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RumenFermentationRumenFermentationRumenFermentationWorld’slargestcommercialfermentationspace100billionlitersorrumenvolumeindomesticanimals1010to1012cells/mLRumencapacityrangesfromlessthan1liter(1quart)inaduikerto200liters(50gallons)inacowRumenFermentationWorld’slargRuminantsContinuousculturefermentersInputandoutputLignocellulosicsubstrates(forages)digestedCellulasecomplexHemicellulasesNitrogencapture(NPN)8x1015mouthstofeedBecauseofthesemicrobialenzymes,ruminantscanutilizefeedstuffsthatprovidelittletononutritionalbenefittonon-ruminantsRuminantsContinuousculturefeFourStepsofRuminationRegurgitationReverseperistalsiscarriesfoodtomouthRemasticationLiquidsqueezedfrombolusandswallowedBoluschewedReinsalivationAddingmoresalivaRedeglutitionSwallowingbolusandliquidsFourStepsofRuminationRegurgRuminationAllowsanimaltoforageandeatfoodrapidly,andthenstoreforlaterdigestionReducesparticlesizeOnlysmallparticlesleavereticulorumenIncreasessurfaceareaformicrobialattachmentanddigestion/fermentationBreaksdownimperviousplantwallsFurtherstimulationofsalivaflow(salivaservestobufferrumen)RuminationAllowsanimaltoforRuminationTimeAveragetimesforagrazinganimalEating–8hoursRuminating–8hoursResting–8hoursRuminatingtimeisquitevariable(highvariation)Reducingforage:concentratedecreasesruminationReducingparticlesizeofforagedecreasestimespentruminatingRuminationTimeAveragetimesfMechanismofRumination:RegurgitationStimulus–digestainfibermatscratchingsurfacenearcardiacsphincterContractionofthereticulumforcesdigestatocardiaAnimalinhaleswithepiglottisclosedtoproduceavacuumCardiasphincteropensandesophagusdilatesNegativepressure(vacuum)sucksdigestaintoesophagusRapidreverseperistalsismovesdigestatomouthMechanismofRumination:RegurMechanismofRumination:Remastication,Reinsalivation,andRedeglutitionBolusisrechewedChewingisslowerandmoredeliberatethanduringinitialeatingphaseDigestareinsalivatedParotidglandssecretemoresalivaduringruminationthaneatingSalivafromparotidglandssecretemoreNaHCO3-thanotherglandsReswallowingAfterreswallowing,therumencontractstomoveswallowedbolusintotherumenMechanismofRumination:RemasRemasticationandRedeglutitionRemasticationReducingParticleSizeofIngestedFeedsChewingduringeating(minimal)PreparationforswallowingReleasesolubleconstituentsDamageplanttissuesformicrobialattachmentChewingduringremastication(extensive)DecreaseparticlesizeforpassageDamageplanttissuesformicrobialattachmentMicrobialdigestionReticuloruminalcontractionsReducingParticleSizeofIngeRumenContractionsInoculateincomingfeedwithmicrobesMixcontentsMinimizeeffectsofstratificationMovefermentationproducts(VFA’s)torumenwallParticlesortingandpassageofsmallparticlestoomasumRuminationEructationoffermentationgasesRumenContractionsInoculateinNeedforEructationPeakgasproductionoccurs30minto2hrpost-feeding(12-27liters/min)Averageis1-2liters/minApproximately30%ofCO2producedinrumenisabsorbedintobloodandremovedthroughthelungsRemainderiseructatedOnly20%oftheCH4isremovedthroughthelungs80%eructatedCompositionofrumengas__Gas___%__CO265.35CH4(variable)27.76N27.00O2(atwall).56H2.18H2S.01NeedforEructationPeakgasprControlofEructationStimulusGaseousdistensionofthereticulumandrumenEsophagusdilates&animalbelches12-30Lperminuteforcattle3-17timesperminuteInhibitionPresenceofdigestanearthecardiacsphincterAffectsallthreesphinctersProtectivemechanismtopreventdigestafromenteringlungsEpinephrine–fightorflightresponseInhibitionofeructationwillcausetheanimalstobloatRuminalpressureswillincreaseupto100mmHgStablefrothorfoamformedinrumenControlofEructationStimulusFeedInVFAMicrobialProteinVitaminsThenutrientspresentedtotheanimalafterruminalfermentationareverydifferentthanthoseenteringtherumenasfeedFeedtheMicrobes,LettheMicrobesFeedtheRuminant!FeedInVFAThenutrientspresenRumenDigestionandFermentation CO2

