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AnimalSentience2017.
013:KujalaonCanineEmotions1CanineemotionsasseenthroughhumansocialcognitionMiiamaariaV.
KujalaDepartmentofEquineandSmallAnimalMedicine,UniversityofHelsinki,FinlandDepartmentofNeuroscienceandBiomedicalEngineering,AaltoUniversity,FinlandAbstract:Itisnotpossibletodemonstratethatdogs(Canisfamiliaris)feelemotions,butthesameistrueforallotherspecies,includingourown.
Theissuemustthereforebeapproachedindirectly,usingpremisessimilartothoseusedwithhumans.
Recentmethodologicaladvancesincanineresearchrevealwhatdogsexperienceandwhattheyderivefromtheemotionsperceptibleinothers.
Dogsattendtosocialcues,theyrespondappropriatelytothevalenceofhumananddogfacialexpressionsandvocalizationsofemotion,andtheirlimbicrewardregionsrespondtotheodoroftheircaretakers.
Theybehavedifferentlyaccordingtotheemotionalsituation,showemotionallydrivenexpectations,haveaffectivedisorders,andexhibitsomesubcomponentsofempathy.
Thecaninebrainincludesarelativelylargeprefrontalcortex,andlikeprimates,dogshaveabrainareaspecializedforfaceperception.
Dogshavemanydegreesofemotion,butthefullextentofdogemotionsremainsunknown.
Humansareasociallymindedspecies;wereadilyimputemindandemotiontoothers,eventovegetablesorrocks.
Hencetheexperimentalresultsneedtobeanalyzedcarefully,sotheemotionallivesofdogsareaccuratelyestimated.
Keywords:Canisfamiliaris,domesticdog,emotion,psychology,socialcognition,comparativecognition,neuroscienceMiiamaariaV.
Kujala(Saarela)isaresearcherincomparativecognitionattheUniversityofHelsinki,FacultyofVeterinaryMedicine,DepartmentofEquineandSmallAnimalMedicine,Helsinki,Finland;andAaltoUniversitySchoolofScience,DepartmentofNeuroscienceandBiomedicalEngineering,Espoo,Finland.
Asocialcognitiveneuroscientist,hercurrentresearchinterestsaresocialcognitionandemotionalexperienceindogs,non-invasivebrainresearchacrossspecies,andhumanperceptionandinterpretationofanimals.
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htmlAnimalSentience2017.
013:KujalaonCanineEmotions21.
IntroductionDogsareourage-olddomesticatedcompanions.
Darwin(1872)considereddogsasacomparativeexampleinhisworkonemotionalexpressionacrossspecies.
Sharingthelivingenvironmentwithus,dogshavedevelopedremarkablesocialskillsininter-speciescommunication(fortheoriginalarticles,seeHareetal.
1998,Sopronietal.
2001,Calletal.
2003,Miklósietal.
2003,Kaminskietal.
2009).
Thus,itisnosurprisethatwehaveinherentinterestinunderstandingdogexperience,behavior,andcognition.
However,provingthatsomeoneexperiencessomethingisanimpossibilitybecauseexperiencesaresubjective(Nagel1974).
Wecanneverknowthatanotherpersonexperiencesthesamethingasus.
Therearealwayssubtledifferencesintheunderlyingpsychologyandphysiology,althoughthelarger-scaleresponsesmaybesimilarwithinaspecies.
How,then,canweknowanythingabouttheinnerlivesofotherspeciessuchasourcompaniondogsInhumanemotionresearch,"emotion"and"feeling"areoftenseparatedbecauseemotioncanbetargetedforobjectiveexperimentalstudythroughbehavioralandneurophysiologicobservations,whereasfeelingissubjective:Whattheemotionfeelslikeandhowitisinterpretedbythesubjectcanonlybeinferredindirectly.
Wecandetectthebehavioralandphysiologicalcorrelatesofbothmyhappinessandyourhappiness,buttheycanbefeltandinterpretedverydifferentlybyeachofus.
Inthiswork,thetopicofcanineemotionsisapproachedusingtheframeworkofAndersonandAdolphs(2014).
Theyarguethatthecapacityforemotioncanexistacrossphylogeny,butemotionsmayconsistofadifferentsetofparallelbehavioral,somatic,physiological,andcognitiveresponsesindifferentspecies.
Webeginbyconsideringtheeffectofthehumanviewpoint.
Canineemotionsareexaminedbothbehaviorallyandbiologically,withabriefreviewoftheneuralbasisforprimaryemotionsandtherespectivestructuresinthedogbrain.
Theneuralbasisforsecondaryemotionsisalsoreviewed,followedbyadiscussionofthecurrentresearchondogs.
Long-termmoodsandcomparativeaspectsarealsoconsidered.
Theultimatepurposeofthistargetarticleistostimulatediscussionaboutthenatureandextentofdogemotionsandtheneedforthisnewfieldofresearch,aswellastoprovidegroundworkfortheapproachfromvariousscientificdisciplines.
Oneofthedifficultiesinconsideringemotionalstatesindogsistheinconsistencyinterminologyacrossstudies.
Avoidinganthropomorphictermshasleftmanycanineaffectivephenomenawithoutastandardizedterminology.
Differentresearchershaveuseddifferenttermsforthesamephenomenon,orsimilartermsforseparatephenomena.
Thisreviewattemptstointegrateresultsacrossstudiesanddisciplines.
2.
HumansocialcognitionaffectsperceptionofdogemotionalstatesTheexistenceofemotionsindogsandtheperceptionofdogemotionsbythehumancaretakersareseparateissues.
Everydaylifepresentsmanypossibilitiesforhumanstomisinterpretthemindbehindadog'sbehavior.
Forexample,guardiansmaymisinterpretthedog'saffectivestateinseparation-relatedanxiety(seeMendletal.
2010a).
Accordingtothethree-factortheoryofanthropomorphism(Epleyetal.
2007),behavioralinterpretationsthatareoftenvalidwithotherhumansarealsoeasilyattributedtonon-humananimalssuchasAnimalSentience2017.
013:KujalaonCanineEmotions3dogs.
Thus,human,biologicallytunedsocialperceptionisthestartingpoint,asitfiltersourunderstandingofdogemotions.
Peoplebelievethatanimalssuchasdogsexperienceemotions(Morrisetal.
2008,Morrisetal.
2012,Walkeretal.
2014).
Humansarealsoquiteconsistentinclassifyingdogs'emotionalbehaviorindifferentcontexts(Pongráczetal.
2005,Tami&Gallagher2009,Walkeretal.
2010,Bucklandetal.
2014,Faragóetal.
2014,Lakestanietal.
2014).
Humansfriendlydogbehaviormosteasilyrecognize,whereasaggressionandfeararemoredifficulttoidentify(Tami&Gallagher2009,Wanetal.
2012,Mirkóetal.
2013,Lakestanietal.
2014)—especiallybychildren(Meintsetal.
2010,Lakestanietal.
2014).
Priorexperienceofdogbehaviourandtraining,ratherthanmereguardianship,enhancestheinterpretationofcaninebehaviourfromthewhole-bodycues(Kujalaetal.
2012,Wanetal.
2012).
Althoughemotionsarevisiblethroughoutdogbodilycues,humanattentionisgenerallydrawntothefacesofbothhumans(Johnsonetal.
1991)anddogs(Quinnetal.
2009).
Humanscanclassifyadog'semotionalvalence(positivity-negativity)fromthefaceirrespectiveofpriorexperiencewithdogs(Bloom&Friedman2013,Schirmeretal.
2013).
