Positive Emotion and the Brain
UniversityPressScholarshipOnline
OxfordScholarshipOnline
PositiveEmotion:IntegratingtheLightSidesandDark
Sides
JuneGruberandJudithTedlieMoskowitz
Printpublicationdate:2014
PrintISBN-13:9780199926725
PublishedtoOxfordScholarshipOnline:April2014
DOI:10.1093/acprof:oso/9780199926725.001.0001
PositiveEmotionandtheBrain
TheNeuroscienceofHappiness
TabithaKirklandTurowski
VincentY.Man
WilliamA.Cunningham
DOI:10.1093/acprof:oso/9780199926725.003.0007
AbstractandKeywords
Happinessisoftensought,butthemechanismsthatunderliehappinessandpositive
emotionareonlybeginningtobeunderstood.Inthischapter,wesuggestthatcurrent
conceptsandtheoriesofpositiveemotioncanbenefitfromaperspectiveinformedby
affectiveneuroscienceduetothenon-relianceonself-reportandtheabilitytospecify
mechanism.Weexploretheneuroscientificbasisofpositiveemotionandhappiness
throughtheorganizationallensoftwobroadthemes:(1)rewardversusthreatsensitivity
and(2)motivatedversushedonicbehavior.Specifically,wereviewtheliteratureon
reward,whichfocusesonbehaviorfollowingstimuluspresentation,includingapproach
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Positive Emotion and the Brain
behaviortowardrewards,aswellastheliteratureonmotivationalversushedonic
processes,whichfocusesonthedifferentiationbetweenmotivationtogainpositivestimuli
andtheenjoymentofthosestimuli.Wepresentevidencethattheneuralmechanismsthat
contributetohappinessareseenthroughoutnearlyallstagesofprocessing,rangingfrom
veryearlyperceptualprocessestoadvancedreflectiveprocessing.Moreover,these
“happinesses”arelinkedtodifferentaspectsofpsychologicalfunctioning.Weconcludeby
discussinganorganizingframeworkforfutureresearch.
Keywords:anticipation,happiness,hedonism,neuroscience,pleasure,reward,savoring,well-being
Whatis“happiness”?Happinesshasbeendefinedinvaryingwaysthroughouthistoryin
termsofitspredictors(Kesebir&Diener,2008),suchasenjoymentofbeauty(Plato,
1999)orlivingvirtuously(Aristotle,1992).However,scientificresearchstilllacksaclear
definitionofhappinessthatcanexplainitsmanycausesandeffects.Multipleformsof
positiveaffectandpositiveemotionarediscussedusingthiscommonterm.Moreover,
exploringthebroaderconceptofhappinesscanhelptoorganizethewaywe
conceptualizeanddiscusspositiveemotion.
Psychologicalresearchhasassuacprofthathappiness(oftencalled“subjectivewellbeing”)isaunitaryconstruct:ratherthanquestioningtheinnerworkingsofhappiness
itself,researchershavefocusedinsteadonansweringquestionssuchashowtoattain
happiness(e.g.,Diener&Seligman,2002)andhowhappinessinfluencesvarious
psychologicalprocessesandbehavioraloutcomes(e.g.,Lyubomirsky&Ross,1999).The
recentadventofneuroscienceandfunctionalneuroimaginginpsychologyhasthe
potentialtoexplorethesequestionssurroundingthenatureofhappinessandpositive
emotioninincreasinglyrigorousandobjectiveways.
Inthischapter,webeginbybrieflysummarizingbehavioralresearchaddressing
psychologicalquestionsabouthappiness,includingtheproblemofdefininghappiness.
Next,wesuggestthatcurrentconceptsandtheoriesofhappinesscanbenefitfroma
neuroscientificperspectivebyspecifyingbiologicalmechanismsandresolvingproblems
relatedtoself-report.Wereviewthehistoryofpositiveneuroscienceandthe
neuroscientificbasisofpositiveaffectthroughtheorganizationallensoftwobroad
themes:reward/threatsensitivityandmotivational/hedonicprocesses.Finally,weuse
neuroscientificevidencetoprovideanorganizingframeworkforfutureresearch.
