Rumen Motility

Church: 67-106, 108-116, 468-474 Van Soest: 237-244 Ruckebusch and Thivend: 35-51; 103-119 Sjersen et al.: 155-164 Susenbeth et al. 1998. Energy Requirement for Eating in Cattle. J. Anim. Sci. 76:2701-2705. http://jas.fass.org/cgi/reprint/76/10/2701.pdf Mawuenyegah et al. 1997. Effect of Ammonia Treatment or Protein Supplementation on Rumination Behavior in Sheep Given Barley Straw. Animal Science 64(3):441-445. http://www.bsas.org.uk/Publications/Animal_Science_PDF_Back_Issues/ Animal_Science_PDF_Back_Issues/ Cheng et al. 1998. A Review of Bloat in Feedlot Cattle. J. Anim. Sci. 76:299-308. http://jas.fass.org/cgi/reprint/76/1/299.pdf Majak et al. 1995. Pasture Management Strategies for Reducing the Risk of Legume Bloat in Cattle. J. Anim. Sci. 73:1493-1498. http://jas.fass.org/cgi/reprint/73/5/1493.pdf

Rumen contractions
Functions
Inoculate incoming feed Mix contents
Minimize effects of stratification Move fermentation products to rumen wall

Particle sorting Particle passage Rumination Eructation of fermentation gases

Location
Contractions are contractions of an entire sac of the reticulorumen, but are the result of muscular contraction primarily on the pillars

Types of reticular contractions

Biphasic contraction
Action
Reticulum contracts to about its size, relaxes, and contracts completely Reticulo-omasal orifice is open during contraction, but closes when reticulum dilates
Assists in digesta passage from the reticulorumen Passage more closely related to duration (r2 = .76) than amplitude (r2 = .56) or frequency (r2 = .15)

Duration
7 to 12 seconds

Frequency
Eating 35 to 45 seconds Resting 75 seconds

Triphasic contraction
Action
An extra complete contraction precedes a biphasic contraction of the reticulum Contraction is associated with the movement of digesta up to the cardiac sphincter before rumination.

Types of ruminal contractions

Primary contraction
Also called A-wave or backward moving Action
A biphasic contraction of the reticulum Anterior pillar contracts lifting the anterior sac.
Anterior fold rises to form barrier

Contraction moves across dorsal sac to dorsal blind sac from contraction of longitudinal pillar and dorsal coronary pillar Dorsal blind sac contracts and dorsal sac relaxes from dorsal coronary pillar Ventral sac contracts along longitudinal pillar Ventral sac relaxed and Ventral blind sac contracts
Ventral contraction absent during rumination

Functions of the primary contraction

Mixing and inoculation of digesta Particle sorting across the reticuloruminal and anterior folds

Duration of contraction
Fed animal 30 to 50 seconds Fasted animal 12 to 18 seconds

The primary contraction of the rumen

 Secondary contraction
Also called the B-wave or forward-moving contraction Usually occurs after a primary contraction Action
Contraction of the ventral blind sac continuing up through the dorsal blind sac using the dorsal coronary pillar Contraction proceeds across dorsal sac forcing gas pocket to the cardiac sphincter

Function
Eructation

Duration
30 seconds

Incidence of pressure waves

Activity Feeding Resting Ruminating D 1 10 22 Contraction DV DSV 27 5 35 25 28 37 DVSV 56 22 6

Reticulum

Cranial sac

Neural control of reticuloruminal contractions

Nerves
Vagus nerve involved both in stimuli and inhibition Sphlanchic nerve involved in inhibition

Stimuli for contractions

Stretch
Most common Receptors
Low threshold receptors Tension receptors Stimulated by pressures greater than 4 mm Hg Stimulate contractions High threshold receptors Epithelial receptors Stimulated by pressures greater than 20 mm Hg Inhibit contractions

Tactile stimulation
Near cardiac sphincter Epithelial receptors
Stimulates triphasic contraction of reticulum

Low abomasal pH
Stimulates ruminal contractions

Hypoglycemia

 Inhibition of ruminal contractions

Abomasal distension Hyperglycemia

Factors affecting ruminal contractions

Feeding
Increases frequency and amplitude of contractions Activity Contractions/min Amplitude P/S mm Hg Resting 1.2 18.2 Feeding 2.0 22.1 1:1 Ruminating 1.1 10.4 Fasting .3 Weak 5:1

Diet
Feeding a finely ground forage
Reduced rate of contractions Reduced amplitude of contractions
Requires 2-6 weeks to adapt

Volatile fatty acids

Acetic, propionic, and butyric acids at 90, 50, and 37 mM given separately or at 33.5, 29.3, and 21.1 mM given as a mixture will inhibit primary contraction

Metabolic problems
Hardware disease, hypocalcemia, or hyperglycemia will inhibit ruminal contractions