Theycandistinguishhappiness(88%ofthetime)andanger/aggressiveness(70%)fromadog'sface,butdiscriminationofotherdiscreteexpressionsislessreliable(fear:45%,sadness:37%,surprise:20%,anddisgust:13%;Bloom&Friedman2013).
Perceptionofothersisaffectedbymanyfactorsinthehumanmind.
Thehumansocialmindisequippedwithapresuppositionofintentionality(forreviews,Blytheetal.
1999,Scholl&Tremoulet2000,Urquiza-Haas&Kotrschal2015),fromwhichanthropomorphismcanarise.
Attributingintentionalityorotherhumancharacteristicstonon-livingthingsisstrengthenedbypersonalconnection(Kiesleretal.
2006),andmentalattributionisfoundinpeople'sdescriptionsofrocks(Kiesler&Kiesler2005),computers(e.
g.
,Nassetal.
1994),animations(Chaminadeetal.
2007),robots(Gazzolaetal.
2007,Imamuraetal.
2015,Martinietal.
2016),orevenvegetables(Vaesetal.
2016).
Humansalsoprojecttheirviewsofthemselvesontodogsmuchastheydowithconspecifics,andtheirperceptionsofdogsaresimilarlyaffectedbystereotypes(Kwanetal.
2008).
Thus,humanseasilyattributementalandemotionalstatestocompaniondogs,andhumaninterpretationofcanineemotionsisfilteredbyhumanpsychologicalcharacteristics(Kujalaetal.
2017).
Humanscanalsodenyhumanityinotherhumans(forreviews,Leyensetal.
2000,Haslam2006).
Theyconsistentlyattributemorecomplexemotionstotheirin-groupthanout-groupmembers(seeLeyensetal.
2000).
Inahumanbrainimagingstudy,theobservationofimagesofextremeout-groupmembers(suchasdrugaddictsorthehomeless)failedtoproducethemedialprefrontalcortexactivationconnectedwithsocialcognition(Harris&Fiske2006).
Thus,thehumanmindisaffectedbyvarioussocialfactors,withthejudgmentssometimesrepresentingmorethejudge'sownideologythanthereality.
Likewise,whenhumansattempttodeciphercanineemotions,dogguardianscanunderestimatetheirdogs'aggressiveness(Mirkóetal.
2013).
Attributingmindstoothersisinnateinhumans,andthehumanbrainappearsremarkablyflexibleregardingthesourceoftheothermind.
Humanempathygeneralizestootherspecies(Ascione1992,Paul2000,Taylor&Signal2005,Norringetal.
2014,WestburyInghametal.
2015,Kujalaetal.
2017)andaffectsourinterpretationofdogbehavior(Meyer&Forkman2014,Meyeretal.
2014).
Empathy(WestburyInghametal.
2015)andtheattributionofmentalstates(Harrison&Hall2010)tonon-humananimalsvarieswiththeirAnimalSentience2017.
013:KujalaonCanineEmotions4phylogeneticrelatednesstohumans.
Mentalattributiontobothhumanandnon-humanspeciesisconnectedtothetemporoparietaljunctionassociatedwithhumantheory-of-mindabilities(Cullenetal.
2014).
Humanbrainresponsestodogscanalsobestrikinglysimilartoresponsestohumanconspecifics,whetherobservingdogs'facialexpressions(Spuntetal.
2016),pain(Franklinetal.
2013),orsocialinteraction(Kujalaetal.
2012).
Humanbrainsseekothermindsandemotions,anddogemotionalbehaviourisfilteredthroughthesamemachinery.
3.
Neuralsupportforthebasic(primary)canineemotionsItisgenerallyagreedthatbasicemotionalstatessuchasanger,happiness,andfearareevolutionarilyadaptive(Ekman1992,Izard1992,Panksepp1998,Plutchik2001),andtheyhaveuniversalfacialexpressionpatternsinhumans(Ekman&Friesen1971).
Basicemotionalstatesareassociatedwithneuralstructureswithinthelimbicsystemanditsconnectionstotheneocortexinmammals(Damasio1994;LeDoux1996;Rolls1999).
Specificbasicemotionsareassociatedwithspecificchemicalneurotransmitterbalanceinthebrain(Panksepp1998).
Figure1.
Keyareasofthedogbrain.
(A)Somekeysubcorticalareasshownontheaxialmagneticresonanceimagingslices(dog'snosepointingupwards;top=anterior,bottom=posterior,left=lefthemisphere,andright=righthemisphere)and(B)thecorticalsurfaceofthedogbrain,withthesulciopenedandthenomenclatureoverlaidbelow.
Thecorticalsurfaceisshownlaterallyfromthelefthemisphere(dog'snosepointingleft;left=anterior,right=posterior,top=dorsal,andbottom=ventral);theimagehasbeenmagnifiedwithstandarddigitalimageprocessingtoshowthegyriandsulci.
Modified,underthetermsoftheCreativeCommonsAttributionLicense,fromfigureshttp://dx.
doi.
org/10.
1371/journal.
pone.
0052140.
g002andhttp://dx.
doi.
org/10.
1371/journal.
pone.
0052140.
g003(Dattaetal.
2012).
AnimalSentience2017.
013:KujalaonCanineEmotions5Thedomesticdog,asamemberofmammalianorderCarnivoraandtheCaninaefamily,hasabrainthatincludesallthemajorstructuresandconnectionssupportingbasicemotionalfunctions(Jensen2007,deLahunta&Glass2009,Evans&deLahunta2013).
Dogbrainsinclude,bilaterally,thelimbicsystemwiththenucleusaccumbens;theamygdalawithitssensoryandcorticalconnections;thecingulatecortex;andthesensory-motorcorticesandtheinsula,deepwithinthepseudosylvianfissure(inhumans,theSylvianfissureoccupiesthetopologicallyequivalentposition).
Dogsalsohavearelativelylargeprefrontalcortexthatisnotdirectlyassociatedwithmotorfunctions(seeFigure1,Palazzi2011,Dattaetal.
2012,Evans&deLahunta2013).
Thecorrespondingstructuresinhumanshavebeenstudiedextensivelyinrecentdecadesregardingtheirrolesinsocialandemotionalfunction(forreviews,seee.
g.
,Bushetal.
2000,Damasioetal.
2000,Adolphs2002,Leppnen&Nelson2009,Etkinetal.
2011,Schilbachetal.
2013).
Fordogs,theresearchismorescattered.
Somedogbrainfunctionisalsoinferredfromneurologicalexperimentswithcats(Feliscatus)andthehomologuesbetweenthebrainanatomiesofthetwospecies.
Manyfunctionsofthesubcorticalnuclei(e.
g.
,theseptalareaorhypothalamus)andthefinerneurophysiologicdetails,aswellasvisualcorticalorganization,areinferredfromcatstudies(King1987,deLahunta&Glass2009,Sjaastadetal.
2010).
Utilizationofthemethodologyforstudyinghumanbrainfunctionhasfacilitatedthestudyofdogbrains.
Arecentstudyshowedthatdogs'nucleusaccumbensisactivatedbytheodoroffamiliarhumans,highlightingthepossibilitiesofmethodologicaladvancesinexaminingdogemotions(Bernsetal.
2015).
Dogandhumanbrainsalsohaveimportantdifferences.
Theassociationareas(brainareasnotdirectlyresponsibleforsensomotorfunctions)coverabout20%ofthedogneocortexbut85%ofthehumanneocortex(Evans&deLahunta2013).
Therhinencephalon,devotedtoprocessingolfactorysignals,coversarelativelylargeareaindogbrains(Evans&deLahunta2013).