VFADegradableRumen MicrobialcellsFeed microbes NH3 CH4 Heat

Long-chain fattyacids H2SProductsinredareusedbythehostanimalProductslistedinblackarenotuseablebytheanimalRumenDigestionandFermentatiRumenMicroorganisms

NutritionalRequirementsCO2EnergyEndproductsfromdigestionofstructuralcarbohydratesFermentationofsugarsNitrogenAmmonia(majorityofnitrogenneeds)Aminoacids(cellulolyticbacteria)MineralsCo,S,P,Na,K,Ca,Mg,Mn,Fe,Zn,Mo,SeVitaminsNonerequiredinmixedculturesofbacteriaRumenMicroorganisms

NutritionSymbioticRelationshipMicrobesprovidetotheruminantDigestionofcelluloseandhemicelluloseProvisionofhighqualityproteinProductionofVFAProvisionofBvitaminsDetoxificationoftoxiccompoundsSymbioticRelationshipMicrobesDigestionofCelluloseandHemicelluloseCellulasesareallofmicrobialoriginWithoutmicrobes,ruminantswouldnotbeabletouseforagecropssuchaspasture,hayorsilageDigestionofCelluloseandHemProvisionofHighQualityProtein50-80%ofabsorbedNisfrommicrobesImprovedmicrobialefficiencywillprovidemoremicrobialproteinCangetover3kgofmicrobialproteinperdayincattleHighbiologicalvalueproteinsourceAminoacidpatternisverysimilartothatrequiredbytheruminantanimalProvisionofHighQualityProtMicrobesAsAFeedSourceMicrobesasafeedsourceBacteriaandprotozoawashedoutoftherumentoomasumandintotheabomasumAcidicenvironmentkillsmicroorganismsDigestedandabsorbedthesameasanyotherfeedsourceinstomachandsmallintestineProvideaminoacidsandsomeenergyMicrobesAsAFeedSourceMicroSourcesofenergyleavingrumen:VFA 70%Microbialcells 10%Digestibleunfermentedfeed 20%NoglucoseavailablefortheruminantConcentrationofVFAinrumen=50to125uM/mlEnergy

