VOL491963PSYCHOLOGYWRMILES841wasnotsetupasaplannedexperimentIthappenedbychanceatOrangeParkinmoderatelygooddaylightonSundayFebruary211954WiththelateDrHenryWNissenamemberofthisAcademywhowas ID: 507161
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CHIMPANZEEBEHAVIOR:REMOVALOFFOREIGNBODYFROMCOMPANION'SEYEBYWALTERR.MILESU.S.NAVALMEDICALRESEARCHLABORATORY,GROTON,CONNECTICUTReadbeforetheAcademy,April24,1963Theintelligenceandsocialbehaviorofapeshavebeenmuchinvestigatedintihelastfiftyyearsthroughorganizedfieldstudiesandespeciallybyexperimentationandcontrolledobservationofmanyanimalsconfinedinlaboratorystationsandzoologicalparks.StudiesconductedonchimpanzeesattheAnthropoidStationinTenerifeandtheYerkesLaboratoriesofPrimateBiologyinFlorida,asexamples,coverawidespectrumofbehavior.Workingforfoodrewards,chimpanzeeshaveamplydemonstratedtheirexcellentsensoryandperceptualcapacities,aswellastheirabilitiesinlearningandinsolvingavarietyofmechanicalproblems.Thesocialbehaviorofgroomingbyanthropoidsremainsasubjectoftheoreticalscientificinterest.Chimpanzeesthatarefriendlyoftenexhibitamutualandex-citeddesiretoengageingrooming.Smallwoundsorsoresfoundbythegroomermaybegiven"treatment"bythelips,orasplinterintheskinmayberemovedbypinchingtherestrictedareabetweenthetwoforefingers.Suchactsorbehaviorarenotnowaccreditedasresultingfrommotivesof"mutualaid."Thegroomer,oroperator,seemstoselecttheobjectivesandperformactionpatternsforhisownpleasure.Thebehaviorsequencereportedinthepresentpaperinvolvedafamouspairofchimpanzees.Ihadvisitedandobservedthispairmanytimesinpastyears.ProfessorRobertM.Yerkes,amemberofthisAcademy,purchasedPanandWendy,maleandfemale,fromaship'sofficerataBostondockin1925.Theirrespectiveagesatthetimeofpurchasewerenotknown,butwereestimatedattwotothreeyears.Panasanadultwasofmediumsize,weighingabout110pounds.Wendywassmaller,withaweightofabout88pounds.Describingthegeneralbe-haviorofthesechimpanzeesin1943,Dr.Yerkeswrote,"Althoughtheyhavelongbeenacquaintedascage-mates,theyhaveoftendisagreedabouttheirrelativesocialstatus.Eachisextremelyaggressiveanddetermined,andneitherhesitatestoprecipitateafight.Usually,Panassumespriorityashisright,forheisthatsortofpersonality.ButifWendymakesuphermindtohaveherway,sheissomuchmoredevastatinginherattackthathealmostinvariablyyieldstemporarily,inordertogetalong,peaceablyandsafely."'Atfirst,eitherofthesechimpswouldstruggledesperatelyifremovedfromtheotheroneandtakenfromtheirlivingcagetoanexperimentalroom.Itrequiredseveralattemptedsessionsbeforeeachbecameaccustomedtotheexperimentaloutlook.Bothlearnedtobegoodsubjectsandworkwellforfoodrewardsonavarietyofproblemsandindifferentlaboratoryenclosures.WhenYerkeswaswritingaboutthemin1943afternineteenyearsofacquaintance,hereferredtoPan'shistoryasabookof60typedpages.andtothatofWendy,themotherofsevenvaluable,goodworkingoffspring,asextendingto75pages.Nowacinderorgrainofsandintheeyeisapowerfulstimulusdemandingtheattentionofthesufferer,behemanorchimpanzee.Theincidentherereported840 VOL.49,1963PSYCHOLOGY:W.R.MILES841wasnotsetupasaplannedexperiment.IthappenedbychanceatOrangeParkinmoderatelygooddaylightonSunday,February21,1954.WiththelateDr.HenryW.Nissen,amemberofthisAcademy,whowasthenDirectoroftheYerklsLaboratories,thewriterNvasstanding15feetormorefromonesideofthelargeout-doorenclosurewhichconfinedPanandWendy,eachofthemaboutthirtyyearsofage.Panwasengagedinhisusualnoisy,belligerentdisplaytacticsforvisitors.2Hewouldclimbandpoundthewirefenceorthelargestripofsheetmetalatthetop.Hewasjumpingabout,stamping,banginghishandstogether,andshouting.Previousexperiencepromptedonetostandwellbackfromthecage,knowingthatPancouldthrowsandwithforceandaccuracy.InthemidstofPan'sexcitedbehavior,Wendycameupbehindhim,whimpering.Pansuddenlyturned,looked,andapproachedher.Shesatdownontheground;heimmediatelycrouchedinfrontofherandwithhistwoforefingersbeganmanipulatingherlefteyelids.Ashortsequenceofimpromptucolorphotographsrecordedthisbehaviorincidentanditspattern.Thisremarkableepisodebeganwithsuchabriefsettingofthestagethatitwasfaradvancedbeforewecouldreachthesideofthecageandthrustthecameralensbetweentheheavyfencewirestoachieveaclearview.Thefirstphotographtaken(Fig.1)showsPancrouchinginfrontofWendyandverynear,asheviewsFIG.1.