Theexistenceoflimbicandcorticalstructuresindogsisconsistentwithhavingthebasicemotions,althoughdogs'qualia–whatitfeelsliketobeadog(Nagel1974)—nodoubtdifferfromourown.
4.
Emotionalreactivityandaffective-behavioraldisordersindogsResearchondogemotionshastraditionallyconcentratedontheproblemsdogbehaviorcausesforthehumanguardians,whichiswhyweknowmoreaboutdogfearandaggression.
Dogemotionalityhasbeenstudiedtopredictgeneralemotionalreactivity(Goddard&Beilharz1986,Sforzinietal.
2009),aggression(Netto&Planta1997,vandenBergetal.
2003,vanderBorgetal.
2010),behavioraldisorders(vanderBorgetal.
1991),anddifferencesamongdogbreeds(Scott&Fuller1965)frompuppiestoadulthood.
Dogaggressivenessistestedbypresentingprovocativestimuli,suchasanunfamiliarbarkingdog(Netto&Planta1997,vandenBergetal.
2003)orstaringthedogintheeyes(Sforzinietal.
2009).
Guardianquestionnairesarealsoused(Netto&Planta1997,Hsu&Serpell2003,Duffyetal.
2008).
Similarly,testingadog'sfearfulnesscanincludepresentingasuddenloudnoise,anovelobject,afallingbag,oragunshot(Melzack1952,Beerdaetal.
1998,Kingetal.
2003,Hydbring-Sandbergetal.
2004,Morrowetal.
2015).
AnimalSentience2017.
013:KujalaonCanineEmotions6Bothfearfulandaggressivebehaviorsindogsareassociatedwithsomephysiologicalorautonomicresponses.
Stimulielicitingfearfulbehaviorindogsincreasetheircortisol(Beerdaetal.
1998,Kingetal.
2003,Hydbring-Sandbergetal.
2004,Dreschel&Granger2005,Morrowetal.
2015)orprogesteronelevels(Hydbring-Sandbergetal.
2004),heartrates(Kingetal.
2003,Hydbring-Sandbergetal.
2004,Ogataetal.
2006),andbodytemperatures(Ogataetal.
2006).
Aggressivebehaviorsareassociatedwithreducedserotonergicfunction(Reisneretal.
1996).
Themostcommonaffective-behavioralclinicallytreateddisordersindogsarerelatedtofearfuloraggressivebehaviorandmaybeinducedbyseparationanxiety,noisesensitivity(forreview,Sherman&Mills2008),anddominance/competitiveaggression(seee.
g.
,Beaver1983,Wright&Nesselrote1987,Cameron1997,Reisner1997,Haug2008).
Treatmentsfortheseconditionsusuallyincludebehaviormodification,oftencombinedwithneuropharmacologicalmedicationasinhumanpsychiatricdisorders(Overall2000).
Takentogether,aggressionandfeararethemoststudiedemotionsindogs,butresearchonotheremotionalstatesisscarce.
Anexceptionamongthepositiveemotionsisdogplaybehavior,whichiswell-documented(Bekoff1974a,Bekoff1974b,Bekoff1995,Rooneyetal.
2000,Horvathetal.
2008,Wardetal.
2008,Horowitz2009a,Palagietal.
2015).
5.
ProductionandperceptionoffacialexpressionsFacesandfacialexpressionsconveydelicateandmeaningfulinformationaboutemotionalstatestoconspecificsinhumans(forreviews,Adolphs2002,Calder&Young2005,Hari&Kujala2009,Leppnen&Nelson2009)aswellasinmanynon-humanspecies(forreviews,Tateetal.
2006;Leopold&Rhodes2010).
FacialexpressionsofemotionindogswerediscussedbyDarwin(1872);theycharacterizedingreatdetailsincethe1960s,notingsimilaritiesintheemotionalexpressionsforaggressionandhappinessbetweendogs,othercarnivores,andprimates(Bolwig1964,Fox1970).
Aprecisecodingofhumanfacialexpressionsbasedonthemovementoffacialmuscles—afacialactioncodingsystem(FACS)—wasdevelopedinthe1970s(Ekman&Friesen1978).
Thesystemhassincebeenappliedtomanyotherprimatespecies(Vicketal.
2007,Parretal.
2010,Walleretal.
2012,Caeiroetal.
2013),horses(Equuscaballus,Wathanetal.
2015),cats(http://www.
catfacs.
com/),anddogs(Walleretal.
2013).
Deviatingfromthehuman-FACS,thenon-human-FACSoftenincludesthemovementofears.
Behavioralandbrainresponsesduringtheperceptionoffacialexpressionshavebeenstudiedinnon-humanprimatesandsheepfordecades(seeTateetal.
2006).
Asasecondnon-primatespeciesaftersheep(Kendrick&Baldwin1987),dogshavebeenshowntopossessadistinguishableface-processingregioninthetemporalcortex,separatingbrainresponsestofacesfromtheresponsestoobjects(Figure2)(Dilksetal.
2015,Cuayaetal.
2016).
Theresponseprofilesareroughlycomparablewiththoseofthehumanfusiformfacearea(Kanwisheretal.
1997),althoughthecorticalregionseemstobemorevariableindogs.
Inhumans,faceprocessingcontinuesfromthefusiformtotheinferotemporalcortexandthesuperiortemporalsulcus,withtheextractionofidentity-andemotion-specificinformation(forreview,Haxbyetal.
2000).
AnimalSentience2017.
013:KujalaonCanineEmotions7Figure2.
Strongerbrainresponsesfromdogsforfacesversusobjects.
Imagesshowthefocusofbrainactivationduringnon-invasivefunctionalmagneticresonanceimaginginsevendogsforfacesversusobjectswithcontrast,overlaidonadigitallyproducedglassbraintorevealfocilocatedwithinthesulci.
(A)Lateralviewfromlefthemisphere,(B)rostralviewfromfront,and(C)lateralviewfromrighthemisphere.
A=anterior,P=posterior,S=superior(ordorsal),I=inferior(andventral),L=left,andR=right.
Modified,underthetermsoftheCreativeCommonsAttributionLicense,fromfigurehttp://dx.
doi.
org/10.
1371/journal.
pone.
0149431.
g005(Cuayaetal.
2016).
Whiledirectinformationondogs'brainprocessingofemotionalexpressionsismissing,agrowingbodyofbehavioralandeye-trackingresearchsupportstheabilityofdogstodistinguishnegativeandpositivefacialexpressionsinbothhumansanddogs,andtorespondappropriatelyaccordingtothevalenceoffaces(Nagasawaetal.
2011,Raccaetal.
2012,Mülleretal.
2015,Barberetal.
2016,Somppietal.
2016)(Figure3).
Figure3.
Gazefixations(circles)andscanningpaths(linesbetweenthecircles)oftwodogs(showninlightanddarkgreen)forfacialexpressionsofdogsandhumans.
Whitecirclesrepresentthetargetsofthefirstfixations;dogstendtogazefirstintotheeyesofbothhumansanddogs.
Figurefromhttp://dx.
doi.
org/10.
1371/journal.
pone.
0149431.
g005(Somppietal.
2016),reprintedunderthetermsoftheCreativeCommonsAttributionLicense.
AnimalSentience2017.
013:KujalaonCanineEmotions8Dogsassociateemotionalvocalizationsofbothhumansanddogswiththecorrespondingfacialexpressions,showingmultisensoryprocessingofemotionalexpressions(Albuquerqueetal.
2016).
Likehumaninfants,dogsusehumanemotionalexpressionsforsocialreferencing,asasourceofapproach/avoidinformationfornovelobjects(Merolaetal.
2012b,Merolaetal.
2012a,Buttelmann&Tomasello2013,Merolaetal.
