11UndesiredBehaviourintheDomesticCat

Introduction

Inthischapterweexaminebehaviourthatcatownersmayfindproblematicorundesirable.Suchbehaviourhasanimpactonthecat–ownerbond,canresultinrelinquishmentoreveneuthanasiaofcatsandcanalsoindicatecompromisedwelfareinthecatsthemselves.Despitethis,relativelylittleresearchhasfocusedontheepidemiology,aetiology,preventionortreatmentofsuchbehaviourincats,andmuchofthepublishedliteratureisanecdotal,basedonindividualcasereportsorderivedfromopinion-basedsources.

Althoughtermssuchas‘behaviourproblem’and‘undesiredbehaviour’arewidelyusedintheliterature,itisdifficulttofindacleardefinitionofwhatsuchtermsmean,oraconsistentideaofwhichtypesorpresentationsofbehaviourareincluded.Sincea‘behaviourproblem’mustbeabehaviourthatanownerfinds‘problematic’,thiswillclearlyvarywitheachindividualowner’ssubjectiveinterpretationofwhatisacceptable.Forexample,whereasoneownermayfindintermittentaggressivebehaviourbetweencatsinahouseholdacceptable,anothermayinterpretthisbehaviourasaproblem.Indeed,furthercomplicationsforsuchdefinitionsoccurwhereownersinterpretcommonbehaviouralsignsshownbytheircatsindifferentways:forexamplethesamebehaviourmaybeperceivedas‘aggression’bysomebutregardedas‘play’byothers.Ourfirstprobleminunderstandingundesiredbehaviour,therefore,istoconsiderhowsuchbehaviourmaybeclassified,andtheextenttowhichownerperceptionof‘undesirability’isanimportantconsiderationinsuchdefinitions.

Theotherpotentiallyperplexingaspectofdefiningundesiredbehaviouristheextenttowhichsuchbehaviourisconsideredtoliewithinthenormalrepertoireofthecat.Indeed,behaviourproblemsaresometimesdividedintheliteratureintothosethatarisefrombehaviourthatisessentially‘normal’or‘adaptive’forthespecies,andthosethatappeartobe‘abnormal’or‘maladaptive’(BorcheltandVoith,1982).Theformerincludesthoseinstanceswherebehaviourislikelytobeshownbycatsinamannercompatiblewiththatseenina‘natural’environment,orwhichmayoccurinothercontextsconsequenttomodulationby

theprocessesofassociativelearning.Foradomesticatedspecies,suchasthecat,itisperhapsdifficulttodefinewhata‘normal’environmentmaybe,andinpracticecomparisonsaregenerallymadewithcatsinafree-rangingorferalenvironment.Abnormalbehavioursaregenerallyconsideredtoincludethosewithaclearlymedicalorigin,orthosewherethebehaviouralsignsdisplayedarenotgenerallyapparentinthatspecies.

Differentiating‘normal’or‘abnormal’behaviour,however,isnotasclearasmightbeexpected.Somepresentationsdofallclearlywithinoneorotherofthesecategories:acatthaturinespraysinresponsetoenforcedclosecontactwithanothercatthatisnotpartofitssocialgroupcanbedescribedasshowinganadaptiveresponse,eventhoughthismaybeundesirabletoanowner.Equally,anextremebehaviouralresponsethatoccursentirelyindependentlyofenvironmentalstimuli,forexamplefromapartialseizurefocuswithinthelimbicsystem,isclearly‘abnormal’.However,therearemanycasesthatfallbetweenthesetwoextremes.Manyanimalsshowspecies-specificbehaviourinresponsetoanaversiveevent,butmayeitherappeartoreactatanunexpectedlylowthreshold,showahigherdegreeofresponsethanwouldnormallybeexpectedatfirstexposureorgeneralizeabehaviourbetweencontextsmorerapidlythanonemightexpectthroughlearning.Theremaybemanyreasonsfortheseapparently‘abnormal’characteristicsof‘normal’behaviours,including:(i)medicalconditionsthatinfluencethethresholdofresponses(seeChapter12,thisvolume);(ii)otherenvironmentalcausesofarousaloccurringconcomitantly;(iii)geneticfactorssuchaspersonalitycharacteristics;(iv)developmentalcausesofincreasedreactivity;(v)epigeneticchangescausedbychronicstress;or(vi)awholehostofotherfactors.Othercasesmightpresentwithbehaviourthatappearsonfirstexaminationtobecompletelyabnormal,butforwhichaclearhistoryemerges,revealinghowthecathaslearntthebehaviourthroughaseriesofuniqueexperiencesovertime.Itisthecomplexinterweavingofgenetic,developmental,environmentalandinternalfactorsthatcanmaketheinterpretationofindividualbehaviourcasesbothfascinatingandchallenging.Theinvolvementofmultiple,interactingfactorsineachcasealsomeansthatthosewhoadviseownersonthetreatmentandmanagementforsuchcasesshouldhavebothdemonstrableknowledgeandclinicalskillinfelinebehaviourbeforeattemptingtotreatcases.Theoverlappingrolesoflearningandmedicaldiseasealsomeanthatnon-veterinarycliniciansshouldworkcloselywithmembersoftheveterinaryprofession.