SourcesofenergyleavingrumeProvisionOfBVitaminsMeetstheruminant’srequirementsundermostconditionsSomesupplementationofspecificvitamins,suchasniacin,maybebeneficialinearlylactationdairycowsProvisionOfBVitaminsMeetstDetoxificationOfToxicCompoundsManypotentialtoxinsarede-toxifiedbyrumenmicrobesExample:MimosineinLeucaenacausesproblemsPoorgrowth,reproductionandhairlossHawaiianruminants,butnotthosefromAustralia,havemicrobesthatdegrademimosinesoLeucaenacouldbefedTransferredrumenfluidobtainefromHawaiiancattletoAustraliaInoculatedrumensofAustraliancattleFedLeucaenasafelytoAustralianruminants!DetoxificationOfToxicCompouSymbioticRelationshipRuminantsprovidetomicrobesHousingGarbageremovalNutrientsOptimalenvironmentforgrowthSymbioticRelationshipRuminantHousingReliableheat(39±2°C)Fluidenvironment(requiresfreewaterintake)85to90%waterGuaranteedhousingfor18to96hoursdependingondietandtypeofanimalStraw-fedwaterbuffalo–longestrumenresidencetimeformicrobesSmallselectivebrowsers(mousedeerorduiker)–shortestresidencetimeformicrobesHousingReliableheat(39±2°CGarbageRemovalAbsorptionofVFAEnergytoruminantEructationCO2andCH4PassageofindigestibleresidueandmicrobestolowerGItractRumenmixingtoseparateandsettlesmallparticlesGarbageRemovalAbsorptionofVNutrientsSubstratescomefromfeedstuffsthatanimalconsumesSalivaprovidesurea(Nsourceforbacteria)NutrientsSubstratescomefromOptimalEnvironmentForGrowthReducedenvironment(littletonooxygen)StrictanaerobicmicrobesinrumeninteriorFunctionalanaerobesnearrumenwallpH6.0to7.0SalivacontainsbicarbonateandphosphatebuffersCowsproduceupto50gallonsofsalivadailyContinuouslysecretedMoreaddedduringeatingandruminationCowruminates10-12hours/dayDecreasesinparticlesizeofforagereduceneedforrumination,decreasechewingtime,decreasesalivaproduction,andrumenpHplummetsOptimalEnvironmentForGrowth-Rumen-Fermentation-大學(xué)動物營養(yǎng)學(xué)課件-英文版OptimalEnvironment(pH)IfpH5.7ratherthan6.550%lessmicrobialsynthesisCellulolyticbacteriafunctionbestatpH~6.8RateofstructuralcarbohydrateuseisdecreasedAmylolyticbacteriafunctionbestatpH~5.8MorelactateandlessacetateisproducedFurtherdownwardpHspiralInconcentrateselectors(likedeer),parotidsalivaryglandsare0.3%ofbodyweightOptimalEnvironment(pH)IfpHBacteriaandpHToleranceSpeciesTypepHRuminococcusflavefaciensFibrobactersuccinogenesMegasphaeraelsdeniiStreptococcusbovisfiberfiberlactateuserlactateproducer6.1564.94.55BacteriaandpHToleranceSpeciMicrobes%ofmassGenerationintervalNo./mLBacteria60-9020min25-80billionProtozoa10-408-36h200-500thousandFungi5-1024hminimalMicrobes%ofmassGenerationinSymbioticRelationshipMicrobesprovidetotheruminantDigestionofcelluloseandhemicelluloseProvisionofhighqualityproteinProductionofVFAProvisionofBvitaminsDetoxificationoftoxiccompoundsRuminantsprovidetomicrobesHousingGarbageremovalNutrientsOptimalenvironmentforgrowthSymbioticRelationshipMicrobesRumenMicrobesBacteria>200specieswithmanysubspecies25speciesatconcentrations>107/mL1010to1012cells/mL99.5%obligateanaerobesRumenMicrobesBacteria

GroupsofbacteriaintherumenFree-livingintheliquidphaseLooselyassociatedwithfeedparticlesFirmlyadheredtofeedparticlesAssociatedwithrumenepitheliumAttachedtosurfaceofprotozoaandfungiEnvironmentalNichesforBacteriaGroupsofbacteriaintherum

AllowsbacteriatocolonizethedigestiblesurfaceoffeedparticlesBringsenzymes(frommicrobes)andsubstrate(fromfeedstuff)together