Specifically,weoutlinetwoprocessingstylesthatmaybeusedwhenmakinginferences
aboutone’sownhappiness:perceptualprocessing,whichreliesoninputfromthe(p.96)
externalworld,andreflectiveprocessing,whichreliesoninternalthoughtsand
sensations.Takentogether,thisframeworkhelpstoexploreourcurrentunderstanding
oftheneurobiologyofpositiveaffectandguidefutureresearchontheconceptualization
ofhappinessandwell-being(seeBox7.1).
Box7.1:ClarifyingTerminology
•Affect:atemporarysubjectiveexperienceofvalence(positive/negative)with
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Positive Emotion and the Brain
someaccompanyingdegreeofarousal;positiveaffectencompassestheclassof
feelingsexperiencedaspositivelyvalenced
•Emotion:anorganizedsetofcognitive,physiological,and/orbehavioralchanges
inresponsetoastimulus;includesthesubjectiveexperienceofvalence
(positive/negative)withsomeaccompanyingdegreeofarousal
•Happiness:agestaltinterpretationofone’sfavorablestandingrelativetothe
world;basedinaffectiveexperienceandconstructedthroughmultiplecognitive
mechanismsthatdifferacrossindividuals;thetraitformisfacilitatedbycognitive
flexibility
•Hedonic:oforrelatedtopleasure(see“pleasure”)
•Pleasure:aconsciousexperienceofenjoymentbasedoncurrentsensoryinput
•Reward:anystimulusthat,whenpresentedfollowingabehavior,increasesthe
futurelikelihoodofperformingthatbehavior
•Savoring:theprocessofdirectingone’sattentiontopositiveexperiencesto
prolongand/orintensifythem
•Subjectivewell-being:see“happiness”
ThePsychologyofHappinessandPositiveAffect:ABriefOverview
Researchinthedomainofhappinessandpositiveaffect/emotionhasoftenfocusedonhow
tobehappy(orhappier).Thisworksuggests,forexample,thatexperiencesaremore
likelytomakepeoplehappythanarematerialgoods(VanBoven&Gilovich,2003);
focusingonmakingothershappybringsmorepositiveaffectthanfocusingononeself
(Dunn,Aknin,&Norton,2008),and,indeed,havinggoodrelationshipswithothersis
essentialforhappiness(Diener&Seligman,2002).Researchhasalsoarticulated
correlatesofhappiness,suchasindividualdifferences(age,personality)andobjectivelife
circumstances(income,socialrelationships;forareview,seeDiener,1984).Other
prominenttheorieshavefocusedonspecificpsychologicalneedsthatmustbefulfilledas
aprerequisiteofhappiness(Ryan&Deci,2000;Ryff&Singer,1996),suchasautonomy
orself-acceptance.Insummary,researchhasbeeneffectiveindemonstratingthesocial
psychologicalfactorsunderlyingincreasedpositiveaffectandhappiness.
Happinessinandofitselfhasalsobeendemonstratedtohavemanybeneficialeffectson
psychological,social,andhealthoutcomes.Forexample,happierpeoplearemore
creative(Isen,Daubman,&Nowicki,1987),moreoptimistic(Campbell,1981),and
viewedmorefavorablyby(p.97) others(Diener&Fujita,1995;Schimmack,Oishi,
Furr,&Funder,2004)thantheirless-happycounterparts.Happierpeoplearealsoin
betterhealth(Richmanetal.,2005;Pressman&Cohen,2005)andmayhavegreater
longevity(Danner,Snowdon,&Friesen,2001;Diener&Chan,2011).Researchsuggests
thatthecognitivestylesofhappierpeoplehelptomaintaintheirsunnydispositions:for
example,happierpeopletendtocasteventsandsituationsinamorepositivelight,are
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Positive Emotion and the Brain
lessresponsivetonegativefeedback,andmorestronglydenigrateopportunitiesthatare
notavailabletothem(Lyubomirsky&Ross,1999).Thus,happierpeoplemayhaveselfenhancingattributionalstylesthatcontributetotherelativestabilityoftheirhappiness.