2014,Turcsánetal.
2015).
Furthermore,theyappeartogeneralizethevalenceinformationoffacialexpressionsacrosshumanindividuals(Mülleretal.
2015,Somppietal.
2016)ratherthanrespondingonlytoguardians'expressions—incontrasttocats,whorespondmainlytothevalenceoftheirguardians'facialexpressions(Galvan&Vonk2016).
Humancross-culturalstudiescouldprovidesomeusefulcluesforstudyingemotionsindogs.
Thebasicemotionsareremarkablysimilararoundtheworld(Ekman&Friesen1971).
Facialexpressionsandtheirrecognition,situationsprovokingemotions,andtheorganizationofemotionsonthevalence-arousaldimensionsareconsistentacrosscultures(Shaveretal.
1992).
However,theperceptionofemotionalintensitydiffersacrosscultures(Ekmanetal.
1987),andthedecodingandencodingofemotionintheculturalin-groupappearsmoreaccurate(formeta-analysis,Elfenbein&Ambady2002).
Similarlytotheculturaldifferencesinhumans,differencesinbreedsorenvironmentmayaffecttheexpressionorperceptionofemotionindogs(forreview,Mehrkam&Wynne2014).
Itappearsunquestionablethatdogscanbothproduceandprocessemotionsthroughfacialexpression,butthequestionistowhatextentOneremainingquestionconcernswhatpartofrecognizinghumanfacialexpressionsbydogsisinnateandwhatislearnedthroughassociationandexperience.
Also,theresearchaddressedmainlythepositive-negativevalenceinformationoffacesratherthanthemorediverse,discreteexpressionsofemotion(e.
g.
,happiness,sadness,surprise,fear,disgust,anger:Ekman&Friesen1971),soinformationondogs'abilitytodiscriminateorrespondtodiscreteexpressionsofemotionislacking.
Arethereuniversalfacialexpressionsofemotionindogsasthereareinhumans(Ekman&Friesen1971)—andifso,whatarethey6.
Fundamentalbasisforsecondary(social)emotionsSecondaryemotions—generatedthroughtheinterpretationofsocialsituationsandrequiringsomesenseofanother'smind—arelesslikelythanbasic(primary)emotionstobeattributedtodogsbypeople,but22to94%ofpeoplebelievethatdogsdohavesecondaryemotionssuchasshameorguilt(Morrisetal.
2008,Morrisetal.
2012).
Asadulthumans,weeffortlesslyattributesecondaryemotionstootherpeople.
Withoutknowledgeofthedifferencesamongmindsacrossspecies,wecanjustaseasilyattributetheemotionstonon-humans,includingdogs.
However,dogsmaybeincapableofexperiencingthemorecomplexsocialemotions,ortheirexperiencesmaybequalitativelyverydifferentfromours.
Thereasonthedogishuman'sbestfriendmaybetheapparentlymissingcaninecapacityforsecondaryemotionssuchascontemptorSchadenfreude(thejoyinothers'misfortune).
Thesecondaryemotionsseemtorequiresomesenseoftheself(Leary2003).
Havingself-awarenesscomplicatestheemotionalexperienceinmanyways—allowingimaginedexperienceswithnobasisinreality.
Leary(2003)clarifiestheeffectofselfonemotionalexperiencethroughfivepoints:"Specifically,havingaselfpermitspeopleto(1)evokeemotionsinthemselvesbyimagingself-relevantevents,(2)reactemotionallytoabstractandAnimalSentience2017.
013:KujalaonCanineEmotions9symbolicimagesofthemselvesintheirownminds,(3)consciouslycontemplatethecauseoftheiremotions,(4)experienceemotionsbythinkingabouthowtheyareperceivedbyotherpeople,and(5)deliberatelyregulatetheiremotionalexperience"(p.
775).
Ashealsopointsout,animalsdonotneedaconceptofselfinordertohaveabasicemotion.
Humansoftenattempttosuppressself-referentialemotionalthoughtinvariousways(e.
g.
,bydrinkingalcohol)sincesuchthoughtscancauseincreasingdistress.
Inchildren,self-awarenessappearstoariseroughlyconcurrentlywiththeabilitytotakeanother'sperspective(Lempersetal.
1977);earlystudiessuggestsimilarco-occurrencesinotherspecies(Gallup&Suarez1986).
Todate,thelevelofdogs'self-awarenessisnotknown—forexample,theyhavenotpassedGallup's(1986)mirrorself-recognitiontest,buttheydospendlesstimeinspectingtheirownurinemarkingsthanthoseofothers(Bekoff2001).
Thebrainregionsresponsibleforsecondaryemotionsincludeanetworkcomprisingthemedialorbitofrontalcortex,thetemporalpole,andthesuperiortemporalsulcusinhumans(Molletal.
2002,Burnett&Blakemore2009).
Inprinciple,homologuesoftheseregionsmayalsobepresentinthebrainsofdogs,inthetemporalandfrontalassociationareas.
Homologuesincorticesaredifficulttoverifybecausethebrainfunctionsofthesecorticalregionscannotbelocalizedanatomically,thusfunctionalbrainimagingisneeded.
Nevertheless,thecorticalassociationareas,associatedwithsecondaryemotionsinhumans,arelargerinhumansthanindogs(20%ofthecortexindogsand85%inhumans;Evans&deLahunta2013).
Thebrainareasrespondingtosecondaryemotionsarealsostronglyconnectedtoareasofthelimbicsystem,andtheconnectionsalterthelevelofcognitiveevaluationoftheemotionalstates(Berridge2003).
Theconnectionsbetweenthecortexandthelimbicsystemaresodifferentinmagnitudebetweenhumansandothermammalsthatcorticallesionshavingminimaleffectsonothermammalsmaycausedrasticchangesinhumanfunction(Berridge2003).
Forexample,acatwithoutacortexmaystillmoveandbehavelikeacat,whereasahumanwithoutacortex,ifalive,liesinahospitalbedcompletelyunresponsive.
Thus,itispossiblethatthere-representationofemotionsthathumanencephalizationproduceswiththeinterconnectionstothelimbicsystemmaybethesourceofsecondaryemotions.
Inotherwords,ashumanswehavethepotentialtobeangry,realizethatweareangry,ponderthecausesofouranger,noticethattheangermomentarilyaffectsourabilitytoworkorcooperate,trytosuppressouranger,thinkabouthowourangerappearstoourcompanionsandhowitaffectsourrelationships,andtrytomodifythesourceoftheanger.
Ifthecerebral-limbicinterconnectionsarethesourceofthisemotionalre-representationalabilityinhumans,theoverallcapacityofdogsforsecondaryemotions,withlessencephalizationthanhumans,maybedramaticallydifferentfromours.
7.
Dodogsdisplayguilt—ormerelyappeasementGuiltisanexampleofasecondaryemotionoftenattributedtodogs,butaccordingtocurrentresearch,itfitsthedogmindpoorly.
Horowitz(2009b)firstrecordedcaninebehaviorandgesturalcuesinasituationwheredogscoulddisobeytheguardian'scommandandeataforbiddentreat.
Bymanipulatingtheguardian'sbeliefaboutwhathappenedinthesituation,sheshowedthatdogs'gestureswerenotdifferentwhetherornottheyobeyed.
Instead,thegesturescommonlyassociatedwithdog"guilt"—forexample,avoidingeyecontact,waggingAnimalSentience2017.
013:KujalaonCanineEmotions10thetaillowandquickly,holdingone'searsorheaddown—wereevidentwhentheguardiansscoldedtheirdogs,regardlessofthedog'sactualbehaviorintheexperiment.