ProportionofCatsShowingUndesiredBehaviours

ProportionofCatsShowingUndesiredBehaviours

Thereislimitedevidenceastotheproportionofdomesticcatsshowingundesiredbehaviour.However,thenumberofcatsreferredtospecialistcentresforbehaviouraladviceissuggestedtobeincreasing:whetherthischangeisasaresultofincreasedownerawarenessthatbehaviourtherapyisanoptionforcats,orduetochangesintheactualnumberofbehaviourproblemsthataredeveloping,isunclear(Heath,2005).Theformerispossible,sincecatsappeartobebecomingincreasinglypopularpets,witharecentsurveyestimatingover10.5millioncatsintheUK(Murrayetal.,2010).Inaddition,theroleofthecatinhouseholdsalsoappearstohavealteredovertime,fromonewherethecatwasrathermoreperipheraltothefamily,possiblykeptforrodentcontrol,toonewherethecatplaysanimportantsocialrolewithinthefamilyandcanprovideconsiderableemotionalsupportforsomeowners(BradshawandLimond,1997).Thismayresultinownersinvestingmoretimeandfinancialresourceinaddressingundesiredbehaviours.

Thesesamefactors,however,mayalsoaccountforanincreasedproportionofcatsdevelopingundesiredbehaviour.Theincreasingnumberofcatsinevitablyleadstohigherlocalpopulationdensities.Thestressthatmanycatsexperiencewhenlivingincloseproximitywithunfamiliarcats(seeChapter8,thisvolume),particularlywherelocalpopulationsareinflux,mayaccountforincreasednumbersofundesiredbehaviours.Morecatsarealsokeptinmulti-cathouseholds,withMurrayetal.(2010)reportingthat42%ofcatownershadtwoormorecats,and2%havingbetweensixand12cats;suchnumbersarelikelytogenerateproblemsassociatedwithsocialstress,asdiscussedfurtherbelow.Thechangingroleofcatswithinfamiliesmayalsobeanimportantfactorinincreasedbehaviourproblems:theinclinationofownerstoshowintenseandcloseboutsofinteractionwithcatscanbeatoddswiththenaturalsocialbehaviourofcats,andtheresultingmiscommunicationmayleadtothedevelopmentofbehavioursthatownersfindundesirable.

TheAssociationofPetBehaviourCounsellors(APBC)intheUKproducesanannualreviewofdataaboutthecasesthathavebeenreferredtomembersfromveterinarysurgeons.Sinceratherfewmembersofthisorganizationspecializeinfelinebehaviour,thedataarelimitedintheirapplication,buttheydoprovidesomeinsightintothetypesofbehavioursforwhichcatownersseekexperthelp.Inthe2005reviewofcases,informationwascollatedon65felinecases,ofwhich30weremaleand35female.Thepresentingproblemfor28%ofthecatswasindoormarking(includingurinespraying,squatmarking,middeningandscratching)and11%presentedwithinappropriatetoileting

problems;22%displayedaggressiontoothercats,ofwhich86%wasdirectedtowardsothercatswithinthehousehold;11%showedsomekindoffearfulor‘phobic’response;10%ofcasespresentedwithaggressiontowardspeople,7%withbehaviourscausedbymedicaldisordersand5%with‘bondingproblems’(presumablymeaningcatsshowingalimitedtoleranceofinteractionwithpeople)(APBC,2005).Datacollectedovera6-monthperiodinareferralclinic(Fig.11.1)similarlyshowedthaturinespraying,inappropriateelimination,aggressionbetweenhouseholdcatsandaggressiondirectedtowardsownerswerethemostcommonreasonsforwhichownerssoughthelp.

Fig.11.1.Numbersofcats,bygender,referredtoaspecialistcentreovera6-monthperiodin2000(N=61).

Surprisinglylittleresearchhasbeendonetoinvestigatetheprevalenceofbehaviourproblemsinthegeneralcatpopulation.Bradshawetal.(2000a)conductedadoor-to-doorsurveyof90householdsintwoareasoftheUK,oneruralandtheothersuburban.Answersfrom15maleand75femalerespondentswereacquiredregardingallcatspresentinthehouseholds,atotalof161cats.AsshowninFig.11.2,notonlywasthereasurprisinglyhighnumberofundesiredbehavioursdisplayedbycatsinthisstudy,butthefrequencyofthevarioustypesofproblemwasverydifferentfromthatdescribedaboveinareferredpopulation.Asurveyof109ownersvisitingafirst-opinionveterinarypracticesimilarlyshowedahigherproportionofbehavioursuchasavoidanceofother

cats,scratchingandavoidanceofvisitors,comparedwiththecommonproblemsseenonreferral(Fig.11.3).

Thedifferencebetweennumbersandtypesofbehaviourproblemsinthereferralpopulationandinthegeneralpopulationsurveyssuggestthatthecatsseenbyreferralpractitionersarenotonlythe‘tipoftheiceberg’intermsofnumbersofcatswithbehaviourproblems,butalsothatownerstendtoseekhelpforparticulartypesofbehaviour.Theseappeartobethosebehavioursthataremorelikelytoimpingeonthelifestyleorenvironmentofowners,suchasurinespraying,inappropriateurinationandaggressiontowardspeopleorothercatsinthehousehold.Confirmingthis,ageneralprevalencesurveyfoundahighproportionofproblemsthathadlessimpactonowners,suchasavoidanceofunfamiliarpeopleorothercats(CaseyandBradshaw,2001).

Otherevidencefortheextenttowhichundesiredbehavioursincatsinfluencethecat–ownerbondcomesfromreasonsforrelinquishmentofcatstorescueorganizations.Thereisconsiderableinconsistencyintheproportionofcatsreportedtoberelinquishedforapparentbehaviourproblemsinthesestudies,rangingfromabout8%toover33%,probablyinfluencednotonlybymethodologicaldifferencesindatacollectionbutalsobyownerperceptionofwhatmaybean‘acceptable’reasontoabandonacat.IntheUK,overa12-monthperiod,only7%ofcatswererelinquishedtoCatsProtectionprimarilyduetoundesiredbehaviours(accordingtoownerreports),althoughamuchhigherproportion(38%)werereturnedtocentresafterhomingbecauseofundesiredbehaviour(Caseyetal.,2009).