Indeed,arecentmeta-analysisof293samplesconcludedthathappinesscausessuccess
andhealthasmuchasitreflectstheseoutcomes(Lyubomirsky,King,&Diener,2005),
suggestingacycleofpositivitywherebypositiveexpectationsleadtopositive
experiences,therebyreinforcingthoseexpectations(Fredrickson&Joiner,2002).
Happiness,then,isbothacauseandaconsequenceofavarietyofpositivelifeoutcomes.
Yet,theconceptofhappinessmaybecurrentlytoopoorlyspecifiedtohaveacomplete
understandingofwhatitmeansto“behappy”orwhatspecificmechanismisdrivingits
beneficialeffects.Therefore,itisnecessarytoexplorehowtodefinehappinessitself.
Happinessismostcommonlydelineatedintoaffectiveandcognitivecomponents.Some
psychologicaltheoriessuggestthatpositivefeelingsaremeaningfullydifferentfrommore
globalevaluationsofone’slife.Forexample,Dienerandcolleagues(Diener,Suh,Lucas,
&Smith,1999)suggestthathappinesscanbeseparatedintohighpositiveaffect,low
negativeaffect,andlifesatisfaction.Thefirsttwocomponentsarecharacterizedasbeing
primarilyaffective,whilethethirdcomponentismorecognitiveinnature.Similarly,
Seligman(2002)suggeststhatastateofflourishingiscomprisedofpositiveemotion,
activeengagementwithactivities,andhavingmeaningorpurposeinlife(affective,
behavioral,andcognitivecomponents),andRyanandDeci(2001)articulateamodel
basedonAristotle’ssuggestionofhedonia(affective—enjoymentoflife)andeudaimonia
(cognitive—meaninginlife)astwodifferentdefinitionsofwell-being.Thesedefinitionsare
broadandvaried,andaneuroscientificperspectivecanbehelpfulforgainingaclearer
pictureoftheessentialaspectsofhappinessaswellasspecifyingthemechanisms
involved.
ANeuroscientificPerspective
Researchtodatehasbeenvaluableinprovidingevidencethatdifferentsubjective
experiencescontributetothegestaltfeelingofhappiness.However,westilllack
specificityregardingthedefinitionofhappinessitself;wecannottrulyunderstandwhy
somethingworkswithoutfirstunderstandinghowitworks.Theproposed
subcomponentsofhappiness(e.g.,affectandcognition)arenotdefinedbeyonda
subjectivelevel,solinksbetweentheseconstructsandtheirunderlyingneurobiology
remainunspecified.Totheextentthatpsychologicalconstructsunderlyinghappinessare
tobetakenseriouslyasmechanisms(ratherthanconceptsthathavetheirgreatestvalue
atthelevelofsubjectiveexperience),theyshouldmapclearlyontobiologicalprocesses.
Inotherwords,afully-developedtheoryofhappinessshouldspecifynotonlythe
psychologicalmechanisms,butalsothebiologicalsystemslinkedtothosemechanisms.
