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TurkJVetAnimSci 29(2005)571-580 ©TÜB‹TAK RewievArticle UseofBovineSomatotropinintheManagementof TransitionCows

MehmetfiükrüGÜLAY*,FatmafiefikaHAT‹PO⁄LU DepartmentofPhysiology,FacultyofVeterinaryMedicine,AkdenizUniversity,Burdur-TURKEY *e-mail:[email protected]

Received:17.03.2004

Abstract: Themajorobjectivesofanefficientdairyfarmoperationincludeasuccessfulperiod,highyieldrelativeto thefeedcosts,reproductivecompetence,andfinallythereturningofthecowtothebodyconditionscore(BCS)thatexistedbe fore lactationsoshewillbepreparedforanotherlactation.Incowsamilkyieldresponsetobovinesomatotropin(bST)treatmentsat60 dafterparturitionhasbeenwellstudiedandfullydocumented.MilkproductionresponsetobSToccursbecauseofitsknowneffects onpartitioningofnutrientsandbecauseagreaterproportionofthenutrientintakeisusedformilksynthesis.Itincreasesl iver glucoseoutput,cardiacoutput,bloodflowtothemammaryglandanduptakeofnutrientsusedformilksynthesisbythemammary glandamongothereffects.Inaddition,STdecreasestherateofoxidationofaminoacidandglucoseandthereforedecreasesglucose clearance.TreatmentwithbSTresultsincoordinatedchangesofvariousorgansandtissuesthatnaturallyoccurduringthetransition fromanonlactatingtolactatingstatewhencirculatingconcentrationofSTishigh.BecauseoftheknowneffectsofbSTdescri bed above,useofbSTduringthetransitionperiodmaycausepositiveandbeneficialeffectssupportingmilksynthesispriortoparturition. ThisreviewrecapsthediversephysiologicaleffectsofSTonlactation,discussesthemajormechanismsthatmediatetheseeffe cts, andsummarizestherecentstudiesusingbSTduringthetransitionperiodtoincreasemilkproductionincows.

KeyWords: Somatotropin,lactation,transitionperiod,metabolism,dairycow.

GeçiflDönemi‹neklerdeBovineSomatotropinKullan›m›

Özet: Sütüreticilerininenönemliamaçlar›baflar›l›birlaktasyondönemi,yemmaliyetineoranladahayükseksütüretimi,do¤um sonras›hayvanlar›ntekrargebeb›rak›lmas›veensonolarakdahayvanlar›ndo¤umsonras›sahipolduklar›kilolaratekrar ulaflt›r›lmas›d›r.Bovinesmatotropin(bST)kullan›ld›¤›ndasütverimindekiart›fl›nnedeni,besinlerinvücutda¤›l›m›naolanetk isiile al›nanbesinlerinbüyükbirbölümününsütsentezlenmesiiçinyönlendirmiflolmas›d›r.bSTbirçoketkisininyan›s›raayr›ca karaci¤erdenglikozsal›verilmesini,toplamkalphacmini,memebezinegelenkanmiktar›n›vesütüretimiiçingerekliolanbesi n al›m›n›art›r›r.Bunlarailaveten,bSTaminoasitveglikozunoksidasyonunuveböyleceglikozkullan›m›n›daazalt›r.bST kullan›ld›¤›ndabirçokdokuveorgandakoordineliolarakde›iflikli¤esebepolur.Bude¤iflikliklergeçifldönemindekandakiST’ni n yüksekoldu¤undagörülende¤iflikliklerilebenzerdir.bST’ninyukar›dadabahsedilenetkilerindendolay›,geçifldöneminde kullan›lmas›sütüretimininart›r›lmas›aç›s›ndanolumluveyararl›olabilir.Buderlemekapsam›nda,ST’ninyararl›etkilerive ana mekanizmalar›tart›fl›larakgeçifldönemindebSTkullan›m›ilesütüretimineetkisiüzerineyap›lanyeniaraflt›rmalarhakk›ndabi lgi verilmesiamaçlanm›flt›r.