Fig.11.2.Prevalenceof‘behaviourproblems’inthegeneralcatpopulation(fromBradshawetal.,2000a).

Fig.11.3.Percentageofcatsshowingarangeofundesiredbehavioursinasurveyofcatownersvisitingafirst-opinionveterinarypractice(N=109)(fromR.Casey,unpublisheddata).

ClassificationofUndesiredBehaviour

Consistentdescriptionsandclassificationsofundesiredbehavioursareessentialforthoseworkingincatbehaviourresearchorclinicalpractice.However,classificationofundesiredbehaviourshastakenanumberofdifferentapproachesintheliterature.Forexample,behaviourproblemsincatshavebeendefinedaccordingto:(i)whatformthebehaviourtakes(e.g.‘excessivevocalization’);(ii)thecontextinwhichthebehaviouroccurs(e.g.‘territorialaggression’);(iii)thetargetforthebehaviour(e.g.‘intraspecificaggression’);or(iv)thelikelymotivationofthebehaviour(e.g.‘redirectedaggression’).Alloftheseapproacheshavepotentialdrawbacks.Forexample,describinganobservedbehaviour,contextortargetgivesnoindicationastopotentialmotivation.Thiscanbemisleading,sinceabehaviouralpresentationcanariseviaanumberofdifferentroutes.Forexample,anexcessivelyvocalizingcatcouldbeshowingthisbehaviourforoneormoreofseveralreasons,includinggainingownerattention,inresponsetoanxiety,orasaconsequenceofhyperthyroidismorage-relatedchanges.Adiagnosticcategorycombiningthesetogethercouldleadto

relatedchanges.Adiagnosticcategorycombiningthesetogethercouldleadtoprescriptiveapproachestotreatmentthatareunlikelytobeappropriateforallcases.

Themaindrawbackofdefiningbehaviouraccordingtomotivationisthatthisapproachrequiresadegreeofsubjectiveinterpretation,gleanedfromhistorical,observationalandcontextualinformation.Becauseoftheinterpretationalnatureofthistypeofclassification,variationmayoccurbetweenauthors,forexamplerelatingtodifferencesinpreferredtheoriesofaetiologywithindifferentcultures.Withthecurrentlimitedknowledgeoftheaetiologyofbehaviouralconditions,thesediagnosesmustberegardedashypotheticalconstructsthatshouldbesubjecttomodificationanddevelopmentasscienceprogressesandknowledgeincreases(SheppardandMills,2003).Onesalientexampleofdifficultiesarisingfromthistypeofclassificationsystemisthedifferencesofopinionintheuseofahierarchicalframeworkwheninterpretinginteractionsbetweengroupsofcats(seeChapter8,thisvolume).Motivationalcategoriesofbehaviourare,however,likelytobemorecloselyrelatedtotreatmentprotocols.

FactorsInfluencingDevelopmentofUndesiredBehaviours

Althoughundesiredbehavioursareoftenconsideredseparately,theprinciplesunderlyingthedevelopmentofsuchbehavioursarenodifferentfromthoseinfluencingbehavioursthatareacceptableforowners.Ininterpretingundesiredbehaviouritisthereforeimportanttoconsiderthenormalbehaviourorethologyofcats,togetherwiththeindividuallearningopportunitiesthathaveinfluencedthedevelopmentofspecificresponsesinparticularcircumstances.Mostoftheundesiredbehavioursthatwediscussinthissectionareessentially‘normal’responsesofmembersofthisspeciestotheenvironmentinwhichtheyfindthemselves.However,ownersoftenseekhelpbecausebehavioursareincompatiblewiththeirlifestyles,areelicitedbyinappropriatestimuli,occurinaninappropriatecontextorareshownatsuchintensitythattheybecomeanuisance.Inmanycases,suchbehaviourhasstartedasanormalresponseandbecomesinadvertentlyreinforcedtothepointthatitbecomesunacceptable.Manybehaviourproblemsalsoariseasaresultofanowner’slackofunderstandingofthenaturalbehaviouroftheirpet.Forexample,manyownersdonotappreciatetheamountofmentalandphysicalstimulationthatcatsrequire,particularlywhenhousedinanindoorenvironment,andtheoccurrence

ofmanybehaviourproblemsisatleastpartlyinfluencedbytheirpet’slackofopportunitytoshownormalspecies-specificbehaviour.Apartfromphysiologicalandpathologicalfactors,whichareaddressedseparatelyinChapter12,thisvolume,themainfactorsinfluencingthedevelopmentofundesiredbehavioursincatsare:(i)thelimitationsofintraspecificsocialbehaviourinthespecies;(ii)relativedifferencesinsocializationexperience;(iii)breedandindividualdifferences;(iv)abilitytoshownaturalbehaviouralresponses;and(v)individuallearntexperiences.Theseareseparatelyaddressedinthesectionsbelow.

Socialinteractionbetweencats

Theuniqueoriginofthedomesticcat,derivedfromtheessentiallyasocialFeliss.lybica,andsubsequentlyadaptedtoliveinsocialgroups,hasimportantimpactsontheabilityofF.catustolivesuccessfullyinthedomesticenvironment.Theirasocialancestryhasresultedinamuchmorelimitedvisualsignallingrepertoirethanthatofspeciesderivedfromcooperativehunters,suchasthedomesticdog(seeChapter5,thisvolume).Complexvisualsignallingevolvedasanimportantelementinenablinggroupcooperationinnaturallysocialspecies,butwasnotanecessityforF.s.lybicawhereadultindividualsmaintainedadistancethatobviatedtheneedforsignalssuitableforface-to-faceencounters.Whereasfortheancestorsofthedomesticdog,socialsuccessdependedontheabilityofgroupmemberstodisplaychangesinemotionalstatesuchthatothergroupmemberscouldadapttheirbehaviouraccordingly,thisabilitywasnotimportantintheevolutionofthecat.