Thisstronglysuggestedthatthedogsrespondedtotheguardian'sbehaviorwithsubmissivegesturesinterpretedbydogguardiansas"guilt,"ratherthandisplayingremorseforamisdeedwiththe"guilty"gestures(Horowitz2009b).
Afteradoghaslearnedtheassociationbetweenacertainunwantedbehavior(e.
g.
,stealingfood)andtheguardian'spunishmentlater,theymaydisplaythesubmissivebehaviorinasimilarsituationevenbeforetheguardian'sscolding(Horowitz2009b).
Thisdoesnotrequireremorseoran"understanding"ofviolatinganorm,butasimplelearnedassociationbetweentwosuccessivesituations.
Whendogsdisplay"guilt"behavior,guardiansarelikelytoscoldtheirdogless,whichsuggeststhat"guilt"behaviormayfunctionaslearnedappeasement(Hechtetal.
2012).
Inarecentwork,petdogs'heartratesweremeasuredduringthe"forbiddentreat"experiment,anddogswhotooktheforbiddentreathadahigherheartratethandogswhodidnot(Torres-Pereira&Broom2014).
Thissuggestsalearnedassociationbetweeneatingthetreatandapossibleconsequentscolding.
Toensurethattheriseinheartratewasnotmerelyafunctionofsympatheticnervoussystemactivationintheactivecondition(eatingaforbiddentreat),asimilarresultshouldbeobtainedwithdogseatingatreatthattheywereallowed.
Alternatively,asbothpositiveandnegativestresscanincreasesympatheticnervoussystemactivation,thetreat-stealingdogsmayjustbemoreexcitedbythetreat.
Nevertheless,evenrepresentationofthecausalityoftheactionplusananticipatoryresponsetotheconsequencedoesnotrequireasenseofguilt.
8.
FairnessorunequaltreatmentofconspecificsAnotherexampleofdogaffectiverepresentationsclosertothesecondaryemotionsisinequityaversion(Figure4).
Instudiesofprimatesocialcooperation,unequaltreatmentofindividualsisrelatedtonegativeresponses(forreview,deWaal&Suchak2010).
Inhumans,thefeelingofinequalityischaracterizedbyanegativeresponsetotheviolationoffairness(forreviews,seeFehr&Rockenbach2004,Fehr&Camerer2007).
Adegreeofinequityaversionwasreportedincapuchinmonkeys(Cebusapella,Brosnan&DeWaal2003)andchimpanzees(Pantroglodytes,Brosnanetal.
2005),andafewstudieshaveinvestigatedthephenomenonindogs(Rangeetal.
2009,Horowitz2012,Rangeetal.
2012,Brucksetal.
2016).
Inasituationwhereaconspecificpartnerwasrewardedforataskandcaninesubjectswerenot,theyrefusedtoperformthetaskorhesitatedlonger(Rangeetal.
2009,Brucksetal.
2016).
AnimalSentience2017.
013:KujalaonCanineEmotions11Figure4.
Testingdoginequityaversion;bothdogsareaskedtogivethepawinturn.
Figurefromhttp://dx.
doi.
org/10.
1371/journal.
pone.
0153799.
g001(Brucksetal.
2016),reprintedunderthetermsoftheCreativeCommonsAttributionLicense.
Thephenomenondoesnotqualifyasasimpleextinctionofalearnedbehaviorbecausethecaninesubjectsrefusedtoobeyearlierafterwitnessingapartnerreceivearewardforobeying,comparedtobeingalone(Rangeetal.
2009,Rangeetal.
2012,Brucksetal.
2016).
Theyalsotendedtorefuseearlierintheunequalsituationthaninthesituationwhereneitherdogreceivedrewards(Rangeetal.
2009,Rangeetal.
2012).
Dogsalsosharedtheirfoodandinteractedlesswithpartnersafterunequalsituations,showingthatthenegativeexperienceofunequaltreatment,ornotbeingrewardedforone'sefforts,diminishessubsequentcooperationandtoleranceofcompany(Brucksetal.
2016).
Humansandsomeotherprimatescanresistunequaltreatment(i.
e.
,refusetocooperate)eitherwhentheygainlessthanthepartnerorwhentheygainmore(Brosnanetal.
2010,Blake&McAuliffe2011).
Incontrast,dogsdonotresisttheinequitywhentheyaremorerewardedthantheircompanions(Horowitz2012).
Takentogether,dogsaresensitivetoconspecificcompanyintheinequityaversiontest.
Theyrefuseearliertoperformthetaskinsituationswhentheyreceivefewerrewardsthantheirpartner,comparedtowhentheyarealone,butunlikesomeprimates,theydonotresistgainingmorethanthepartner.
Althoughdogsbehavedifferentlytowardthecompanionandexperimenterafterunequalandequalconditions(Brucksetal.
2016),thiscouldalsoreflectthenegativeoverallmoodcreatedbynotbeingrewarded.
Thedatatodatesuggestthatdogshavethecapacityforinequityaversion,butfutureworkisneededwithmoreconditionssuchasvaryingfoodquantitiesandthedirectionoftheinequality,learningthroughasocialmodel,expectationviolation,negativesituationsaffectingsubsequentbehavior,andindividualfactorssuchaspersonalityorbreed.
AnimalSentience2017.
013:KujalaonCanineEmotions129.
ResourcecompetitionasaprecursorofjealousyAnothercaseofpossiblesecondaryemotionsindogsisjealousy.
Dogguardiansreportbehaviorsrelatedtojealousyasoftenasbehaviorsrelatedtothebasicemotionofanger(Morrisetal.
2008).
However,thisalsoillustratesanthropomorphicmisunderstandings:Couplesmayreportthat"Ontherareoccasionthatwehaveacuddlehe'llstartbarkingandwhining.
"Buthuggingisnotindogs'naturalbehaviorrepertoire,sopeoplecuddlingcanappeartodogsasathreatbetweenpackmembers,towhichtheyreactbywhiningortryingtoseparatethe"fightingpackmembers.
"Inhumans,jealousyusuallyconcernsromanticrelationshipsandextendstoimaginarysituationsofarivalthreateningasignificantrelationship(Leary2003).
Aprecursorofjealousyindogsmayexistinasituationofdefendingapreviouslygainedresourcesuchasahumancompanion.
Inarecentbehavioralstudywheredogguardiansignoredtheirdogsandattendedtorealistictoydogsorotherobjects,thedogsexhibitedsignificantlymorebehaviorssuchasgoingbetweentheguardianandthetargetoftheirattention,orpushing/touchingtheguardianorthetarget,whenthetargetwasarealistictoydogratherthananobject(Harris&Prouvost2014).
Similarly,humaninfantsshowedmorenegativeresponseswhenamother'sattentionwasdirectedtowardsalife-likedollthananobjectlikeabook(Hartetal.
1998,Hart&Carrington2002).
Althoughthecurrentdataareconsistentwiththepossibilityofsituation-basedresourcedefensebeingaprecursorofjealousy,theevidenceforenvyorjealousyindogsisinconclusiveandmorerigorousresearchisrequired.
Unfortunately,thebehaviorsassociatedwithdogjealousycanalsoappearindogsasreplacementbehaviors,whenthedogisconfusedastohowtoreact.
10.
DivisionsofempathyTheneuroscientificstudyofhumanempathyexplodedinthebeginningofthe2000s(seeSingeretal.
2004,Jacksonetal.
2005,Gazzolaetal.
2006,Saarelaetal.
2007),revealingthattheemotionalaspectofempathyisprocessedinthelimbicsystem,insula,andanteriorcingulatecortex.
Similarpatternsmaybealsopossibleinthecaninebrain.