AnahtarSözcükler: Somatotropin,laktasyon,geçifldönemi,metabolizma,sütine¤i

Introduction production.ResearchcontinuedduringandafterWorld Asearlyasthe1930s,thegrowth-promotingeffects WarIIasscientistssoughtaneffectivemeansof ofsomatotropin(ST)werecharacterized.Evansand increasingfoodproduction.However,theamountofST Simpson(1)showedthatcrudeextractsofbovine fromeachpituitarywassolowthatthissourcewouldnot pituitaryincreasedthegrowthrateofrats.In1937,two bepracticalforimprovingtotalmilkproduction(3). Russianscientists,AsimovandKrouze(2),performedthe Thearrivalofmoderntechnologyenabledthe firstbSTresearch,using600lactatingdairycows.These developmentofrecombinantbovinesomatotrophin researchersfoundthatinjectingcowswithextractsofthe (rbST),whichprovidedanunlimitedsourceofSTfor pituitariesofslaughteredcowsresultedinincreasedmilk researchand,potentially,forcommercialapplication.This

571 UseofBovineSomatotropinintheManagementofTransitionDairyCows

allowedresearcherstoconductmanystudiesusing graduallyreturnstonormalpre-injectionlevelsafter recombinantbST.Fromthesestudiesitwasconcluded cessationoftreatment(11). thatexogenousSTcouldincreasemilkproductionofdairy ObtainingaMYresponsetobSTdoesnotrequire cowsatleast10%to15%.Obtainingamilkyield specialdietsordifferentfeedingredients.SubstantialMY responsetobSTdidnotrequirespecialdietsordifferent responseshavebeenobservedondietsrangingfrom feedingredients.However,treatedcowsdidneed pasturetothemoretypicaltotalmixedration(TMR) adequateamountsofbalanceddietthatcontainsall diets.However,drymatterintake(DMI)increasesinbST nutrientsnecessaryforsupportingexpectedmilk treatedcowsafterafewweeksofthesupplementation production(4). andpersiststhroughouttheintervalofbSTuse.Thus, TreatmentofcowswithbSTduringlactationwas treatedcowsrequireadequateamountsofabalanceddiet approvedforuseintheUSin1994.Sincethenithas ratherthanaspecialdiet(10). beenusedextensively.Inaddition,morethan34 a)EffectsofbSTontheMammaryGland countrieshavereachedsimilarconclusionswithrespectto foodsafety,andapprovedtheuseofbST,suchasAlgeria, InearlystudiesconcerningtheactionsofST,itwas Brazil,Bulgaria,Columbia,CostaRica,CzechRepublic, concludedthatadirecteffectofSTonmammarygland Honduras,Hungary,Jamaica,Kenya,Korea,Malaysia, developmentwasunlikely.StudiesfailedtodetectanST Mexico,Namibia,Pakistan,Peru,Romania,Russia, receptoronthemammaryepithelialcellofthecow(12). Slovakia,SouthAfrica,Turkey,UnitedArabEmirates, Moreover,infusionsofSTdirectlyintomammaryartery