Further,mostresearchpresupposesthathappinessitselfisasingleconstructthat(a)
canbeoptimizedand(b)isoptimizedinthesamewayforallpeople.Inotherwords,this
researchassumesthathappinessisthesame—inbothexperienceandprocess—for
everyone.Thisassumptionmaybeunjustified.(p.98) Peopledifferinmanymeaningful
ways,butthesedifferencesareoftenignored(Wegner&Gilbert,2000);averaging
acrossindividualscanpotentiallyfailtocaptureorevencanceloutconsequential
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individualdifferences(e.g.,Cunningham&Kirkland,2013).Differencesbetween
individualsmaybefunctionallyadaptivegivenaparticularsocialenvironment;for
example,inapervasivelyunsafeorunpredictableenvironment,sensitivitytolossesand
threat(e.g.,Higgins,1997)maybethemostadaptiveorientation.Giventremendous
individualvariability,itdoesnotnecessarilyfollowthateveryoneexperienceshappiness
inthesameway(e.g.,Barrett,2009).However,behavioralresearchtypicallymeasures
happinessinonlyoneway—selfreport—anditsdefinitionisnottypicallyspecifiedbythe
researcher,soitislefttoparticipantstointerpret“happiness”howevertheywish.This
methodcanmakeitdifficulttobescientificallyrigorous.Assuch,aneuroscientific
perspectivecanbehelpfulforaddressingsomeoftheseconcernsandprovidingamore
objectivewayofspecifyingthemechanismsinvolvedintheexperienceofhappiness.
Researchinvestigatingtheneurobiologyofhappinesshasblossoacprofinthepasttwenty
years,focusingontheneurophysiologicalandneurochemicalsubstratesofdifferent
behaviorsreflectingpositiveaffect.Integratingthisperspectivewithbehavioralresearch
hasthepotentialtousefullyaddresssomeoftheconcernsoutlinedabove.Therefore,we
turnnexttoanoverviewoftheneuroscientificliteraturerelevanttothestudyof
happinessandwell-being.
TheFunctionalNeuroscienceofPositiveAffect
Varioustheoristshavehaddifferentideasabouthowbesttocaptureorstudypositive
affectivefeelingsfromaneurobiologicalperspective.Althoughthisliteraturetendsto
focusonstatemanipulationsorexperiencesofaffect,linksaremadetotraitaffectwhere
appropriate.Twobroadthemesunderliethisresearch:(1)thestudyofreward
sensitivity(versusthreatsensitivity),and(2)thestudyofmotivationalandhedonic
processes,includingtheneuroscienceofpleasure.Thestudyofreward/threatsensitivity
focusesonbehaviorfollowingstimuluspresentation,includingapproachbehaviortoward
rewards(and,conversely,avoidancebehaviortowardthreats).Bycontrast,thestudyof
motivational/hedonicprocessesfocusesonthetemporalaspectsofsubjectiveexperience
relatedtoastimulus,differentiatingbetweenmotivationtogainpositivestimuliandthe
enjoymentofthosestimuli.Despitetheirseparatepresentation,thesetwoliteraturesare
relatedandlinksbetweenthetwoaredrawnwhereappropriate.Themajorstructures
involvedacrossbothofthesethemesareshowninFigure7.1.
RewardSensitivity
ApproachandAvoidance:BehaviorActivationandInhibition
Rewardofteninducespositiveaffect.Indeed,oneofthemostcommonexperimental
methodsforinducingpositiveaffectisthroughgivingareward(e.g.,anunanticipatedgift)
toparticipants.Rewardsandpunishers(orthreatsofpunishment)oftenguidebehavior:
anextensiveliteratureusingbothhumansandnonhumananimals(e.g.,rats)hasshown
thatorganismstend(p.99)
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Figure7.1 :Majorstructuresandneurotransmittersinvolvedin
processingpositiveaffectandemotion.Structures:NAcc=nucleus
accumbens;OFC=orbitofrontalcortex;PFC=prefrontalcortex;
VTA=ventraltegmentalarea.Neurotransmitters:DA=dopamine;
OP=opioids.