Althoughsimpleformsofnon-humanempathyhadbeenstudiedinpreviousdecades(Church1959,Rice&Gainer1962,Massermanetal.
1964,Watanabe&Ono1986),interestinanimalandhumanempathygrewwiththestudyofnon-humanprimates(forreviews,Preston&deWaal2002,deWaal&Ferrari2010),andalsoextendedtonon-primates,includingrats(Rattusnorvegicus,Ben-AmiBartaletal.
2011).
Inhumans,empathyhasthreecomponents:emotionalempathy,cognitiveempathy,andtheseparationoftheselffromtheother(seeDecety&Jackson2004).
Emotionalempathycanbefurtherdividedintoemotionalcontagion/self-distressandempathicconcern:emotionalcontagionoriginatesfromautomaticallytriggeredresponsestoothers'emotions,andempathicconcernincludesexpressingaworryaboutothers'wellbeing(Davis1980).
Emotionalcontagionisimportantforcompassion,butathighlevelsitmayleadtoanxietyandpassivityoraggressionandantisocialityratherthanhelpingbehavior(Roberts&Strayer1996).
Cognitiveempathyinvolvesatheory-of-mind-likemeta-representationofanother'semotionalstate.
Tohighlightthedifferencebetweenemotionalandcognitiveempathy,theAnimalSentience2017.
013:KujalaonCanineEmotions13lattercanbefullypreservedinhumansdiagnosedwithpsychopathictendencies,whereasthispopulationshowsmuchlessemotionalcontagionandempathicconcern(Blair2005),possiblyduetoalteredlimbicfunction(Birbaumeretal.
2005).
11.
FromemotionalcontagiontoprosocialbehaviorindogsAlthoughlongrecognizedbysome(seee.
g.
,Bekoff2007),thecapabilityofdogstoempathizehasbeenreceivingmorescientificattentionrecently.
Anecdotalreportsofdogsapparentlyconsolingconspecificsorhumansareabundant,butthetopichasbeenthoroughlyexaminedonlyinafewexperiments.
Mostresearchondogs'empathy-relatedbehaviortowardconspecificsconcernsbehaviorthatresemblesconsolation,thatis,reconciliationorpost-conflictaffiliation.
Incooperativespecies,aggressiontowardconspecificsmaybecostlyforthewholegroup.
Animportantmechanismformanagingtheeffectsofaggressionispost-conflictaffiliativebehaviorbetweentheformeropponents(reconciliation),sometimesthroughmediationbyathirdparty(deWaal&vanRoosmalen1979).
Bothreconciliationbehaviorandthird-partypost-conflictaffiliationarepresentingroup-livingdogsandwolvesinheightenedgreeting;sittingorlyingdowntogether;andsniffing,playing,orlickingthevictimofaggression(Coolsetal.
2007,Palagi&Cordoni2009,Cordoni&Palagi2015).
Emotionalcontagionacrossdogswasrecentlystudiedbyplayingfamiliarandunfamiliarconspecificwhinestodogsandexaminingtheirbehaviorduringtheplaybackandreunionwiththefamiliardogs(Quervel-Chaumetteetal.
2016).
Whenexposedtodogwhines(recordedwhenthedogwasleftalone),thecaninesubjectsweremorealertandexhibitedmorestress-relatedbehaviorscomparedwithexposuretoacousticallymatchedcontrolsounds(Quervel-Chaumetteetal.
2016).
Additionally,exposuretofamiliardogwhinestriggeredmorecomfort-offeringbehaviorstowardthepartnerdogsinreunion,resemblingpost-conflictaffiliativebehaviorobservedinnaturalgroups(Coolsetal.
2007,Palagi&Cordoni2009,Cordoni&Palagi2015).
Post-conflictaffiliationhighlightsthepossibilityofemotionalcontagionorempathicconcernindogs,althoughitsmechanismsareunknown.
Across-speciesaffiliativeinteractionbetweenadogandtheircaretaker(e.
g.
,guardianpettingthedog)cancausehormonalandphysiologicalsynchronization,loweringcortisollevelsandincreasingoxytocinanddopaminelevelsinbothspecies(Odendaal&Meintjes2003,Milleretal.
2009,Nagasawaetal.
2009,Handlinetal.
2011,Nagasawaetal.
2015).
Thisacross-speciesemotionalsynchronizationsuggestsapossiblephysiologicalmechanismfortheemotionalcontagionbothinhumansanddogs.
Inasimilarexamplerelatedtothestressresponse,cortisollevelsinbothhumansanddogsincreasedsignificantlyafterlisteningtoacryinghumaninfantcomparedwithababblinginfantorwhitenoise(Yong&Ruffman2014).
Dogsmayalsoactprosocially,pullingaropethatdeliversapartnerdogarewardevenwhenthepullerdogsthemselvesarenotrewarded,butonlyiftherecipientdogisfamiliar(Quervel-Chaumetteetal.
2015).
Similarhelpingbehaviorfromdogstohumanswaspreviouslyfound(forreview,Bruer2015).
Theseresultsshowthepossibilityofdogs'altruisticbehavior.
Morestudiesontheextentofthiskindofbehavior.
Overall,thestudiesshowemotionalcontagionfromhumansacrossspeciestodogs,aswellasfromdogstotheirconspecifics,althoughtheunderlyingmechanismsofcontagionarecurrentlynotclear.
AnimalSentience2017.
013:KujalaonCanineEmotions1412.
Yawningcontagiousnessindogs:EmpathyorsocialrelaxationContagiousyawningisnotusedasameasureofempathyinhumanpsychology,sincewell-validatedquestionnaires(e.
g.
,Mehrabian&Epstein1972,Davis1980,Lawrenceetal.
2004,Daddsetal.
2008)canbecombinedwitheitherbehavioralstudies(especiallyinchildren,seee.
g.
,Eisenberg&Miller1987,Eisenberg&Fabes1990,Roberts&Strayer1996,Eisenbergetal.
1999)orbrainimagingstudies(forreviews,Decety&Jackson2004,Bernhardt&Singer2012)toshowacorrespondencebetweenself-reportedreactivityandbehavioralorphysiologicalchanges.
EarlystudiesreporteddiminishedcontagiousnessofyawninginchildrenwithAutismSpectrumDisorders(ASD)(Senjuetal.
2007,Giganti&EspositoZiello2009,Heltetal.
2010),whichpromptedspeculationregardingthepossiblerelationofcontagiousyawningtoempathy.
NewerstudieshavereportedthecapacityforcontagiousyawninginchildrenwithASD(Senjuetal.
2009,Usuietal.
2013)andtheindependenceofyawningcontagiousnessfromempathy(Bartholomew&Cirulli2014).
However,sincetheinauguralstudyshowingyawncontagionacrossspeciesfromhumanstodogs(Joly-Mascheronietal.
2008),arangeofstudieshasexploredthepossibleconnectionofcontagiousyawningandempathyincanines(O'Hara&Reeve2011,Silvaetal.
2012,Madsen&Persson2013,Romeroetal.
2013,Silvaetal.
2013,Buttner&Strasser2014,Romeroetal.
2014).
Yawningisasomewhatproblematicmeasureindogsbecausethecanineyawnservesasatension-releasingstressresponse(seee.
g.
,Beerdaetal.
1998).
Nevertheless,manystudiesfoundahigherfrequencyofcanineyawningafterobservingorhearingahumanyawn(Joly-Mascheronietal.
2008,Silvaetal.
2012,Madsen&Persson2013,Romeroetal.
2013).
Somestudiesalsoreportedstrongeryawncontagiousnessindogsafterperceivingafamiliarratherthannon-familiarpersonyawning(Silvaetal.