UkraineandZimbabwe. ofsheepdidnotstimulatemilkproduction(13). However,inrodents,adirecteffectofSTonmammary Bovinesomatotropinhadbeengivento335,000 developmentandfunctionwasestablished(14,15). cowsinNewYork(45%ofthestatetotal),themajor AlthoughmostevidencesuggestedanindirecteffectofST dairystateintheeasternUS,afteroverayearperiod. onmammaryglanddevelopmentinruminants,itwas About55%ofallsalesofbSThavebeentofarmerswho foundthatruminantsalsoexpressedmRNAfortheST have100orfewercows(5).In1998,ofnearly9million receptorinmammarygland(16).Furthermore, dairycowsintheUS,about25%wereinbSTtreated immunologicstainingofSTreceptorsinmammarytissue herds,and300additionaldairyfarmerswerereportedto ofpregnantandlactatingcowswasalsoreported(17). beadoptinguseofbSTpermonth.Theaveragedairy Collieretal.(18)observedasignificanteffectofSTon farmerusedthecommercialbSTproduct(POSILAC®)to mammarygrowthofpregnantheiferswhenitwas supplementmorethan50%oftheherddependingupon administeredthroughtheteatcanal.However,as individualherdmanagementpracticesandstageofthe indicated,unilateralclosearterialinfusionofSTintoone- adoption. halfofthemammaryglandofsheepdidnotincreasemilk yieldperformancecomparedwiththeuninjectedhalf Responsestosomatotropinsupplementation (13).Furthermore,STadministrationthroughtheteat canaloflactatinggoatsdidnotresultinincreasedmilk Somatotropinisamajorregulatorofgrowthin productionresponse(19).Thislatterresponsewouldnot mammals.IntheliverSTregulatestheexpressionofa beexpectedunlesstherewasuptakeofSTacrossthe widerangeofincludinghormonereceptorsand apicalmembraneoftheepithelialcellsoruptakeintothe growthfactors,secretoryproducts,andenzymessuchas generalcirculationandsubsequentactioninmammary cytochromeP450(6).Somatotropinalsohasaunique gland. effectonstimulatingmammaryglanddevelopment(7) andlactation(8). TheacuteriseinMYinresponsetobSTandrapid declineafterstoppingbSTinjectionarguesagainstcell Typicalmilkyield(MY)responsetobSTisanincrease proliferationbeingcausedbySTintheshort-term. ofatleast10-15%.However,theresponsecanbemuch Knightetal.(20)observedthatSTtreatmentdidnot greaterwithbettercareandmanagementofthecows affecttheamountofmammaryparenchymaandBrdU- (9,10).AfterthefirstfewdaysofbSTtreatment,MY labelingofmammaryepithelialcellsinvivoduringthe increasesgraduallyandreachesapeakandtheincreaseis first6wkoflactation.Ontheotherhand,alarge maintainedwithcontinuousinjectionsofbST.Milkyield