ProtectsmicrobialenzymesfromproteasesintherumenIfattachmentpreventedorreduced,digestionofcellulosegreatlyreducedRetentiontimeofmicrobesintherumenisincreasedtoprolongdigestionReducespredatoryactivityofprotozoaOver-feedingfattoruminantscancoatforages,reducingbacterialattachmentBenefitsofBacterialAttachmentAllowsbacteriatocolonizetBacterialPopulationsCellulolyticbacteria(fiberdigesters)DigestcelluloseRequirepH6-7UtilizeNinformofNH3RequireSforsynthesisofsulfur-containingaminoacids(cysteineandmethionine)Produceacetate,propionate,littlebutyrate,CO2PredominatefromroughagedietsBacterialPopulationsCellulolyMicrobialPopulationsAmylolyticbacteria(starch,sugardigesters)DigeststarchRequirepH5-6UtilizeNasNH3orpeptidesProducepropionate,butyrateandlactatePredominatefromgraindietsRapidchangetograindietcauseslacticacidosis(rapidlydecreasespH)MicrobialPopulationsAmylolytiMicrobialPopulationsMethane-producingbacteriaProducemethane(CH4)UtilizedbymicrobesforenergyRepresentlossofenergytoanimalReleasedbyeructationMicrobialPopulationsMethane-pRumenMicrobesProtozoaLarge(20-200microns)unicellularorganismsIngestbacteriaandfeedparticlesEngulffeedparticlesanddigestcarbohydrates,proteinsandfatsNumbersaffectedbydietRumenMicrobesProtozoaEntodinium(RumenProtozoa)Entodinium(RumenProtozoa)RumenMicrobesFungiKnownonlyforabout20yearsNumbersusuallylowDigestrecalcitrantfiberRumenMicrobesFungiLocationofMicrobesRumenWallRumenFluidFiberMatGasPhaseLocationofMicrobesRumenWallDietaryFactorsThatReduceMicrobialGrowthRapid,dramaticrationchangesTakes3-4weeksformicrobestostabilizeRestrictedamountsoffeedExcessiveunsaturatedfatBacteriadonotusefatforenergyInhibitfiberdigestionandmicrobialgrowthDifferenttypesoffathavedifferenteffectsDietaryFactorsThatReduceMiDietaryFactorsThatReduceMicrobialGrowthExcessivenon-structuralcarbohydrateLowersrumenpH(rumenacidosis)SlugfeedingFeedbarleyorwheat(rapidlyfermented)Topreventacidosis,mustbalancelactateusersandproducersDietaryFactorsThatReduceMiDietaryFactorsThatMaximizeMicrobialGrowthMaximumdrymatterintakeBalancedcarbohydrateandproteinfractionsatthesametimeBacterianeedbothenergyandNforaminoacidsynthesisGradualrationchangesFeedavailableatalltimesMaintainsstablerumenpHDietaryFactorsThatMaximize-Rumen-Fermentation-大學(xué)動物營養(yǎng)學(xué)課件-英文版RumenFunctionOverviewRumenFunctionOverviewRumenFermentationRumenFermentationRumenFermentationWorld’slargestcommercialfermentationspace100billionlitersorrumenvolumeindomesticanimals1010to1012cells/mLRumencapacityrangesfromlessthan1liter(1quart)inaduikerto200liters(50gallons)inacowRumenFermentationWorld’slargRuminantsContinuousculturefermentersInputandoutputLignocellulosicsubstrates(forages)digestedCellulasecomplexHemicellulasesNitrogencapture(NPN)8x1015mouthstofeedBecauseofthesemicrobialenzymes,ruminantscanutilizefeedstuffsthatprovidelittletononutritionalbenefittonon-ruminantsRuminantsContinuousculturefeFourStepsofRuminationRegurgitationReverseperistalsiscarriesfoodtomouthRemasticationLiquidsqueezedfrombolusandswallowedBoluschewedReinsalivationAddingmoresalivaRedeglutitionSwallowingbolusandliquidsFourStepsofRuminationRegurgRuminationAllowsanimaltoforageandeatfoodrapidly,andthenstoreforlaterdigestionReducesparticlesizeOnlysmallparticlesleavereticulorumenIncreasessurfaceareaformicrobialattachmentanddigestion/fermentationBreaksdownimperviousplantwallsFurtherstimulationofsalivaflow(salivaservestobufferrumen)RuminationAllowsanimaltoforRuminationTimeAveragetimesforagrazinganimalEating–8hoursRuminating–8hoursResting–8hoursRuminatingtimeisquitevariable(highvariation)Reducingforage:concentratedecreasesruminationReducingparticlesizeofforagedecreasestimespentruminatingRuminationTimeAveragetimesfMechanismofRumination:RegurgitationStimulus–digestainfibermatscratchingsurfacenearcardiacsphincterContractionofthereticulumforcesdigestatocardiaAnimalinhaleswithepiglottisclosedtoproduceavacuumCardiasphincteropensandesophagusdilatesNegativepressure(vacuum)sucksdigestaintoesophagusRapidreverseperistalsismovesdigestatomouthMechanismofRumination:RegurMechanismofRumination:Remastication,Reinsalivation,andRedeglutitionBolusisrechewedChewingisslowerandmoredeliberatethanduringinitialeatingphaseDigestareinsalivatedParotidglandssecretemoresalivaduringruminationthaneatingSalivafromparotidglandssecretemoreNaHCO3-thanotherglandsReswallowingAfterreswallowing,therumencontractstomoveswallowedbolusintotherumenMechanismofRumination:RemasRemasticationandRedeglutitionRemasticationReducingParticleSizeofIngestedFeedsChewingduringeating(minimal)PreparationforswallowingReleasesolubleconstituentsDamageplanttissuesformicrobialattachmentChewingduringremastication(extensive)DecreaseparticlesizeforpassageDamageplanttissuesformicrobialattachmentMicrobialdigestionReticuloruminalcontractionsReducingParticleSizeofIngeRumenContractionsInoculateincomingfeedwithmicrobesMixcontentsMinimizeeffectsofstratificationMovefermentationproducts(VFA’s)torumenwallParticlesortingandpassageofsmallparticlestoomasumRuminationEructationoffermentationgasesRumenContractionsInoculateinNeedforEructationPeakgasproductionoccurs30minto2hrpost-feeding(12-27liters/min)Averageis1-2liters/minApproximately30%ofCO2producedinrumenisabsorbedintobloodandremovedthroughthelungsRemainderiseructatedOnly20%oftheCH4isremovedthroughthelungs80%eructatedCompositionofrumengas__Gas___%__CO265.35CH4(variable)27.76N27.00O2(atwall).56H2.18H2S.01NeedforEructationPeakgasprControlofEructationStimulusGaseousdistensionofthereticulumandrumenEsophagusdilates&animalbelches12-30Lperminuteforcattle3-17timesperminuteInhibitionPresenceofdigestanearthecardiacsphincterAffectsallthreesphinctersProtectivemechanismtopreventdigestafromenteringlungsEpinephrine–fightorflightresponseInhibitionofeructationwillcausetheanimalstobloatRuminalpressureswillincreaseupto100mmHgStablefrothorfoamformedinrumenControlofEructationStimulusFeedInVFAMicrobialProteinVitaminsThenutrientspresentedtotheanimalafterruminalfermentationareverydifferentthanthoseenteringtherumenasfeedFeedtheMicrobes,LettheMicrobesFeedtheRuminant!FeedInVFAThenutrientspresenRumenDigestionandFermentation CO2