toapproachrewardsandavoidorescapepunishers(Rolls,2000).Aneurobiological
approachtothestudyofrewardandthreatdescribesthebasicsystemsunderlying
approachandavoidancebehavior.Accordingtothismodel,thebehavioralactivation
system(BAS)representsrewardsensitivityandgovernselationandapproachmotivation
andbehavior.Bycontrast,thebehavioralinhibitionsystem(BIS)representsthreat
sensitivityandgovernsanxiety,caution,andvigilanceforthreat(Gray,1991).Athird
system,thefight-flight-freezesystem(FFFS),maygovernavoidantresponsestothreat
(Gray&McNaughton,2000).Insituationsofthreat,BISassessesthesituationand
resolvesconflictbyengagingeithertheBAStoapproachortheFFFStoescapeas
appropriate,withsomepreferentialtendencytowardengagementofFFFS(escape)
throughanegativitybias.
Individualsdifferintheirdegreeofresponsivenessofthesesystems,reflectingvariation
inthebiologicalresponsesthatunderliestabledifferencesinpositiveemotionality(Gray,
1994;Smillie,Pickering,&Jackson,2006).Forexample,dopaminemayplayanimportant
roleintheneuralcommunicationthatsupportstheBASsystem.Dopamine,likeother
neurotransmitters,isachemicalsecretedbythebrainthathelpsdifferentneural
structurestocommunicate.TheBAShasbeenlinkedtodopaminergicpathways(i.e.,
neuronsthattransmitdopaminetooneanother)ascendingfromtheventraltegmental
areatohigher-levelstructures(Depue&Zald,1993;Winters,Scott,&Beevers,2000).
Individualswhoaremorebiologicallysensitivetorewardstendtohaveamore
responsiveBAS(i.e.,morelikelytotransmitdopamine),whichisreflectedasatendency
towardhighpositiveaffectandreward-seeingbehavior.AnunderactiveBASislinkedwith
unresponsivenesstoincentives,lowpositiveaffect,andalackofengagementwiththe
environment(Depue&Zald,1993).
(p.100) RegulatoryFocus
Anotherwayofexaminingresponsestorewardsandthreatsisbystudyingthe
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Positive Emotion and the Brain
subjectiveinterpretationofstimuliasrewardingorthreatening.Accordingtoregulatory
focustheory,thewaypeopleframetheirgoalsinfluenceshowthosegoalsare
experienced(Higgins,1997).Incominginformationcanbeinterpretedindifferentways
dependingonwhetheroneismotivatedtogainrewards(promotionfocus)ortoavoid
punishments(preventionfocus)—inotherwords,toseekorapproachpleasureandto
avoidpain.Preventionfocusisnotnecessarilya“bad”thing;insomesituationsandfor
somepeople,itcanbethemostadaptivewayofreachingone’sgoals.However,thetype
ofpositiveaffectthatisexperienceduponthesuccessfulattainmentofarewardisquite
differentfromthetypeofpositiveaffectthatisexperienceduponthesuccessful
avoidanceofapunishment(Rolls,2000).Importantly,thesameindividualmayexperience
eitherpromotionorpreventionfocusinagivensituation,dependingonhisorher
mindset.Thus,orientationtowardspromotionorpreventionfocuscandifferacross
situations(i.e.,apersonmayexperiencepromotionfocusinonesituationandprevention
focusinanother)andindividuals(i.e.,acrosssituations,somepeopletendtobe
chronicallymorepromotion-focused;others,moreprevention-focused).
Somepositiveaffectderivesfromgainingrewardsandisassociatedwithamotivational
systemthatispreferentiallysensitivetogains(rewards)ratherthanlosses
(punishments).Higgins(1997,1998)suggestedthatthepromotionsystemorientsthe
individualtowardexplorationandopportunity;thepresenceofpositiveinformationisthe
primaryconcernforpromotion-focusedindividuals.Forexample,apromotion-focused
individualrunningahalf-marathonmaybefocusedonreachingthefinishlinewithina
designatedamountoftime;shewillfeelelationifsheaccomplishesthisgoalwithtimeto
spareanddisappointmentifshefails(seeCarver,Johnson,&Scheier,2014).