572 M.fi.GÜLAY,F.fi.HAT‹PO⁄LU

increaseinparenchymalvolumewasobservedinmid- suggeststhatthemechanismofactionofbSTwithregard lactationgoatstreatedovera22wkperiodwith tostimulatoryeffectsonMYcannotbelimitedonlytothe exogenousST(21).Otherstudieswithcowsandgoats knowneffectsofIGF-I. reportedtrendsand/orsignificantincreasesinkey b)EffectsofSomatotropinonOtherTissues enzymesinthemammarygland(suchasacetylCoA carboxylase,acetylCoAsynthetaseandfattyacid Somatotropinexertsmanydifferenteffectson synthetase)duetobSTsupplementation.Thus,bST metabolisminbothmammaryandnon-mammarytissues. directlyand/orindirectlycausesanincreaseintheratesof BovinesomatotropinincreasesMYwithnochangeinmilk milksynthesispercellandimprovedmaintenanceof composition,unlessthereisenergydeficiency.It secretorycells(22,23). provokesMYincreasebyincreasingoverallbloodsupply tomammarygland(wateruptake),alteringglucose Somatotropinshowsoneofitseffectsonmammary metabolism(lactosesynthesisinthemammarygland), glandfunctionindirectlyviatheactionoflike enhancinglipolysisandincreasinglipogenesisinthe growthfactor-I(IGF-I)onthetissue(24). mammarygland(supplyoflipidprecurcors),andalso ConcentrationsandactionsofIGF-Iarelikelythemost modifyingproteinmetabolism(supplyofaminoacids).As importantlinktotissueresponsewithhigher aresult,milkyieldincreasesandthecompositionofmilk concentrationsofST.Somatotropinhasamajorrolein doesnotchangefromnormal(31). regulatingtheconcentrationsofIGFsthatareinthe circulation.ConcentrationsofIGF-IincreaseduringbST IncreasedMYresponseduetoSTresultsmainlyfrom treatmentsbecauseofbothinducedreleaseofIGF-Ifrom alteredpartitioningofnutrientsinfavorofmammary ahepaticstoragepoolandgreaterbiosynthesisofIGF-I. glandsandfromanincreaseinthesynthesizingcapacity BiosynthesisisregulatedbyincreasedmRNAlevelsand and/orlongevityofthemilksynthesizingcells(22). mRNAstability(25).IncreaseinIGF-Iconcentrationwas Somatotropinisaprimaryhomeorheticregulatorduring maintainedwithcontinuedinjectionsofSTandthe pregnancyandlactation;itregulatespartitioningof increaseinMYwasparalleltotheincreaseinIGF-I. nutrients(carbohydrates,lipids,proteins,andminerals) CessationofSTsupplementationcausedaparallel andplaysanimportantroleinthecoordinationofvarious decreaseinbothMYandbloodconcentrationsofIGF-I organsandtissues(4).Tosupportmilksynthesis,the (25). metabolismofothertissuesisstimulatedtoprovidethe necessaryprecursors. Insulinlikegrowthfactor-Ihasbothautocrineand paracrineactionsinadditiontoitsendocrineactions(26). Carbohydratemetabolism Atthecellularlevel,IGF-Ilocallystimulatesaminoacid Duringearlylactation,glucoseisusedalmost transport,synthesisofRNAandDNA,andsynthesisof exclusivelybythemammaryglands(exceptionsare cellularproteins(27).ReceptorsforIGF-Iwere nervoussystemandbrain),andMYisheavilydependent demonstratedinmammarytissueandIGF-Iisalocal uponincreasedglucosesupplytothegland.IfbSTisto mediatorofmammaryepithelialgrowthanddevelopment increaseMY,thenitmustactinawaytodirectmore (28).Intralobularstromalcells,smallbloodvessels,and glucosetothemammarygland.Thiscanoccurthrougha capillariescontainIGF-Ireceptorsinthemammarygland varietyofindividualactionsofbST.First,bSTincreases (29).DuringSTtreatments,IGF-Ibindsinthecytoplasm mammarybloodflowsothatmorebloodperfusesthe andthestromaofepithelialcells.Insulinlikegrowth mammaryglandandincreaseduptakeofglucosecan factor-Istimulatesthecellularactivityofmammarygland, occur(31).Adecreasedabilityofinsulin(INS)toinhibit itincreasessynthesisofRNAandDNA,andthesynthesis gluconeogenesisisobservedfollowingbSTtreatments. ofcellularproteins.Therefore,itisalocalmediatorof Bothinvivo(32)andinvitro(33)studiesdemonstrated mammaryepithelialgrowthanddevelopment(27). thathepaticratesofgluconeogenesiswereincreased Interestingly,closearterialinfusionsofIGF-Iinto duringtreatmentofdairycowswithbST.Furthermore, mammaryglandofgoat,butnotsystemicinfusionsof STdecreasedsensitivityofINSreceptorstoINSinthe IGF-I,increasedMYdramatically(30).However, peripheraltissuesandthisresultedindecreasedoverall lactationalresponsetoclosearterialinfusionofIGF-Iwas uptakeofglucoseinperipheraltissues.Thisminimized muchlessthanthatofsystematicbSTtreatmentswhich theoxidationofglucosetoCO 2,andasaconsequence