Thelimitationsofvisualsignallingincatsappeartohavebeenafactorinrestrictingthetypesofsocialgroupingstowhichthespeciescansuccessfullyadapt.Feralorfarmcoloniesaremadeupofcooperativesocialgroupsoffemales,basedontheconcentrationoffoodresources,andprovidingadvantagesofcooperativerearingofkittens(seeChapter8,thisvolume).Thesegroupsworkwell,despitetherelativelypoorabilityofcatstodisplaychangesinemotionalstateusingsubtlevisualsignals,fortwobroadreasons.Thefirstisthatcompetitionbetweenindividualsisminimizedbythegroupsizecloselymatchingresourceavailability,andhuntingbehaviourremainingasolitaryexercise.Thesecondisthatmembersofsocialgroupsarepredominantlyfamilial.Individualsareveryfamiliarwitheachother,oftendevelopingtogethersincebirth.Furthermore,socialbondsarerepeatedlyreinforcedwithaffiliativetactilebehavioursandtheswappingofscentsignalsthroughrubbingand

grooming.Thisfamiliarityenablesgroupmembersto‘know’eachotherwell,inthesensethattheycaneffectivelypredicthowothergroupmembersarelikelytorespondindifferentcircumstances,makingtheabilitytojudgeresponsesthroughinterpretationofvisualsignalslessimportant.Encounterswithcatsfromothercoloniesareunlikelytoresultinaffiliativeresponses;rather,suchinteractionsgenerallyresultinstrategiesthatavoidanycontactatall,orinvolveovertsignsofaggression.Studiesofthesenaturalgroupingoffree-rangingcatsthereforesuggestthattheresponseofcatstoconspecificsishighlydependentontheirrelativeperceptionofothersasa‘groupmember’or‘non-groupmember’.

Giventhelimitationsofsocialcompatibilityforcats,itisperhapsunsurprisingthatthediversityofsocialsituationsinwhichpetcatsfindthemselvesisthemostcommoncauseofundesiredbehaviours.Inareasofhighhumanpopulationdensity,catsmayliveincloseproximitywithanumberofcatswhichtheydonotperceiveaspartofthesamesocialgroup.Thiscanleadtohighlevelsofvigilanceandanxietyascatsattempttooccupyhomerangeswhileatthesametimeavoidingcontactwithothercats.Predictingthetemporalandspatialactivityofothercats,forexamplethroughthemonitoringofscentsignals,cansometimesenablecatstooccupyoverlappingterritoriesbutnotcomeintocontact.However,wherealteredpatternsofactivityarecausedbynewcatsarrivinginanarea,owneractivitiessuchasholidaysoralteringcats’accessoutside,thiscanhaveanimpactoncatsacrossaconsiderablearea.Thegreatestimpactisprobablycausedbycatsthatmaintainalargeterritorysize(e.g.unneuteredorlate-neuteredtoms),sincetheywillcomeintocontactwithothercatsoverawidearea.

Behavioursarisingfromconflictbetweenneighbourhoodcats

Conflictbetweencatsinaneighbourhoodcanhaveanumberofconsequencesforthebehaviourofindividuals.Mostobviousistheoccurrenceofaggressionbetweencats:althoughownersmayhearevidenceofcatsfighting,thefirstsignofaproblemisoftentheabscessesoccurringasaresultofcatbites.Thelimitedabilityofcatstoshowappeasementbehaviourmeansthatencountersbetweencatsoftenresultinovertaggression,chasingandbiting.However,otherbehaviouralconsequencesarealsocommon:catsthatareanxiousaboutcontactwithothercatscannotpredicttheactivityofothercats;catsthathavehadpreviousnegativeencounterswithothercatsmayaltertheirpatternsofoutdooractivity.Thiscanconsistofreducingtimeoutside,theavoidanceofparticularareasorreducedoverallareaofrange.Theconsequencesofsuchalterationswill

varywithotherfactors,butcaninclude,forexample,increasedamountsoftimeseekinginteractionwithowners.Suchcatsmayseekattentionthroughexcessivevocalization,orshowabnormalplay/predatoryresponsestowardsowners.Alternatively,avoidanceofcontactwithneighbourhoodcatsmayresultinsomecatsnolongerleavingthesafetyofthehome,oronlydashingoutforshortperiods.Catsthatareanxiousabouttheactivityofothercatsincloseproximitytothehomemayspendconsiderableamountsoftimeshowingvigilantbehaviour,suchas‘checking’outofwindows,orlookingoutthroughclearcatdoors.Theymayonlyventureoutsidewhenownersgointothegarden,orprefertoleavethehousethroughanopendoororwindowtoavoidtheuncertaintyassociatedwithgoingoutsidethroughacatdoorwithlimitedvisualaccesstotheoutsidearea.

Reluctancetoleavethesafetyofthehouseduetothepresenceofothercatscanalsoleadtothedevelopmentofinappropriateelimination.Whereacat’snormaltoiletinglocationisoutside,reluctancetoleavethehouseisacommonreasonforseekinganalternativetoiletinglocationinside.Catsarevulnerablewhentoileting,andinsecurityaboutaccesstoausualtoiletingsiteisonereasonforcatstoaltertheirtoiletinglocation(Table11.1).Hence,ifacatnormallytoiletsoutside,anxietyaboutneighbourhoodcatswillcommonlyleadtoashifttotoiletinginside;wherealittertrayisnotprovidedthiswillbeonothersurfaces,andperceivedasundesirablebyowners.