Need for eructation

Gas production
Peak
Occurs 30 min to 2 hr post-feeding 12-27 l/min

Average
1-2 l/min

Approximately to 1/3 of CO2 produced in rumen is absorbed into blood and removed through the lungs Only 1/5 of the CH4 is removed through the lungs

Composition of rumen gas __Gas__ _%__ CO2 65.35 CH4 (variable) 27.76 N2 7.00 O2 (at wall) .56 H2 .18 H2S .01

Mechanism of eructation
Biphasic contraction of reticulum Modified primary contraction of rumen Secondary contraction of rumen Ruminal gas pocket forced forward Anterior fold and reticuloruminal fold rise to hold digesta away from cardiac sphincter Cardiac sphincter relaxes allowing esophagus to fill with gas Rapid reverse peristallsis of the esophagus
Diaphragmatic and Pharyngeal sphincters open Nasopharyngeal sphincter closes

Epiglottis opens while thoracic muscles contract

Cause 80% of gas to enter trachea Acts as a muffler

Animal exhales

Control of eructation
Stimulus
Gaseous distension of the reticulum and rumen

Inhibition
Presence of digesta near the cardiac sphincter
Affects all three sphincters Prevents digesta from entering lungs

Epinephrine Histamine

Inhibition of eructation will cause the animals to bloat

Ruminal pressures will increase to 45 to 100 mm Hg.

Bloat
Types of bloat
Free gas bloat
Gas pocket over the digesta is normal, but cant be eructated because of a physical obstruction in the esophagus or anatomical abnormality

Frothy bloat
Gases form a foam over the digesta that inhibits eructation when it touches the reticular wall near the cardiac spincter Two types
Feedlot bloat Pasture or legume bloat

Feedlot bloat
Etiology
Occurs in ruminant animals fed high grain diets particularly during adaptation to the diets

Causes
Digesta becomes viscous and gel-like that trap gas as a foam
Viscosity increased by
The presence of bacterial capsules on some gram + starch-digesting bacteria such as Streptococcus bovis and lactobacilli Streptococcus bovis and lactobacilli only found in large numbers in animals not adapted to high grain diets The presence of short-chain starches (dextrans)

Increased concentrations of VFAs

Reduce ruminal motility

Increased intake of a dense diet

Lowers cardiac sphincter

Increased numbers of mucinolytic bacteria

Mucin will prevent foam

Reduced outflow rate

 Prevention
Slowly adapt animals to high grain diets Feed adequate fiber Use less fermentable grains
Wheat or barley > Corn or sorghum Steam-flaked > Coarse ground

Feed ionophores (Monensin or Salinomycin)

Decreased growth of Streptococcus bovis and lactobacilli Decreased feed intake Decreased methane production

Pasture bloat
Occurs in ruminant animal consuming fresh legumes such as alfalfa, white clover and red clover or the small grain wheat when immature or immediately after a frost Causes
Presence of high concentrations of soluble protein present in chloroplasts increases surface tension causing foam
Earlier research implied that a protein named ribulose-1,5biphosphate carboxylase oxygenase (also called the 18S protein)
3 6% of DM in bloat-causing legumes

Recent research shows that total soluble protein concentration is more highly related to bloat

Rapid cell wall digestion

Rapid release of chloroplasts Rapid release of cell wall fragments that get trapped in foam

Pectin
A cell wall component in high concentration in legume cell walls Pectin metabolism
Pectin fermented to Pectic acid (binds 2000x its weight in water increasing viscosity) Pectic acid rapidly fermented to methane

 Minerals
K, Ca, Ni, Zn, and Mg stabilize foam Frosted legumes
K is released into cytoplasm Decreased Na:K ratio results in increased binding between proteins Increases surface tension causing bloat

Decreased chewing
Decreased salivary mucin and Na Mucin prevents foaming

Reduced ruminal outflow

Retention time of CoEDTA, hr Nonsusceptible 8 Susceptible 12-17 Increases concentrations of chloroplasts and chlorophyll

Saponin
Surfactant glucoside Minor role in stabilizing foam

pH less than 6
Stabilizes foam

Toxic factors
Amines like histamine reduce motility

Legume bloat prevention

Mix grasses with legumes in pastures Allow legumes to mature before grazing Use of non-bloating legume species
Birdsfoot trefoil, sanfoin, crownvetch, berseem clover, and some cultivars of red clover Produce high concentrations of tannins
Polymers of phenolic compounds Actions
Binds proteins preventing release Reduce rate of digestion

Feed some dry hay before putting animals on pastures Move to new paddocks after early morning
Not related to dew Appears related to selective retention of chloroplasts

Increase grazing pressure

Forces animals to increase consumption of stems to leaves

Close availability of water

 Use of surfactants
Vegetable oil Poloxalene (Bloatguard)
Effective at 10 gm/1000 lb/day May be used as drench, block or loose supplement Questionable palatability