573 UseofBovineSomatotropinintheManagementofTransitionDairyCows

moreglucosewasmadeavailabletotheudder. ofothersubstratesavailable,inlargepart,dueto Somatotropinalsostimulatedincreasedfeedintake,and combinedactionsofSTandINS. asaresult,morepropionatewasproducedinthereticulo- WhencowsareinnegativeEB,STstimulateslipolysis; rumenandbecameavailableforgluconeogenesis(34).In italtersthesensitivityofadiposetissueto β-adrenergic vitrostudiesoflivertissueofSTtreatedcowsshoweda agents(22).Therefore,forcowsthatareinnegativeEB 60%increaseincapacitytousepropionateforglucose andarebeingtreatedwithbST,increasedlipid synthesis(33). mobilizationwouldbeamajorsourceofenergyneeded Pyruvatecarboxylase(PC)andphosphoenolpyruvate tosupportmilkproduction(37).Lipolysisisregulatedby carboxykinase(PEPCK)arepotentialrate-limiting asignaltransductionsystemthatincludescAMP, enzymesforhepaticgluconeogenesisduringthe stimulatoryGproteins(Gs)andinhibitoryGproteins(Gi). transitionperiod(35).Itwasspeculatedthatexogenous CatecholaminesactthroughtheG s systemtostimulate STwouldincreasemRNAsynthesisintheliverthatcoded lipolysis,whereasexertsitsantilipolyticaffects fortheseenzymes(11).Ontheotherhand,Pershinget viatheGi system.Whenadenosinebindstoitsreceptorit al.(36)concludedbSTstimulationofmilkproductionwas stimulatesG i whichuncouplesG s proteinactivationof notmediatedthroughenhancedlivergluconeogensisin adenylcyclasecatalyzedbycatecholamines.Thisleadsto cowswith80daysinmilk(DIM).Somatotropinalso theinhibitionofthelipolyticpathwaythatisstimulatedby increasedlipidmobilizationandmoreglycerolthenwas catecholamines.Somatotropinalterslipolysisthroughan increaseinresponsetocatecholamineswithnochangein availableasaprecursorforgluconeogenesis.Tosumup, sensitivity.EpinephrinechallengefollowingSTtreatment thetotaloftheseactionswouldbethathigherblood dramaticallyincreasednon-esterifiedfattyacid(NEFA) glucoseconcentrationsoccurredandthisglucosecouldbe concentrationsinplasma.Interestingly,STtreatment directedtothemammaryglandtosupportlactation.In resultedinmodestchangesin β and α adrenergic additiontoincreasedglucoseproductionintheliver, 2 receptornumbers.However,theactivityofG proteins glucoseusagebyotherleanbodytissuesdecreased. i wasreducedsignificantlybySTtreatments.Asaresult,it Becauseglucoseisusedasaprimaryenergymetabolite hasbeensuggestedthatSTimpairedtheabilityofG i to andasasubstrateforsynthesizingmilkconstituents, interactwithadenylcyclase.This,inturn,wouldincrease energyneededbyotherperipheralbodytissueswouldbe theeffectivenessofG s systemstimulatedby derivedfromproductsoflipolysisormetabolismofnon- catecholamines(11). gluconeogeniccompoundsarisingfromtherumenand Theeventsdescribedabovewoulddramatically lowerdigestivetract. increasemobilizationoflipidsfromtheadiposetissue, Lipidmetabolism andincreasebloodNEFAandglycerol.Thus,therewould Nutritionalstatusplaysamajorroleintheregulation begreatlyreducedfattyacidsynthesisornonet oflipidmetabolism.ExogenousbSTgiventoanimals synthesis,andhence,lessacetateandglyceroluseinthe altersbothlipogenesisandlipolysisinadiposetissuewith adiposetissue.Therefore,thenetresultwouldbeashift theneteffectbeingrelatedtoenergybalance(EB)(11). intheavailabilityofthesemetabolitesinthemammary WhencowsareinpositiveEB,synthesisanddepositionof glandwheretheycanbeusedforsynthesisofshortand lipidsinadiposetissuearereducedbyST,which,inturn, mediumchain-fattyacidsthatarethemselvesusedforTG increasestheavailabilityofnutrientsandtheirutilization synthesisandmilkproduction.Therefore,lipolysisalso formilkproduction(22).Insulinperformsimportant mustbeanimportantpathwaytoprovideneeded precursorsintheearlypostpartumperiodofcows homeostaticcontrolintheregulationoflipidmetabolism. especiallytosupplytheenergyneededformilk SomatotropinreducestheabilityofINStostimulate production(11). lipogenesisinadiposetissue.Thus,STreducestheaction ofINS,suppresseslipogenicenzymeactivity,andreduces Aminoacidmetabolism glucoseuptake(22).ThesecoordinatedchangesinINS Somatotropintreatmentincreasesmilkprotein actionswouldsupportproductionofglucosefrom synthesisinlactatingcowsviaimprovedefficiencyof availableprecursorsandconservationofglucosefor aminoacidutilization.Areductionincirculatingurea mammaryusebyshiftingperipheraltissuestoutilization andinurinarynitrogenlosswerereported