Table11.1.Characteristicsofsuitabletoiletingsitesforcats.

Featuresofasuitabletoiletingsiteforacat

SuitablematerialforburyingQuietlocationhiddenawayLocationawayfromthethreatofothercatsthatarenotperceivedaspartofthesamesocialgroupSomecatspreferaseparatelocationfordefecationandurination

Littermatchesthecat’ssubstratepreferenceLocationawayfromfeedingsite,butnotsofarfromthecoreareaastofeelunsafe

Urinesprayingisanotherconsequenceofsocialincompatibilitybetweenneighbourhoodcats.AsdiscussedinChapter5,urinesprayingisanolfactorysignalthatisconsideredtoserveanumberoffunctions.Inentirecats,thesesignalsappeartohaveasexualsignallingfunction.However,urinesprayingalso

commonlyoccursinneutereddomesticcats.Althoughhistoricallyoftendescribedasa‘territorial’behaviour,urinesprayingrarelyoccursattheperipheryofacat’srange.Rather,spraymarksusuallyoccurinlocationsthatare‘sociallysignificant’(Herron,2010).Forexample,themostcommonlysprayeditemsincludefurnitureandwallsorwindowsneartovisualaccessoutdoors(Pryoretal.,2001),whichareareaswhereconflictwithothercatsislikelytooccur,orwheretheactivityofothercatsmaybeobserved.Althoughcats‘check’boththeirownscentmarksandthoseofothers,theyrarely‘over-mark’thescentofothercatswiththeirown(Hart,1974),alsosuggestingthatthisbehaviourisnotpurelyterritorial.Therateof‘checking’tendstodeclinewiththeageofthescentmark,suchthatfreshurineismostinterestingtocats,andinterestdecreasesasthescentfades(DeBoer,1977b).Astimesincedepositionappearstobeimportanttocats,itseemslikelythatoneimportantfunctionofthesemarksistoenablecatstoavoideachotherwhentheyco-occupythesamearea(Cooper,1997)inordertoavoidovertaggression.

Sincecatswillurinesprayinsocialisolation,itisalsopossiblethatthesemarkshaveafunctioninprovidinginformationtothemarkingcatitself.Leavingamarkinlocationsassociatedwithperceivedthreatmayprovidetheanimalwithagreaterdegreeofpredictabilityandcontroloveritsenvironment,asitenablesitto‘know’areaswherethereispotentialdanger,andvigilanceisneeded.Hence,aspraymarkonthecatflapwouldalertacattobecautiousinthatareaifconflictwithothercatshadpreviouslyoccurredatthatlocation.

Catsthathavetheiractivityinhibitedbyothercatsintheneighbourhoodmayalsoshowotherbehaviouralsignsassociatedwithstress,suchasover-grooming.Thisisparticularlylikelytobethecasewhereothercatsactuallyenterthehouse,ortheenvironmentissuchthattheresidentcatcannotpredictoravoidencounterswithothercatsperceivedasthreatening.

Behavioursoccurringinmulti-cathouseholds

Wheremultiplecatsliveinthesamehousehold,similarissuesariseamongthemasthosecausedbysocialincompatibilitybetweencatsinaneighbourhood.Whetherundesiredbehavioursariseinmulti-cathouseholdsdependsontheextenttowhichcatsperceiveeachothertobemembersofthesamesocialgroup,andalsoontheextenttowhichtheycanaccessresourcesindependently.Aswithfree-rangingcats,individualsinsocialgroupsgenerallyshowhighlevelsofaffiliativebehaviours(suchasallogroomingandallorubbing)andwillchoosetobeincloseproximitytoeachother,forexamplesleepingincontactwithone

another(Table11.2).Alsoconsistentwithfree-rangingcats,individualsthataresiblingsorthathavedevelopedtogetheraskittensaremorelikelytoformsocialbondsthancatsintroducedasadults(seeChapter8,thisvolume).

Wheretwoormorecatswithinahouseholdformseparatesocialgroups,theywillgenerallyestablishdifferentcoreareasindifferentpartsofthehouse,andoftenessentiallyliveseparatelives.Thismaybeachievedbyactiveaggressionorwithdrawalonencounteringtheothercat,butmoreoftenoccursthroughestablishingroutineswherebyactualencountersareminimized(Table11.2).Insuchcases,evenwherethereisnoactiveaggression,theactivityofeachcatmaybeinhibitedbytheothers.Forexample,onecatinthehouseholdmaybeunwillingtopassthroughadoorwaywhenanothercatisnearby,oritmaynotenterthroughacatflapwhentheothercatisinthevicinity.Ownerscanoftenbeoblivioustothesepatternsofavoidance.Thisisparticularlythecasewhereincompatiblecatsoccupythesamespaceinordertoaccessessentialresources.Forexample,ownerswilloftenprovidefoodforallcatsinahouseholdinthesameroomatthesametime.Inordertoobtainsufficientnutrition,catsmayhavetoeatclosetoothercats,althoughtheywouldnotchoosetodosoinothercircumstances.Often,signsofanxietyshownbycatswhenforcedtoeatincloseproximityareoftennotappreciatedbyowners.Thismayincludeindividualcatsboltingfoodveryfast,oreatingexcessiveamountsatatimetoavoidthenecessityofreturningfrequentlytothefeedingarea,factorspotentiallyimportantinthedevelopmentofobesity.Otherexamplesofunwillingproximitymayoccurwherecatsareattractedtolimitedsourcesofheat,orwheremultiplecatsinahouseholdvaluehumancontactandmaysiteithersideofanownerinordertoachieveattention,whilealsoattemptingtoavoiddirectcontactwitheachother.