-Anoldmalechimpanzeerespondspromptlyandinanadequatemannertothedistressofhiscage-mate,whohasacinderinherlefteye.Hehaspulleddownthelowereyelid,exposingthemucousmembrane.herlefteye.Heisusingbothhishandsforthetaskofsearching.WithhislefthandheappearstostabilizeWendy'shead.Withhisrightforefingerhehasdrawndownthelowereyelidexposingthepinkmucousmembrane.Wendy'sheadistippedtowardPan,whoisviewinghereyefromadistancethatweestimateas5to6inches.Thesecondphotograph(Fig.2)wastakenabriefintervalafterthefirstone.Thesearchisjustabouttoconclude.Wendyhasdrawnherheadbacksomewhat.Panstillretainsverynearlyhissamestanceofheadandshoulders,but 842PSYCHOLOGY:W.R.MILESPROC.N.A.S._~~~~~~~~~~~~~~~~~~~~~~~~i...:,...............................i~~~~~~u~ljIf.........._...'Fi-(~~~~~_92'TescacE.11tlMS1dIs_'list'11I(IucehisrighthandreachesoutfurtherasiftotouchWendy'sleftcheek.WecanseethesmallthumbofPan'srighthand.ThefocaldistancebetweenPan'seyesandWendy'slefteyehasincreased.ThemomentwhenPanwithdrewhisfingersfromWendy'sface,signalingsuccess'ofhisattempt,occurredwithsuchsuddennessitwasnotcapturedbyaphotograph.AsWendyexperiencedrelief,therewasaquickchangeinherposturefromsittingtolyingdown.Panturnedsothat.shewasinfroaatofhim,whileheatthesametimecouldagainseehistwohumanobservers.HeglancedbrieflyatWendy'sgenitalareaandthenassumedtherelaxedsittingpostureimmediatelybehindWendy.Shewaslyingontheground,herheadrest-ingonsomestrawandherrightarmlimpandrestingonherbody.Panseemedtolookathishumanobserverswithamuchalteredperceptualattitude.Hewasnotthreatening.Hesatquietlyrestingonhisrumpandrightthighwithhisleftlegflexedandhisrightforearmrestingonhisknee.Le&aningforward,hewaswellbalancedand,judgingfromtheflexionofhisleftwrist,quiterelaxed.ThecameraviewforFigures1and2wasfromthesamepositionthroughtheheavywirefence.BeforerecordingthephotographforFigure3,thecamerapositionwasloweredabouttwofeet.TheperspectivebetweenPan'sheadandthebackgroundhasthusbeenchanged.Pan'spostureasshowninFigure3hasremainedthesameforaboutfiveminutes.Justasthisfinalphotographwastaken,WendyliftsherheadtolookatPan.Itheplottingsomemischief,orishequietlyenjoyingasenseofachieve-ment?Thereisnoclearanswer.Pancouldhaveusedhismercyactasthepreludeforanenjoyableperiodofgrooming.Wendycouldhavewithdrawnimmediatelyafterexperiencingrelief.Theirremaininginphysicalcontactinsuchpassivestatesforseveralminutesaftereye-reliefwasachievedwouldseemtoemphasizethepsychologicalsignificanceofthisbehavioralevent.Itisdistinctlydifferentfromgrooming.Itwassomethingspecial.Manhasbecomeorisbecomingwisetothevalueofsuccorantbehavior.'Hedidnotinventit;therewereearlierpioneers. VOL.49,1963BIOCHEMISTRY:BALTIMOREETAL.843FIG.3.-Comforthasbeenachieved,andoperatorandpatientbothrelax.'Yerkes,RobertM.,Chimpanzee8ALaboratoryColony(NewHaven:YaleUmiversityPress1943),pp.47-48.2Fmlen,JohnT.,"Thedisplayofthegorilla,Proc.Amer.Philos.Soc.,106,516-519(1962).Scott,JohnP.,AnimalBehavior(U~niversityofChicagoPress,1958),pp.xiand281.POLIOVIRUS-INDUCEDRNAPOLYMERASEANDTHEEFFECTSOFVIRUS-SPECIFICINHIBITORSONITSPRODUCTION*ByDAVIDBALTIMORE,HANSJ.EGGERS,RICHARDM.FRANILIN,tANDIGORTAMMTHEROCKEFELLERINSTITUTECommunicatedbyFrankL.Horsfall,Jr.,April19,1963Inthelastfewyearsnewexperimentalapproacheshavebeenfoundtothestudyofvirus-specificbiosynthesisinanimal-virusreproduction.Thisreportdescribesrecentresultsobtainedinthestudyofthemechanismofactionoftwovirus-specificinhibitors1ii6andprovidesevidencewhichbearsonthevirus-specificnatureofthevirus-inducedRNApolymerase.i17-202-(a-Hydroxybenzyl)-benzimnidazole(HB'-,andguanidine4'10-16inhibitcompletelyandspecificallythemultiplicationofmanysmall,RNA-containing,lipid-freeanimalviruses(picornaviniseS21).Thesecompoundshavenoeffectoronlyminoreffectsonthemultiplicationofviruseswhichbelongtoothermajorgroups.1'2,10Similarly,themetabolicactivitiesandgrowthrateofhostcellsareunaffectedoroniyminimallyaffectedbyHBB,Ioguanidine.12,13,16Mutantsof