2012,Romeroetal.
2013).
Thiseffectofsocialconnectednesswasalsodemonstratedwithinwolves(Romeroetal.
2014).
Inhumans,thetendencytoyawnafterwitnessinganotherperson'syawnisnegativelycorrelatedwithamygdalaactivation(Schürmannetal.
2005).
Thus,higheramygdalaactivation,whichmayoccurnaturallyinunknowncompanyasvigilanceforathreat(e.
g.
,Whalen1998,Hartetal.
2000),actsagainstyawningcontagion.
Ifasimilarconnectionexistedincanids,familiaritywouldincreasecontagiousyawningmerelyasafunctionoftheindividual'slevelofsocialrelaxedness.
Theyawncontagiousnessindogs,however,addstothepossibilityofinterspecificemotionalsynchronizationandcontagioninrelaxation(Odendaal&Meintjes2003,Milleretal.
2009,Nagasawaetal.
2009,Handlinetal.
2011,Nagasawaetal.
2015).
Altogether,thestudiesofempathyshowthatatleastemotionalcontagionispossibleindogs.
Studiesoncaninepost-conflictaffiliativebehaviorandprosocialityalsosuggestsomeempathicconcern.
Arecentstudyoncanineprosocialtendencies(Quervel-Chaumetteetal.
2015)alsoreportedsituation-dependentperspective-taking.
Theextentofempathiccapacityindogsisunknown,however(fordiscussion,seealsoSilva&deSousa2011).
Thereareprobablylimitstothecognitivecomponentofempathyindogs:Theylackmeta-representationalandself-representationalskillsbecausetheirbrainshavelessencephalizationandconnectivitythanhumans.
However,coupledwiththerudimentarytheoryofmindhypothesizedbyHorowitz(2011),empathicrespondingmaynotbeanall-or-nonefunctionbutanabilitythatoccurstovariousdegreesacrosssocialspecies.
AnimalSentience2017.
013:KujalaonCanineEmotions1513.
Long-termmoods:A"cognitivejudgmentbias"testTheeffectsofrearingenvironmentonemotionalresponseinanimalswerenotedinthe1950sbypsychologistssuchasHebb(1955).
Raisingdogsinarestrictedenvironmentaffectedtheirsubsequentemotionalresponses(Melzack1954).
Fiftyyearslater,theworkwasextendedintothefieldofanimalwelfare,andtheeffectoftheenvironmentonthepositiveornegativeaffectivestatesinanimalswasstudiedusinganemotionaljudgmentbiastest(Hardingetal.
2004).
Theeffectiscalled"cognitivejudgmentbias"or"cognitivebias"intheanimalsciences(Mendletal.
2010a,Rygulaetal.
2015),althoughhumanpsychologyhasamultitudeofdifferentcognitivebiases(seee.
g.
,Tversky&Kahneman1974,Haseltonetal.
2005).
Thisexpectation-related"cognitivebias"isalsobasedonstudiesofhumansinwhichthephenomenoniswidelyknownasaffectivecongruence(e.
g.
,Bower1991,Fazio2010):anxiousordepressedpeopletendtointerpretambiguousstimulinegatively(Eysencketal.
1991,Wright&Bower1992,MacLeod&Byrne1996,Gotlib&Krasnoperova1998).
Non-humananimalsmayalsobebiasedintheirexpectationsafternegativeorpositiveexperiences(Mendletal.
2010b).
Thebasictestissimple:animalsarefirsttrainedthatonestimulus(e.
g.
,ablackcard)signalsapositiveevent(reward,e.
g.
,food),andanother(awhitecard)signalsanegativeevent(apunishment).
Aftersuchtraining,theyarepresentedwithambiguous,intermediatesignals(e.
g.
,agreycard),andtheirreactions(e.
g.
,timeofapproachingthestimulus)totheambiguoussignalsarerecorded.
Indogs,afoodbowlisplacedinonecornerofaroomandanemptybowlinanothercorner(Mendletal.
2010a).
Whendogslearntodiscriminatethetwolocations,abowlisplacedbetweenthem.
Inthetesttrials,approachtimetotheambiguouslocationsismeasured:aquickapproachindicatesanticipationoffood,an"optimistic"judgment,andaslowapproachindicatesa"pessimistic"judgment(Mendletal.
2010a).
Dogswithhigherseparation-relatedanxietyapproachtheambiguousbowllocationsmoreslowly,showinganegativecognitivebias(Mendletal.
2010a).
Anxiolyticmedicationwiththehumananti-depressantfluoxetinecombinedwithbehavioralmodificationdiminishesthebias(Karagiannisetal.
2015).
However,leavingdogsaloneforabrieftimedoesnotgeneratenegativeexpectations(Mülleretal.
2012),suggestingthatitisaprolongedemotionalstatethatinducesnegativebiasindogs(Mendletal.
2010a).
Likewise,brieflysearchingfortreatspriortotestingwasnotenoughtoinducepositivebiasindogs(Burmanetal.
2011),whereasadministeringoxytocinpriortotestingcausedpositivebias(Kisetal.
2015).
Todate,itseemsthatthecognitivebiastestmeasureslong-termtendencies,expectations,andmoodsratherthansuddenemotions.
Inducingthepositiveornegativeexpectationsexperimentallyhasproventricky,leavingroomforfurtherinvestigation.
AnimalSentience2017.
013:KujalaonCanineEmotions1614.
Emotionalandsocialdataacrossspecies:EmotionalevolutionAlthoughdirectdataondogemotionsarecurrentlyquiterare,therearemorestudiesondogskillsininterspecificcooperative-communicativesocialtasks.
Inthesetasks,parallelexperimentsindogsandothercanids,carnivores,andnon-humanprimateshavebeenextremelyinformative.
Regardingevolutionarychangesinemotionleadingtochangesinsocialcognition,Hare(2007)hassuggestedthat"dogs'specializedsocial-problem-solvingskillsmayhavefirstappearedaftersystemsmediatingfearandaggressionwerealtered"(p.
62).
Inthelong-termstudiesonexperimentaldomesticationoffoxes,selectivebreedingforlowlevelsoffearandaggressiontowardhumanswereassociatedwithchangesinfoxes'limbicsystems(Trut2001),andasaside-effect,theirsocialcognitiveabilities(Hareetal.
2005).
ThisEmotionalReactivityHypothesis(Hare&Tomasello2005)wasrecentlytestedwithdogsandwolvesbyRange,Ritter,andViranyi(2015).
Inacooperativesituation,wolveswerenotmoreaggressivetowardsconspecificsthandogs.
Thus,themodifiedCanineCooperationHypothesisisthatdog-humancooperationmighthaveoriginatedfromwolf-wolftoleranceandcooperation(Range&Viranyi2014,Rangeetal.
2015).
Thesuggestionthatthereisinterplaybetweenemotionalandsocialskillsindogsisintriguing,andmeritsacloserexaminationofthebraincircuitriesinvolved.
15.
MethodologicaladvancesandfuturedirectionsHumanandnonhumanemotionshavealonghistoryofbeingstudiedusingdifferentmethods(Berridge2003).
Therefore,usingcomparablemethodsinhumansanddogsshouldprovidevaluablenewinsights(forexample,Raccaetal.
2012,Andicsetal.
2014,Trnqvistetal.
2015,Yong&Ruffman2016).
Methodologicaladvancesincludenon-invasivebrainimagingusuallyusedwithhumans,suchasfunctionalmagneticresonanceimaging(Bernsetal.
2012,Andicsetal.
2014,Jiaetal.
2014,Bernsetal.
2015,Dilksetal.