Table11.2.Behaviouralsignsshownbycatsinmulti-cathouseholdsindicativeofsocialgroupings.

Theproblemsforcatsoflivinginahouseholdwithothercatsnotperceivedtobeinthesamesocialgroupareoftenexacerbatedbytheirowners’tendencytoclusterotherimportantresourcestogether.Inadditiontofeedingcatstogether,ownersoftenprovidelittertrays,waterbowlsandentrance/exitpointstotheoutsideinsinglelocations.Thecombinationoflimitedresourceaccessandavoidingothercatsinthehouseholdcommonlyleadstoarangeofundesiredbehaviours.Forexample,acatthathasacoreareaupstairsinahouseholdwillhaverestrictedaccesstoimportantresourcesifthesearealldownstairs,inthecoreareaofanothercat.Onecommonconsequencemightbethe‘upstairscat’startingtoeliminateupstairsifaccesstoadownstairstoiletinglocationbecomeslimited.Insomecasesthetoiletingbehaviourisprecipitatedbysomeotherchange,suchasachangeintheroutineofthe‘downstairscat’.Forexample,the‘upstairscat’maylearnthatitissafetogodownstairstousethetoiletfirstthinginthemorningandlastthingatnight,becausethesearetimeswhenthe

‘downstairscat’isouthunting.However,ifanothercatmovesintotheneighbourhoodandinhibitstheoutsideactivitiesofthe‘downstairscat’,thismayinduceinappropriatetoiletinginthecatlivingupstairs.Equally,ifthecatlivingupstairsisintheearlystagesofhyperthyroidism,itwillstarttodrinkmore,andhenceneedtourinatemorethantheprevioustwicedaily,leadingtotheonsetofinappropriateeliminationupstairs.

Aswithconflictbetweencatsintheneighbourhood,anxietyarisingduetoincompatiblecatslivinginthesamehousecanleadtovariousotherundesiredbehaviours.Onecommonconsequenceistheoccurrenceofurinespraying,withurinemarkslocatedinsignificantareassuchaswherecatsneedtopasseachotherinnarrowpassagewaysinordertoaccessimportantresources.Asdescribedforneighbouringcats,thesesignalsmayhavebothacommunicativefunctiontoavoiddirectconfrontationandenablemarkingcatsthemselvestosubsequentlyidentifyareaswherepotentialconflictmayoccur.

Behaviouralsignsofchronicstressalsooccur.Thesemayincludecatsspendingprolongedperiodsoftimehidingtoavoidcontactwithothercats,orshowingabnormalresponsessuchasovergrooming.Inaddition,somecatsmoveawayfromhouseholdswherecontactwithothercatsisstressful,forexamplepredominantlylivingoutsideormovingtoliveinotherhouseholds.

Becausecatsdonotappeartobemotivatedtomaintainsocialbondsinthesamewayasotherspecies,thebreakdownofestablishedrelationshipsoftenresultsinapermanentsplitofcatsintoseparatesocialgroups.Sincetherecognitionofotherindividualsasgroupmembersislikelytobeatleastpartiallythroughtheirodourprofiles,maintainedbyaffiliativerubbingandgrooming,situationsthatalterthescentofagroupmembercanresultinthebreakingofsocialbonds.Shouldonecatfromasocialgroupleaveahouseholdforaperiod,forexampleifitishospitalized,evensiblingsmayfailtorecognizethatcatonitsreturn.Inextremecases,thiscanleadtothecompletebreakdownoftherelationship,becausetheinitialaggressiveresponseleadstoareciprocalresponsefromthehome-comer.Itisthereforesensibleforcatownerstorubcatsreturningfromatriptothevetswithatowelorpieceofbeddingthatiscoveredinthe‘groupscent’,priortoreintroduction,ortoactively‘swapscent’bystrokingonecatandthentheother(Crowell-Davisetal.,1997).

Relativesocializationexperience

TheimportanceofearlysensitiveperiodsforlearningaboutbothconspecificsandpeoplehasbeendiscussedinChapters4and9,respectively.Experiences

duringtheweeksimmediatelyafterakittenisbornappeartohaveaprofoundeffectontheoccurrenceoffear-associatedbehaviourslaterinlife(CaseyandBradshaw,2008).Mostoftheevidenceaboutsocializationrelatestotheextenttowhichkittensneedtoexperiencepeopleinordertoacceptcontactwithhumansasadults.Wherecontactwithpeoplebefore8weeksofagehasbeenlimitedorrestrictedtopeopleofaparticulargenderoragecategory,anxietyaboutpeopleismorelikelytooccursubsequently.

Probablyoneofthemostcommonbehaviouralconsequencesoflimitedsocializationexperienceistheavoidanceofunfamiliarpeople.Althoughrarelyseeninthereferralbehaviourclinic,forreasonsdiscussedearlier,avoidingcontactwithvisitorsappearstobeparticularlyprevalentinthecatpopulation.Manycatsarereportedto‘disappear’whenvisitorscometothehouse,butveryfewownersperceivethistobeaproblem.Catsthatarefearfuloftheirownersarepresentedasclinicalcasesmorefrequently.Theseareoftencatsthathavehadlittleornoexperienceofhumansintheirearlysensitiveperiodforlearningbecause,forexample,theywereferalorfarmkittens.