Justasonetypeofpositiveaffectcanbederivedfromthepleasurableexperienceof
rewardingstimuli,anothertypeofpositiveaffectcanalsobederivedfromavoiding
punishments.Thispositiveaffectisassociatedwithamotivationalsystemthatis
preferentiallysensitivetolosses(punishments)ratherthangains(rewards).Higgins
(1997,1998)suggestedthatthispreventionsystemorientstheindividualtoward
concernsofsafetyandsecurity,withtheabsenceofnegativeinformationbeingthemost
importanttoprevention-focusedindividuals.Forexample,aprevention-focused
individualrunningahalf-marathonmaybefocusedoncompletingtheracewithout
exceedingacertaintime;shewillfeelreliefifsheaccomplishesthisgoalandagitationif
shefails.
Neuroscientificresearchhassupportedthisfunctionaldistinctionbetweenpromotionand
preventionorientation:ourbrainsaredesignedasevaluativesystemsorientedtoward
rewardsandpunishmentsbecausethisisthemostadaptiveorientationforsurvival
(Rolls,2005).
MultipleSystemsforReward
Becauserewardisstronglylinkedwithpositiveaffect,manyofthebehavioralinfluences
ofpositiveaffectareacprofiatedbythesameneuralmechanismsthatacprofiatereward.
Atearlylevelsofprocessing,theamygdalaandnucleusaccumbens(NAcc)1 areboth
heavilyinvolvedintheprocessingofrewardingstimuli(e.g.,Blood&Zatorre,2001;
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O’Doherty,Rolls,Francis,Bowtell,&McGlone,2001;Rollsetal.,2003)through
projectionsfromdopamine-producingcellsintheventraltegmentalarea,whichprojects
totheamygdala,whichsubsequentlyprojectstoboththe(p.101) NAccandthe
hippocampus(Ashby,Isen,&Turken,1999).Theamygdalamayplayaparticularly
importantroleinsignalingthemotivationalvalueofstimuli(Cunningham&Kirkland,2013;
Cunningham,VanBavel,&Johnsen,2008).Researchhasalsoimplicatedtwoadditional
areasasactiveinresponsetopositivestimuliduringpromotionfocusandnegativestimuli
duringpreventionfocus,supportingthismotivationalfitexplanation(Cunningham,Raye,
&Johnson,2005).Theseareasaretheanteriorcingulatecortex,whichisinvolvedin
autonomicregulationaswellashighercognitivefunctionssuchasdecision-making,and
extrastriatecortex,anareasensitivetovisualandmotioncues.
Researchwithhumansandevolutionarilysimilarnonhumananimalshasprovided
increasedinsightintotheroleofhigher-orderregionsofthebrainforprocessing
rewardsandpunishers.Neuralactivitycorrespondingtorewardsandpunishments
becomesincreasinglyintegratedacrosslevelsofprocessing.Beginningwiththeamygdala
andventralstriatumasdiscussedabove,dopaminetransmissionfacilitatesprocessing
throughtheprefrontalcortex,particularlyacprofialorbitofrontalcortex(OFC;
Grabenhorst&Rolls,2001;Kawabata&Zeki,2004).Moreover,dopamineisinvolvedin
severalbrainareasimplicatedinrewarddetection,suchastheacprofialtemporal,
dorsolateral,prefrontal,premotor,andorbitofrontalcortices(Schultz,2000).Inprimates,
theOFCisinvolvedinmakingstimulus-reinforcerassociationstoguidebehaviors
towardsrewardandawayfrompunishment(Rolls,2000).Similarly,inhumans,viewing
promotiongoalsiscorrelatedwithactivationofacprofialOFC(O’Doherty,2004).Further,
activationintheleftPFCtopromotiongoalsisstrongerinindividualswithchronic
promotionfocus(Eddington,Dolcos,Cabeza,Krishnan,&Strauman,2007).Through