VFADegradableRumen MicrobialcellsFeed microbes NH3 CH4 Heat

Long-chain fattyacids H2SProductsinredareusedbythehostanimalProductslistedinblackarenotuseablebytheanimalRumenDigestionandFermentatiRumenMicroorganisms

NutritionalRequirementsCO2EnergyEndproductsfromdigestionofstructuralcarbohydratesFermentationofsugarsNitrogenAmmonia(majorityofnitrogenneeds)Aminoacids(cellulolyticbacteria)MineralsCo,S,P,Na,K,Ca,Mg,Mn,Fe,Zn,Mo,SeVitaminsNonerequiredinmixedculturesofbacteriaRumenMicroorganisms

NutritionSymbioticRelationshipMicrobesprovidetotheruminantDigestionofcelluloseandhemicelluloseProvisionofhighqualityproteinProductionofVFAProvisionofBvitaminsDetoxificationoftoxiccompoundsSymbioticRelationshipMicrobesDigestionofCelluloseandHemicelluloseCellulasesareallofmicrobialoriginWithoutmicrobes,ruminantswouldnotbeabletouseforagecropssuchaspasture,hayorsilageDigestionofCelluloseandHemProvisionofHighQualityProtein50-80%ofabsorbedNisfrommicrobesImprovedmicrobialefficiencywillprovidemoremicrobialproteinCangetover3kgofmicrobialproteinperdayincattleHighbiologicalvalueproteinsourceAminoacidpatternisverysimilartothatrequiredbytheruminantanimalProvisionofHighQualityProtMicrobesAsAFeedSourceMicrobesasafeedsourceBacteriaandprotozoawashedoutoftherumentoomasumandintotheabomasumAcidicenvironmentkillsmicroorganismsDigestedandabsorbedthesameasanyotherfeedsourceinstomachandsmallintestineProvideaminoacidsandsomeenergyMicrobesAsAFeedSourceMicroSourcesofenergyleavingrumen:VFA 70%Microbialcells 10%Digestibleunfermentedfeed 20%NoglucoseavailablefortheruminantConcentrationofVFAinrumen=50to125uM/mlEnergy