Catsmayalsodisplayaggressivebehaviourtowardspeopleasaresultoflimitedsocialization.Aggressionoccursasanalternativestrategytoavoidancewhenthecatisattemptingtopreventananticipatednegativeoutcome.Ingeneralterms,catstendtoavoidthreateningstimulibywithdrawing,hidingorclimbing.However,aggressionisusedasadefensivestrategywhenotherbehavioursarenotsuccessful.Forexample,thismighthappenwhereowners‘pursue’catsthatarefrightened,andtrytointeractwiththem.Onceacathaslearnedthataggressionisaneffectivestrategytokeeppeopleawayitwillbecomemoreconfidentintheexpressionofthisbehaviour.Overrepeatedlearningopportunities,therefore,catsmaydevelopapparently‘offensive’aggression,evenlaunchingthemselvesatpeople,orshowingthebehaviourimmediatelyonperceivingaparticularperson.Itisparticularlyimportant,therefore,thatopportunitiesforwithdrawalareprovidedforcatshousedinconfinedspaces,suchasrescuecentres(KryandCasey,2007),topreventthedevelopmentofaggressioninthiscontext.

Althoughthereismuchlessevidenceastohowsocialtoleranceforothercatsmaybelearnedduringthesensitiveperiod,itislikelythatlearningaboutconspecificsoccursinparallelwithlearningaboutpeople.Differencesintherelativetoleranceofindividualcatsforothercatsmaythereforebemodifiedbyearlyexperience:thisisanareawherefurtherresearchisimportant,giventheimportanceofsocialstressforboththeoccurrenceofundesiredbehaviourandthewelfareofdomesticcats.Asdiscussedwithrespecttointeractionwithpeople,ongoingsocialexperienceofothercatsmaymodifyperceptionsformed

people,ongoingsocialexperienceofothercatsmaymodifyperceptionsformedduringthesensitiveperiod.Forexample,acatthatisrepeatedlyattackedbyanothercatmaysubsequentlyfeelevenmorethreatenedbytheproximityofothercats.

Afurtherimportantconsequenceoftheearlyenvironmentistheformationofthekitten’spreferencesfortoiletinglocation.Apreferenceforsubstratesonwhichtotoiletformsinthefirstweeksoflife.Thisoccursaskittensformanassociationbetweentheactoftoiletingandthematerialundertheirfeetatthetime.Aswithotherassociationsmadeduringthisperiodofdevelopment,thiswillofteninfluencetoiletingbehaviourthroughoutlife.Forexample,wherebreedersuseaparticulartypeoflittermaterial,kittensmaynotrecognizeothersubstratesas‘toilets’afterhoming.Itisprobablyalsoforthisreasonthathand-rearedkittenssometimeshaveapropensityforeliminatingonsoftfurnishings.Sincetheircarerstendtoholdtheminatowelwhilststimulatingtheurogenitalreflex(emulatingthemother’susualbehaviour),someappeartoformanassociationbetweentoiletingandsoftmaterials.Althoughsubstratepreferencesformedduringdevelopmentcanbestrong,preferencescanalsochangeintheadultcat.Generallythisisdesirable,asitenablesownerstograduallychangefromonelittermaterialtoanotheriftheywishto.However,ifacathaspersistentlytoiletedinaninappropriatelocation,suchasacarpet,overaprolongedperiod,itcanformapreferenceforthissurface,makingresolutionoftheproblemmoredifficult.

Abilitytoshowanormalbehaviouralrepertoire

Anotherimportantfactorinthedevelopmentofundesiredbehaviouristheextenttowhichcatsareabletoshowafullnormalbehaviouralrepertoire.Thenormalethologyofthecatanditsspecies-specificbehavioursshouldthereforebeconsideredaspotentiallyrelevantinindividualcaseswherenaturalbehaviouralopportunitiesarerestricted.Theextenttowhichcatstoleraterestrictionsinactivity,socialinteractionandpredatorybehaviourappearstovaryconsiderablybetweenindividuals,andislikelytobeinfluencedbothbypersonalitycharacteristics(seeChapter9,thisvolume)aswellas‘expectations’derivedfrompreviousexperiences(seeChapter3,thisvolume).Forexample,catsappeartovaryconsiderablyintheextenttowhichbehaviourisalteredbyindoor-onlyhousing,althoughanecdotalevidencesuggeststhatcatsthatwerepreviouslyactiveoutsidearemorelikelytoshowfrustration-associatedresponsestospatialrestriction.However,toleranceofspatialrestrictionwillalsovarywiththeextenttowhichownersareabletoenrichtheindoorenvironment–

forexample,throughtheprovisionofplaytosimulatehuntingactivity,feedingenrichmentandopportunitiesforclimbingutilizingathree-dimensionalspace(Ellis,2009).

Theabilitytodisplaypredatorybehaviour,ortodirectpredatory-typeresponsesintoplay,appearstobeparticularlyimportantforthecat.Restrictedabilitytoshowthistypeofbehaviourcanresultinfrustration,andthustoundesiredbehavioursincludinghuman-directedaggression.Inappropriatepredatory/playaggressiontowardsownersoftenfirstdevelopswherethesebehaviourpatternsaremisdirectedinthekitten.Playbehaviourinkittensisimportantinthedevelopmentofthemotorresponsesneededforpredatorybehaviour(seeChapter7,thisvolume).Inaferalsituation,this‘practising’isinitiallydirectedatinanimateobjects,butislaterdirectedbythequeentowardspreyitemsthatshebringsbacktothenestsite(Kitchener,1991).Thekittensthereforelearntheappropriateconditionedcuesthatstimulatethesebehaviours.Inthedomesticenvironment,ownersarefrequentlytemptedtoplaywiththeirkittensby,forexample,wigglingtheirfingersormovingtheirfeetaroundunderaduvet.Whilethisisrelativelyharmlesswithakitten,itcanleadtoinappropriateplay/predatoryaggressionbeingdirectedtowardshandsorfeetoncethecatmaturestoanadult.Inthesecases,catsoften‘ambush’theirownersastheywalkpastbyrushingoutfrombehindfurnitureandgrabbingtheirfeetorlegs,orswipingatthemastheywalkpast.Oncethistypeofbehaviourstartsitisoftenreinforcedbytheresponseoftheowner–shrieking,pullingarmsawayorrunningawaytendtoreinforcetheresponse,justasmovementandsquealingofapreyitemwouldencouragefurtherattack.