574 M.fi.GÜLAY,F.fi.HAT‹PO⁄LU

followingbSTtreatment(31).Duringnegativeenergy AlthoughincreasedMYresponsestypicallyarefound balanceSTwillspareproteinuseasasourceofenergyin whencowsareinjectedwithbSTbeyond60d tissuesbecauseitincreaseslipidmobilizationand postpartum,nosuchincreaseinMYwasobservedwhen enhancesglucosemetabolism.Proteinsthatare exogenousbSTwasinjectedduringprepartumandearly mobilizedfromthemusclescanbeusedintheliver,inthe postpartumperiods,whichincludedtheso-called gut,andinthebloodandthiswillincreaseoverall transitionperiod.BinesandHart(41)speculatedthat metabolismandefficiencyofproteinuse.Aminoacids becauseofdelayedincreaseinDMI,treatmentwithbST mobilizedhaveanimportantroleinsupportinggrowthof duringthetransitionperiodmightleadtoacuteanimal someorgans(liver,heart,digestivetract)ratherthanto problemssuchasketosis,fattylivers,wasting,and supportmilksynthesis(38).Amountsofaminoacidsthat increasedsusceptibilitytootherdiseases,oritcould areoxidizedtoprovideenergywillbereducedandthe resultinlowerthanexpectedMYresponses. proteinmobilizedcanbeusedforgrowthofspecific a)bSTinjectionsduringprepartumperiod tissuesandmilkproteinsynthesis.Asindicated,STcauses anincreaseinfeedintake,whichwillmakemore StudiesbyEppardetal.(42)failedtoshowan nutrientsandaminoacidsavailabletoincreaseMY,and increaseinMYwhentheyinjectedHolsteinandJersey thiswilllessentheneedfortissuemobilizationprocesses. cowsduringtheprepartumperiodwithafullstandard doseofbST(POSILAC®).Simpsonetal.(43) Mammarybloodflow administeredgrowthhormonereleasingfactor(GRF) Ithasbeenestablishedthatnutrientsupplytothe prepartumtobeefheiferstoincreasesecretionofST mammaryglandisoneofthemajorlimitationsforthe beforeparturitionandduringearlylactation.Treated activityofsecretorycellsandmilksynthesis.Alactating heiferslostmorebodyweights(BW)andhaddelayed mammaryglandplacesaheavydemandontheanimalto ovarianactivity,whereasnodifferencewasobservedin providesubstratesformilksynthesis(31).WithbST MY.Inanothertrial,Holsteincowsreceived0,5or14 injections,alongwiththeincreasesthatoccurinMY, mgbST/dduringthelast46dbeforeparturition(44). cardiacoutputalsoincreases.Insulinlikegrowthfactor-I Cowstreatedwith14mgbST/dhadincreasedyieldsof hasaroleintheincreasedbloodflowtothemammary solids-correctedmilk(SCM)onlyduringwk1.However, glandthatappearstobemediatedbyproductionofnitric theyfoundnodifferencesinSCMamongtreatments. oxide(30).SomatotropinalsoincreasesconversionofT 4 Exceptforthecowstreatedwith5mg/dofbSTduring tothetissueactivethyroidhormoneT specificallyinthe 3 wk10oflactation,EBwasnegativeforallcowsduring mammaryglandandtheincreaseinT alterslocal 3 thefirst70doflactation.Bachmanetal.(45)evaluated metabolismandhelpsmediatingthegalactopoietic whetheradoseof25mgbST/dadministeredprepartum response(39).Thus,bothincreasedmilksecretionand affectedpostpartumMYofHolsteincows.After localmetabolismdrivemorebloodtomammarygland covarianceadjustmentforprevioustotallactationMY, andthis,inturn,suppliesgreaterquantitiesofwaterand nutrientsneededformilksynthesis. 3.5%correctedmilk(FCM)yieldsoftreatedand controlcowsdidnotdiffer.TheyconcludedthatbST treatmentsduringprepartumperioddidnothaveeither bSTduringtransitionperiod apositiveoranegativeeffectonMYduringthefollowing Theapprovedtimeduringthelactationcycleofcows lactation.NoneofthestudiesthatusedbSTtreatments tostartuseofbSTisataboutpeakMY(~60±3d)and duringthetransitionperiodhavereportedacuteanimal itsuseiscontinuedthroughouttheremainderofthe healthproblemssuchasketosis,fattylivers,wasting,and lactation(40).Milkyieldgraduallyincreasedoverthefirst increasedsusceptibilitytodiseasesinthetreatedcows. fewdaysfollowingbSTtreatmentandreachedto Stelwagenetal.(46)administered20or40mgbST maximumconcentrationduringthefirstweek.Despite dailytoHolsteinheifersduringthelasttrimesterofthe largeincreasesinmilkproduction,feedintakedidnot firstpregnancy.TheyfoundasignificantincreaseinFCM increaseimmediatelyfollowingbSTtreatment.Early productionoftheheifersinjectedwith20mgbST/dbut productionresponsestherefore,wereduemostlyto onlyafter90doflactation.Putnametal.(47)reported partitioningofnutrientsawayfrombodytissuesto asignificanteffectofprepartumbSTtreatmentsonmilk mammaryglandusetosupportincreasedmilksynthesis. productionduringearlylactationwhichappearedto