Antibiotics
Penicillin and tylosin
Prevents bloat for a short period Bacteria adapt causing bloat

Ionophores
Decreased incidence by 50%, but didnt prevent it

Genetic selection
After 10 years, bloat removed from herd Phenotype difference
Increased rumen liquid volume

Rumination
Functions
Rechewing to increase particle size Saliva flow

Particle breakdown
Chewing during eating
Functions
Preparation for swallowing Release soluble constituents Damage plant tissues for microbial attachment

Chewing during rumination

Functions
Decrease particle size for passage Damage plant tissues for microbial attachment

Microbial digestion Reticuloruminal contractions

Particle size reduction

Alfalfa Meadow Ryegrass Red clover Alfalfa hay hay ( % of intake) DM digested in RR 59.9 61.7 55.3 37.1 49.3 Soluble DM released 36.8 37.6 31.9 22.9 20.4 by chewing during eating Large particles reduced to < 1 mm by: Eating 11.8 14.0 14.5 14.3 14.4 Ruminating 12.0 13.7 16.4 26.6 38.5 (% of particle reduction) Particle reduction by 50.4 49.4 53.1 65.0 72.8 rumination

Mechanism of rumination
Triphasic contraction of the reticulum
Forces digesta to cardia

Animal inhales with epiglottis closed

Produces a vacuum of 60 to 80 mm water in esophagus

Cardia opens and esophagus dilates

Negative pressure sucks digesta into esophagus

Rapid reverse peristalsis moves digesta to mouth Bolus is rechewed

Chewing is slower and more deliberate than during eating

Digesta reinsalivated
Saliva composition is different than during eating
Parotid glands secrete more saliva during rumination than eating
Saliva from parotid glands secrete more HCO3- than other glands

Reswallowing

After reswallowing, the rumen undergoes a primary contraction to move it back in the rumen 20 to 65% of the DM is released in a fine state and will not return to the mouth again

Control of rumination
Controlled by tactile receptors (epithelial receptors) near the cardiac sphincter, reticuloruminal fold and anterior sac Stimulated by scratching of feed against the rumen wall Reflex is semiautomatic
Can be stopped anytime

EEG resembles sleeping

Rumination time Average times for a grazing animal

Eating 8 hours Ruminating 8 hours Resting 8 hours

Ruminating time is quite variable Factors affecting rumination time

Fiber content of diet Physical form of diet

Effects of diet on rumination

Increasing the proportion of grain in the diet will decrease rumination
Hay Dried grass Concentrate Chews/day 50,100 36,100 11,000

Increasing the maturity of the forage in a diet will increase rumination Decreasing the particle size of the diet will decrease the rumination time Increasing feed intake will reduce the rumination time per gm of feed consumed Cattle ruminate less per kg NDF than sheep or goats

Roughage value index

Total chewing, min/kg DM Alfalfa hay, Chopped Long Pelleted Oat straw Cottonseed hulls Alfalfa silage Fine chop Medium chop Corn-ground Soybean meal Minerals Molasses Urea 44.3 61.5 36.9 160 30.1 22.3 26.0 5.1 6.0 0 0 0

Effects of rumination of the animal

Saliva flow
More saliva secreted during rumination than eating Affects
Rumen pH
Fiber digestion Microbial growth VFA and methane production Maintenance of intake Structure of rumen epithelium Prevention of liver abscess and laminitis

Liquid rate of passage

Efficiency of microbial growth within the rumen

DM intake
Ruminating time is a limiting factor controlling intake of high fiber diets Maximum time, hours Ruminating 10-11 Grazing 13 Lower rumination efficiency (min/g CWC) limits intake of smaller and growing animals

Body weight Animal kg kg.75 Lambs 40 16.0 Goats 39 15.6 Mature sheep 82 27.3 Calves 119 35.9 Heifers 1 213 55.7 Heifers 2 342 79.6 Heifers 3 456 98.6 Mature cows 561 115.0

Rumination min/g CWC 2.05 1.30 1.18 .78 .42 .19 .16 .10

Max. CWC intake/d g/kg BW g/kg.75 BW 5.8 14.6 9.4 23.7 4.9 14.9 5.2 17.2 5.4 20.8 7.4 31.8 6.5 29.9 8.2 40.1

 Energy cost
Total chewing
increases maintenance cost by .24 kcal/hr/kg BW and accounts for 30% of maintenance requirement Accounts for difference in maintenance energy requirements of cattle fed high grain or high forage diets.

Contribution of rumination to energy cost of total chewing varies Time, min/kg DM consumed Feedstuff Eating Ruminating Oat straw 41-58 94-133 Medium quality hay 20-40 63-87 Good quality hay 27-31 55-74 Concentrates, pelleted 4-10 0-25 Finely ground oat straw 11-24 0-20 Finely ground hay 13 0-6