Theotherimportantaspectofthenormalrequirementsofcatsthatmayleadtoundesiredbehaviourispatternsoftoileting.Eliminatingawayfromnormaltoiletingsitescommonlyoccurswherecatscannoteasilyaccessthesesites,asdiscussedearlier.However,toiletingbehaviourcanalsochangewheretoiletinglocationsnolongerfulfilthecat’srequirementsoftoiletinglocations,suchasasuitablehiddenlocationandlittersubstrate(Table11.1).Forexample,sincecatsaregenerallyfastidiousintheireliminatinghabit,alittertraythatiscleanedinfrequentlyisapossiblereasonforchoosinganalternativelocation(Herron,2010).However,sincetheyareattractedtoatoiletingsitepartlybyolfactorycues,over-zealouscleaning,particularlywithastrong-smellingdisinfectant,canalsocauseproblems.

Learntexperience

Learntexperienceplaysanimportantroleinthedevelopmentofalmostallundesiredresponses.Forexample,althoughrelevantsocializationexperiencemayleadtoageneralacceptabilityofsocialcontactwithpeople,itisindividuallearningexperienceonfirstcontactthatdetermineshowanindividualcatrespondstoanyperceivedthreatfromaspecifichumancontact.Furthermore,learningopportunitiesthroughoutlifeconstantlymodifyhowacatrespondstovariousaspectsofitssocialandphysicalenvironment.Hencealthoughfirstexperiencesstronglyinfluenceongoingresponses,behaviourcanbesubsequentlymodified.Indeed,theabilityofcatstolearnnewassociationsformsthebasisofbehaviourmodificationprogrammes.Inmanycases,relativesocializationexperienceisreinforcedbysubsequentexperience–forexample,acatthatiswaryofpeopleduetolimitedsocializationislikelytoavoidpeoplewhoapproachit.Wherethisstrategysuccessfullyavoidstheperceivedthreat,theresponsewillbereinforced,andhencemorelikelytobeusedonsubsequentencounters(seeChapter3,thisvolume).Insomecases,however,fearresponsesoccurentirelyasaresultofspecificaversiveexperiences.Thesebehavioursincatsbecomereinforcedastheyaresuccessfulatavoidingtheperceivedaversiveevent.Throughthenormalprocessesofassociativelearning,catswillalsobecomegraduallymoresensitizedtoaversiveevents,suchthattheyrespondatalowerthresholdofstimulus,andwillalsogeneralizeanavoidanceresponsetosimilarstimuli.

Inadditiontotheavoidanceresponse,catsthatvaluehumanattentionwillcommonlylearnbehavioursthataresuccessfulatachievingthis.Forexample,acatthatvalueshumanattentionmaylearnthatwalkingalongashelfand‘wobbling’valuedornamentsisaveryeffectivewaytoachieveinteractionwithownerswhowereotherwisewatchingtheTV.

Learningisalsoimportantinunderstandinghowthebehaviourofindividualcatsmaydevelopovertime.Urinesprayingincatsoftenfirstdevelopsasaresponsetotheproximityofsociallyincompatiblecats,asdiscussedearlier.However,onceestablished,theoccurrenceofthebehaviourcanbemodifiedthroughlearning.Forexample,thefrequencyandlocationofurine-sprayingbehaviourcanalterdependingonhowownersrespondtothebehaviour,andhowcatsperceivethisresponse.Becauseownersfindthebehaviourundesirable,manycatsthaturinesprayarepunished,forexamplebybeingshoutedatorchasedoutoftheroom.Mostcatsfindthisresponseaversive,andwilllearntoleavescentmarksonlywhenownersarenotpresent.However,otherconsequencesarepossiblefromthisresponsebyowners.Somecatsmaybecomeveryanxiousaboutthechangeinthebehaviouroftheirowner:sincetheydonotunderstandthereasonforthepunishment,theyperceiveonlythattheirowneris

understandthereasonforthepunishment,theyperceiveonlythattheirownerisbehavingstrangelytowardsthem.Sincespraymarkingisusedto‘identify’contextsinwhichvigilanceisrequired,catswillsometimesusethebehaviourtoidentifyownersassomethingofwhichtobewary,andurinespraydirectlyontheirowners,oronitemsthatsmelloftheirowners.Unfortunatelymanyownersanthropomorphicallyassumethatthisbehaviouris‘vindictive’or‘spiteful’,therebyleadingtoanincreaseinpunishment,andhenceanxietyinthecat,andsotherateofspraymarkingspiralsupwards.Afurtherpotentialconsequenceofownerspunishingtheircatforurinesprayingiswherecatsdonotperceivetheowner’sresponseaspunishing,butratherreinforcing.Thisoccursoccasionallyincatsthatarehighlymotivatedtoachieveownerattention,andinsuchcasesurinesprayingmaydevelopasaresponsethatoccursinfrontofownersinordertoachievearesponsefromthem.

Themannerinwhichcatsperceivetheconsequencesoftheirownactions,andbehaviouralstrategieslearnttoresolvesituationsofperceivedthreatorachievedesiredgoals,arethereforeimportantfactorsinmodulatinghowindividualcatsrespondindifferentcircumstances.