575 UseofBovineSomatotropinintheManagementofTransitionDairyCows

increaseaslactationprogressed.TheexogenousbST MY.Inanotherstudy,thesamedoseofbSTwasinjected injections(500mgover14dperiod)wereinitiated28d every14dfrom10to150DIM(51).Theyconcluded priortoexpectedcalvingdateandwerecontinueduntil thatbSTinjectedearlyinlactationincreasedMY. parturition.CowstreatedwithbSTintheirtrialproduced However,increaseinMYoccurredattheexpenseofan 3.3kg/dmoremilkthanuninjectedcontrolsduringthe extensiveperiodofNEBanddecreaseinBWandBCS first42doflactation.However,cowsinthebSTtreated despiteanincreaseinDMIwasobservedduetobST grouphadsignificantlyhigherinitialbodyconditionscore treatment. (BCS)thanthecontrolswhentheywereassignedtothe Richardetal.(52)reporteda6%increaseinMY trial.Thisallowedtreatedcowstomobilizemorebody whencowswereinjectedwith50IUofbSTstarting reservesthancontrolsandtolosemoreBCSwithouta postpartum20th d;milkfatalsowaselevatedby25%.In negativeeffectduetobST.Thisisaveryimportant thesametrial,whencowswereinjectedbeginningon0 factorbecausebSTuselikelyincreasesnegativeenergy d,MYresponsewasgreater(12%)withnochangein balanceandgreaterlossofbodyweight,whichsupports milkfat.Chalupaetal.(53)treatedcowsfedadietthat thegreatermilkproductionuntiltheDMIincreases. contained0%or1.2%sodiumbicarbonatewith50IU b)bSTinjectionsduringpostpartumperiod bSTstartingat4 th wkoflactation.Theyreportedthat Asreportedforprepartumtreatments,early bSTtreatmentincreasedMYby4.7kgovercontrolcows postpartuminjectionsofbSTalsoshowedinconsistent andtherealsowasa1.05pointincreaseinmilkfat results.deBoeretal.(48)injectedcowswith20.6mg percentage.Feedintakeofinjectedcowstendedto bST/dstartingpostpartum4-9d.Nosignificant increase(17.6kgvs.16.1,P<0.11).Inoneofthe differencesweredetectedforMYofcontrolvs.bST largesttrials,Stanisiewskietal.(54)injected5mgor14 treatedcows.Unfortunately,thecowsassignedtobST mgbST/dfrom14dpostpartumthrough60DIM.Cows hadlowerMYpotentialbasedontherateandextentof thatreceivedeither5or14mgofbST/dproducedmore declineinMYaftercessationofbSTinjection.Thus,bST FCMthancontrols,butFCMofthetwobSTtreated injectionsenhancedMYtolevelssimilartothoseof groupsdidnotdiffer.However,cowsreceivingthelower controls(48).Moallemetal.(49)studiedthe dose(5mg/d)hadhigherpregnancyrateandhigher mechanismsofhowCasoapsoffattyacidsandbSTaffect conceptionratethanallotherexperimentalcows.Cows productionandreproductionofhighproducingcows. receiving5mgbST/dalsomaintainedBCSasuninjected Theyinjected500mgofbSTevery14dfrom10to150 controls. DIM.Milkyieldduringthefirst60ddidnotdiffer Variableresultswithinandamongtrialsthatevaluated betweentreatments.However,bSTtreatment useofbSTeitherpre-orpostpartumperiodmayhave significantlyenhancedMYbeyond60dandthepeakmilk beendue,inpart,todifferencesamongthedoses,diets productionincreasedforinjectedcows.Ontheother theanimalswerefed,orduetodifferencesinBWand hand,theeffectoftreatmentonBCSwassevere.The BCSoftheanimals.Usually,ahighdoseofbSTincreased BCSofinjectedcowsdecreasedmoreandwas thelengthoftimecowswereinNEB;lossofBWandBCS considerablylessfortreatedcowsandpostpartum occurredeventhoughanincreaseinDMImighthave conceptionratewasaffectedadverselycomparedto occurred.Thus,adequateBCS(55)forcowsinjectedwith uninjectedcows.Santosetal.(50)investigatedeffectsof bSTpre-and/orpostpartumisrequiredbecausethecows bSTontheperformanceofearlylactationcowsfeddiets requiregoodmanagementandadequatenutritionto differinginruminallydegradablestarch.Holsteincows produceandreproducewell. receivedbiweeklyinjectionsof500mgbSTfor90d c)bSTinjectionsduringbothprepartumand startingat5DIM.ApositiveMYresponsewasobserved postpartumperiods duringthefirst45dandforthetotaltreatmentperiod (90d).Interestingly,MYresponsetobSTwasless TreatmentwithalowerdoseofbSTduringboth between7and13wkthanfrom1to6wk.NeitherEB prepartumandpostpartumperiodshasapotentialto norBCSweredeterminedinthisstudy.Authors causemetabolicchangesafterparturitionthatare concludedthattheresponsetobSTwaslessthanusually beneficialtothehealthandperformanceofthecows observedforcowswhenbSTinjectionsbeganatpeak (56).Inonestudy,pre-andpostpartuminjectionsof

576 M.fi.GÜLAY,F.fi.HAT‹PO⁄LU

15.3mgbST/dshowedincreasedconcentrationsofST, first28dofpostpartumperiod;theincreaseinDMIwas