SourcesofenergyleavingrumeProvisionOfBVitaminsMeetstheruminant’srequirementsundermostconditionsSomesupplementationofspecificvitamins,suchasniacin,maybebeneficialinearlylactationdairycowsProvisionOfBVitaminsMeetstDetoxificationOfToxicCompoundsManypotentialtoxinsarede-toxifiedbyrumenmicrobesExample:MimosineinLeucaenacausesproblemsPoorgrowth,reproductionandhairlossHawaiianruminants,butnotthosefromAustralia,havemicrobesthatdegrademimosinesoLeucaenacouldbefedTransferredrumenfluidobtainefromHawaiiancattletoAustraliaInoculatedrumensofAustraliancattleFedLeucaenasafelytoAustralianruminants!DetoxificationOfToxicCompouSymbioticRelationshipRuminantsprovidetomicrobesHousingGarbageremovalNutrientsOptimalenvironmentforgrowthSymbioticRelationshipRuminantHousingReliableheat(39±2°C)Fluidenvironment(requiresfreewaterintake)85to90%waterGuaranteedhousingfor18to96hoursdependingondietandtypeofanimalStraw-fedwaterbuffalo–longestrumenresidencetimeformicrobesSmallselectivebrowsers(mousedeerorduiker)–shortestresidencetimeformicrobesHousingReliableheat(39±2°CGarbageRemovalAbsorptionofVFAEnergytoruminantEructationCO2andCH4PassageofindigestibleresidueandmicrobestolowerGItractRumenmixingtoseparateandsettlesmallparticlesGarbageRemovalAbsorptionofVNutrientsSubstratescomefromfeedstuffsthatanimalconsumesSalivaprovidesurea(Nsourceforbacteria)NutrientsSubstratescomefromOptimalEnvironmentForGrowthReducedenvironment(littletonooxygen)StrictanaerobicmicrobesinrumeninteriorFunctionalanaerobesnearrumenwallpH6.0to7.0SalivacontainsbicarbonateandphosphatebuffersCowsproduceupto50gallonsofsalivadailyContinuouslysecretedMoreaddedduringeatingandruminationCowruminates10-12hours/dayDecreasesinparticlesizeofforagereduceneedforrumination,decreasechewingtime,decreasesalivaproduction,andrumenpHplummetsOptimalEnvironmentForGrowth-Rumen-Fermentation-大學(xué)動物營養(yǎng)學(xué)課件-英文版OptimalEnvironment(pH)IfpH5.7ratherthan6.550%lessmicrobialsynthesisCellulolyticbacteriafunctionbestatpH~6.8RateofstructuralcarbohydrateuseisdecreasedAmylolyticbacteriafunctionbestatpH~5.8MorelactateandlessacetateisproducedFurtherdownwardpHspiralInconcentrateselectors(likedeer),parotidsalivaryglandsare0.3%ofbodyweightOptimalEnvironment(pH)IfpHBacteriaandpHToleranceSpeciesTypepHRuminococcusflavefaciensFibrobactersuccinogenesMegasphaeraelsdeniiStreptococcusbovisfiberfiberlactateuserlactateproducer6.1564.94.55BacteriaandpHToleranceSpeciMicrobes%ofmassGenerationintervalNo./mLBacteria60-9020min25-80billionProtozoa10-408-36h200-500thousandFungi5-1024hminimalMicrobes%ofmassGenerationinSymbioticRelationshipMicrobesprovidetotheruminantDigestionofcelluloseandhemicelluloseProvisionofhighqualityproteinProductionofVFAProvisionofBvitaminsDetoxificationoftoxiccompoundsRuminantsprovidetomicrobesHousingGarbageremovalNutrientsOptimalenvironmentforgrowthSymbioticRelationshipMicrobesRumenMicrobesBacteria>200specieswithmanysubspecies25speciesatconcentrations>107/mL1010to1012cells/mL99.5%obligateanaerobesRumenMicrobesBacteria

GroupsofbacteriaintherumenFree-livingintheliquidphaseLooselyassociatedwithfeedparticlesFirmlyadheredtofeedparticlesAssociatedwithrumenepitheliumAttachedtosurfaceofprotozoaandfungiEnvironmentalNichesforBacteriaGroupsofbacteriaintherum

AllowsbacteriatocolonizethedigestiblesurfaceoffeedparticlesBringsenzymes(frommicrobes)andsubstrate(fromfeedstuff)together

Protectsmicrobialenzyme

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