2015,Cuayaetal.
2016)andsurface-electroencephalography(Howelletal.
2012,Kujalaetal.
2013,Trnqvistetal.
2013,Kisetal.
2014).
Thesecannowbeusedwithdogstogetherwithpositiveoperant-conditionaltraining.
Thermographicimagingalsoappearspromisingfordetectingemotionally-stimulatedchangesinbodysurfacetemperature(Travainetal.
2015,Riemeretal.
2016,Travainetal.
2016).
Allthesenewtechniquesrequirecarefulexperimentationtoavoidthepossibleconfoundsreportedforhumanresearch(importantly,seeBennettetal.
2009,Kriegeskorteetal.
2009,Poldrack&Mumford2009,Vuletal.
2009).
Numeroustopics,suchastheoryofmindindogs,andpossibleemotionallateralizationorgendereffects,deservedmorediscussion.
Manypersonalandenvironmentalfactorsunderlieindividualdifferencesinhumanemotionalprocesses(e.
g.
,Tomarkenetal.
1992,Canlietal.
2002,deRosnay&Harris2002,Gross&John2003),andsimilarvariationalsooccursindogs(seee.
g.
,Goslingetal.
2003,Fratkinetal.
2013,Cooketal.
2014).
Dominancerelationsmayaffectcanineemotionsthroughcerebralneurochemicalconcentrations(forreview,Chichinadzeetal.
2014).
Skullshapeindogscanalsoaffectbrainformationandhencecognitionandemotion(McGreevyetal.
2004,Helton2009,Robertsetal.
2010,McGreevyetal.
2013).
Thereisalsonoreasondogscouldnothaveuniqueemotionalstatesthathumansdonothave,forexample,statesrelatedtotheirolfactoryworldandthefunctionofthepiriformcortex.
Thesetopicswillcertainlyreceivemoreattentioninthefuture.
WeneedAnimalSentience2017.
013:KujalaonCanineEmotions17morestudiesoftheemotionalworldofdogsalongwithsensiblecautionininterpretingandgeneralizingtheresults.
AcknowledgmentsThisworkwasenabledbytheBrainResearchcollaborationbetweentheAaltoUniversityandUniversityofHelsinki(BRAHEinitiative);EmilAaltonenFoundationPersonalGrant(#160121);andTekesgrant#7244/31/2016.
Inadditiontotheeditorsandreviewers,Iwishtothankmycolleaguesfromvariousfieldsinthehumanandanimalsciencesfortheircommentsonanearlierdraftofthearticle:OttoLappi,SanniSomppi,HelenaTelknranta,RiittaHari,JariHietanen,JukkaLeppnen,OutiVainio,andKatriinaTiira.
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htmlUQM/ISCCognitiveScienceSummerSchoolJune26-July62018,Montreal,CanadaTheOtherMindsProblem:AnimalSentienceandCognitionOverview.
SinceDescartes,philosophersknowthereisnowaytoknowforsurewhat—orwhether—othersfeel(noteveniftheytellyou).
Science,however,isnotaboutcertaintybutaboutprobabilityandevidence.
The7.
5billionindividualmembersofthehumanspeciescantelluswhattheyarefeeling.
Butthereare9millionotherspeciesontheplanet(20quintillionindividuals),fromelephantstojellyfish,withwhichhumanssharebiologicalandcognitiveancestry,butnotoneotherspeciescanspeak:Whichofthemcanfeel—andwhatdotheyfeelTheirhumanspokespersons—thecomparativepsychologists,ethologists,evolutionists,andcognitiveneurobiologistswhoaretheworld'sleadingexpertsin"mind-reading"otherspecies—willprovideasweepingpanoramaofwhatitfeelsliketobeanelephant,ape,whale,cow,pig,dog,bat,chicken,fish,lizard,lobster,snail:Thisgrowingbodyoffactsaboutnonhumansentiencehasprofoundimplicationsnotonlyforourunderstandingofhumancognition,butforourtreatmentofothersentientspecies.
GregoryBerns:DecodingtheDog'sMindwithAwakeNeuroimagingGordonBurghardt:ProbingtheUmweltofReptilesJonSakata:AudienceEffectsonCommunicationSignalsPANEL1:Reptiles,BirdsandMammalsWORKSHOP1:KristinAndrews:The"Other"Problems:Mind,Behavior,andAgency9SarahBrosnan:HowDoPrimatesFeelAboutTheirSocialPartnersAlexanderOphir:TheCognitiveEcologyofMonogamyMichaelHendricks:IntegratingActionandPerceptioninaSmallNervousSystemPANEL2:Primates,VolesandWormsWORKSHOP2:JonathanBirch:AnimalSentienceandthePrecautionaryPrincipleMalcolmMacIver:HowSentienceChangedAfterFishInvadedLand385MillionYearsAgoSarahWoolley:NeuralMechanismsofPreferenceinFemaleSongbirdSimonReader:AnimalSocialLearning:ImplicationsforUnderstandingOthersPANEL3:SeatoLandtoAirWORKSHOP3:StevenM.
Wise:NonhumanPersonhoodTomokoOhyama:ActionSelectioninaSmallBrain(DrosophilaMaggot)MikeRyan:"CrazyLove":NonlinearityandIrrationalityinMateChoiceLouisLefebvre:AnimalInnovation:FromEcologytoNeurotransmittersPANEL4:Maggots,FrogsandBirds:FlexibilityEvolvingSPECIALEVENT:MarioCyr:PolarBearsColinChapman:WhyDoWeWanttoThinkPeopleAreDifferentVladimirPradosudov:ChickadeeSpatialCognitionJonathanBalcombe:TheSentientWorldofFishesPANEL5:Like-MindednessandUnlike-MindednessWORKSHOP5(part1):GaryComstock:ACow'sConceptofHerFutureWORKSHOP5(part2):Jean-JacquesKona-Boun:PhysicalandMentalRiskstoCattleandHorsesinRodeosJoshuaPlotnik:ThoughtfulTrunks:ApplicationofElephantCognitionforElephantConservationLoriMarino:WhoAreDolphinsPANEL6:MammalsAll,GreatandSmallLarryYoung:TheNeurobiologyofSocialBonding,EmpathyandSocialLossinMonogamousVolesWORKSHOP6:LoriMarino:TheInconvenientTruthAboutThinkingChickensAndrewAdamatzky:SlimeMould:CognitionThroughComputationFrantisekBaluska&StefanoMancuso:WhataPlantKnowsandPerceivesArthurReber:ANovelTheoryoftheOriginofMind:ConversationsWithaCaterpillarandaBacteriumPANEL7:Microbes,MoldsandPlantsWORKSHOP7:SuzanneHeld&MichaelMendl:PigCognitionandWhyItMattersJamesSimmons:WhatIsItLikeToBeABatDebbieKelly:SpatialCognitioninFood-StoringStevePhelps:SocialCognitionAcrossSpeciesPANEL8:SocialSpaceWORKSHOP8:TobeannouncedLarsChittka:TheMindoftheBeeReuvenDukas:InsectEmotions:MechanismsandEvolutionaryBiologyAdamShriver:DoHumanLesionStudiesTellUstheCortexisRequiredforPainExperiencesPANEL9:TheInvertebrateMindWORKSHOP9:DelciannaWinders:NonhumanAnimalsinSportandEntertainmentCareltenCate:AvianCapacityforCategorizationandAbstractionJenniferMather:DoSquidHaveaSenseofSelfSteveChang:NeurobiologyofMonkeysThinkingAboutOtherMonkeysPANEL10:OthersinMindWORKSHOP10:TheLegalStatusofSentientNonhumanSpecies