IGF-IandT 3 inplasma(57).Evidencesuggeststhat sameforbothtreatedanduntreatedcows(59).Inboth changesincirculatingconcentrationsofhormones, studies,bSTtreatedcowsproducedmoremilkand3.5% growthfactorandglucosecanbebeneficial.Inthesame FCMduringtheinjectionperiod.Inthefirststudy,no study,pre-andpostpartuminjectionsofbSTincreased carryovereffectsofbSTweredetectedonMYas DMIofcowsafterparturition,andtherewasasmaller evidencedbythefactthattheincreaseinMYdidnot decreaseinBCSandBW.Thisallowedthecowsto persistafterbSTinjectionswerestoppedaroundd42 recovertosatisfactoryBWandBCSmorerapidlyduring postpartum(58).Ontheotherhand,duringthesecond earlylactation.Cowsalsoproducedquantitativelygreater study,treatedcowsalsoproducedmoremilkwhenall dailyMYand3.5%FCM.Thus,thechangesin cows(controlsandtreated)wereinjectedwithfulldose concentrationsofmetabolichormoneslikelyhadarolein ofbST(>60doflactation).Theyalsohadhigher thepositiveeffectsonDMI,BCS,BWandMYofthese concentrationsofST,INSandIGF-Iprepartumandhigher cows. STandIGF-Ipostpartum(59).Thefindingsofthese Gulayetal.conductedtwoconsecutiveexperiments studiesshowthatbSTinjectionhasgeneralandlikely withbSTduringthetransitionperiod.Duringthefirst morediverseeffects,whichresultsingreaterMYand experiment(58),injectionsofbST(10.2mgbST/d) DMIwithoutapparentnegativeeffects.Indeed,results beganapproximately21±3dbeforeexpectedcalving indicatedtheoveralleffectsofprepartumandearly datesandbiweeklyinjectionsofbSTcontinuedupto postpartumbSTwerebeneficialsinceBWandBCSwere postpartum42 nd d.NoinjectionsofbSTweregiven bettermaintainedandconcentrationsofmetabolic betweenpostpartum42 nd dandd100±4postpartum, hormoneswereimproved. butallcowsreceivedfulldoseofPOSILAC®beginningat d100±4postpartum.Duringthesecondexperiment SummaryandConclusions (59)injectionsbeganapproximately21d(±3d)before expectedcalvingdatesandwerecontinueduptod42(± Wheninjectedduringongoinglactation,bSTresulted 2d)postpartum.After60d,allcowsonexperiment inanexceptionalincreaseinmilkproductionofdairy receivedthefulldoseofPOSILAC®biweekly(500mg/14 cows.Becauseofitspositiveeffectsonbloodglucose, d).Datafromboththefirstandsecondstudiessuggest lipidsandaminoacidconcentrationsandblood thatuseof10.2mgbST/dduringlateprepartumand circulation,bSTincreasesmilkyieldwithoutaffectingthe earlypostpartumperiodscausednoapparentnegative overallcompositionofthemilk.Thisincreasewouldbe effectsonthetreatedcows.AlthoughEBwasmeasured equaltothatnormallyachievedbyAIandgenetic onlyinthesecondstudy(59)duringfirst4wkof selectionover10-20yperiod.Somatotropinalsohasa postpartum,NEBwasnotgreaterininjectedcowsthanin pivotalroleinhomeorheticcontrolofmetabolismand uninjectedcowsineitherstudy(58,59).Thiswasan nutrientpartitioning(carbohydrates,lipids,proteins,and indirectconclusionforthefirststudysincetheincreasein minerals)ofthecowduringthetransitionperiod.The MYandBWandthechangesinBCSwereequalorbetter transitionperiodandtheearlylactationperiodwere ininjectedrelativetoinuninjectedcows.InjectionofbST consideredastimeperiodsthathavethepotentialto resultedinbetterrecoveryofBWandBCSduringearly enhancelactationalperformance.Increasing lactation,especiallyafterinjectionswerestoppedaround concentrationsofSThasapotentialtoaugmentthe d42postpartum(58).Duringthesecondstudy,low metabolicchangesthatfavorthemammaryglandand doseofbSTdidnotprovokeagreaterlossofBWor haveapositiveeffectonDMI.Inabroadview,bST decreaseinBCScomparedtountreatedcows.Cowsin affectsthesysteminapositivewayandmakesitpossible bothgroupsappearedequallycapableofreplenishing forcowstorespondbetterlaterinlactationwithout theirbodyreserveseventhoughallcowsstartedinjection apparentnegativeeffectsonhealth.Changesin ofafulldosebSTaroundpostpartum60th d;adailydose concentrationsofmetabolichormones,likelycoupled wasthreetimesgreaterthanthatinjectedbefored42. witheffectsonvariousorgans,suggeststronglya Bovinesomatotropintreatmentsdidnotadverselyor beneficialeffectofbSTduringthetransitionperiod.This positivelyaffecttherateofincreaseinDMIduringthe canbeinterpretedthatinjectionsofbSTmayhavea

577 UseofBovineSomatotropinintheManagementofTransitionDairyCows

potentialuseduringthepostpartumperiodandprobably Acknowledgments duringtheprepartumperiod,toimprovemetabolicstatus TheauthorsthankAkdenizUniversityforitssupport. andimproveoverallmilkyieldsduringearlylactation. AppreciationisextendedtoDr.H.H.Head.

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