrin increases and there may also be raised serum pepsinogen levels. Diarrhoea, reduced digestibility and disturbed protein metabolism and utilization The high serum gastrin may reduce both rumino reticular and abomasal motility and also the animals appetite. H. contortus is a blood-sucker and can cause serious anaemia and death.

Ruminant Digestion

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Integrative Veterinary Physiology Ruminant Digestion

GWR 227206 Integrative Veterinary Physiology

GWR 227206 Integrative Veterinary Physiology

GWR 227206 Integrative Veterinary Physiology

The microbes synthesis relatively high quality proteins from

GWR 227206 Integrative Veterinary Physiology

Ruminants must spend

GWR 227206 Integrative Veterinary Physiology

acetic, propionic and butyric acids,

GWR 227206 Integrative Veterinary Physiology

GWR 227206 Integrative Veterinary Physiology

GWR 227206 Integrative Veterinary Physiology

Ruminant Stomach Right Side

GWR 227206 Integrative Veterinary Physiology

Ruminant Stomach left Side

Caudal Blind sac

abomasum reticulum Ventral rumen

GWR 227206 Integrative Veterinary Physiology

Innervation of Ruminant Stomach

The innervation to the stomach

GWR 227206 Integrative Veterinary Physiology

Innervation of Ruminant Stomach

to liver & pylorus Rumen Dorsal vagus

Ventral vagus Omasum Abomasum Reticulum

GWR 227206 Integrative Veterinary Physiology

Innervation of Ruminant Stomach

slowly adapting mechanoreceptors located in the muscle layers activated by

Blood Supply to Ruminant Stomach

GWR 227206 Integrative Veterinary Physiology

Blood Supply to Ruminant Stomach

GWR 227206 Integrative Veterinary Physiology

GWR 227206 Integrative Veterinary Physiology

Slowly broken down by the ruminal cellulolytic bacteria to VFAs.

Carbohydrate in grain-based feed contains linked glucose units

rapidly degraded by amylolytic bacteria to VFAs and lactic acid.

Quickly used up by facultative anaerobic bacteria

GWR 227206 Integrative Veterinary Physiology

GWR 227206 Integrative Veterinary Physiology

Digestion in the Rumen

Animal Sheep Ox Horse Dog

Organ Rumen Rumen Caecum Colon

GWR 227206 Integrative Veterinary Physiology

Digestion in the Rumen

GWR 227206 Integrative Veterinary Physiology

Dietary Nitrogen Salivary urea

Tissue proteins Amino acid synthesis

GWR 227206 Integrative Veterinary Physiology

GWR 227206 Integrative Veterinary Physiology

Ruminants produce large volumes of saliva

Sheep: 6-16 L/day

cf blood volume ~ 3.5 litres cf blood volume ~ 40 litres

Cattle: 60-160 L/day

The secretions from the major glands are

GWR 227206 Integrative Veterinary Physiology

water contained in food water in drink saliva

The parotid glands are serous glands

Parotid saliva is approximately isotonic with tissue fluid

helps stabilise the osmotic environment for the microbes.

Has a high electrolyte content:

Na+ Saliva (mmol/l)

Control of Salivary Secretion

Salivary glands are innervated by parasympathetic and sympathetic divisions of ANS