denseinterconnectionsfromacprofialOFCtolimbic(e.g.,amygdala)andstriatal(e.g.,
NAcc)regions,incominginformationissharedandinterpretedacrosslevelsof
processing.ReciprocalconnectionsbetweenacprofialOFCandtheselower-order
rewardcircuitsimplythatactivityinacprofialOFCcaninfluencefurtherprocessingat
earlierstages.Forexample,goal-directedbehaviorisdrivenprimarilybyNAcc,which
integratesinformationfrombothamygdalaandprefrontalregions,allowingforthe
convergenceofaffectivelysalientinformationwiththecurrentmotivationalstate(Goto&
Grace,2008).Insummary,informationisintegratedacrossmultiplechannels,beginning
inlower-orderregionssuchastheamygdalaandNAcc,andmovingthroughhigherorderregionssuchastheOFC,whichinturncanreciprocallyinfluenceprocessingin
thoselower-orderregions.Forreward,theseareasseemtorespondmorestronglyto
positiveinformation,signalingthewaysinwhichtheseregionscodeforthemotivational
valueofstimuli.
PredictionofFutureRewards
Theexpectationoffuturerewardscanalsoimpactaffectandbehavior.Researchintothe
systemsinvolvedinreinforcementlearninghasfocusednotonlyoncurrentresponsesto
rewardsandpunishers,butalsothefullertimecourseofresponsesincludingthe
anticipationorpredictionoftheseoutcomes.Whereasanticipationofbothpositiveand
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negativeoutcomesislinkedwithactivationoftheacprofialcaudate,reward-specific
anticipationislinkedwithactivationoftheNAcc(Knutson,Adams,Fong,&Hommer,
2001;Knutson,Fong,Adams,Varner,&Hommer,2001)and/oracprofialOFC(Hare,
O’Doherty,Camerer,Schultz,&Rangel,2008).
Researchhasfurtherexaminedtheneuralresponsestounpredictedoutcomes.
Predictionerrors,inwhichoutcomesaredifferentfromwhatwaspredicted,are
associatedwithactivityintheventralstriatum(Hareetal.,2008).Specifically,dopamine
activityinthestriatummay(p.102) becentraltothecalculationofpredictionerrorsto
informfuturedecision-making(Pessiglione,Seymour,Flandin,Dolan,&Frith,2006);
eventsthataremorerewardingthanpredictedactivatedopamineneurons,whilethose
thatareworsethanpredicteddepressdopamineneurons(Schultz,1998).Other
research,bycontrast,hassuggestedthatprocessingofunanticipatedoutcomesoccurs
inthehighercortices;forexample,inonestudy,theomissionofanexpectedreward
suppressedactivityintheventroacprofialPFC(Knutson,Fong,Adams,Varner,&
Hommer,2001).Clearly,furtherworkremainstobedonetointegratethesepotentially
conflictingfindings.
Motivational&HedonicBehaviors
Asecondneuroscientificapproachtothestudyofpositiveaffecthasarticulatedthe
differentcontributionstopositiveaffectbymotivationalandhedonicprocesses.Whereas
sometypesofpositiveaffectmaystemfromtheanticipationofafuturedelight,other
typesofpositiveaffectmaystemfromtheenjoymentandsavoringofthatdelight.
Researchhasexploredthedifferencebetweenthemotivationtogainrewards(“wanting”
oranticipatorybehaviors)andtheenjoymentofthoserewards(“liking”or
consummatorybehaviors)(Berridge&Kringelbach,2008).Thus,anticipatorybehaviors
includefuture-focusedactionsinvolvedinseekingrewards,whereasconsummatory
behaviorsincludepresent-focusedactionsinvolvedinthepleasurableexperienceofthe
reward(Craig,1918;Gard,Gard,Kring,&John,2006).2Thesetypesofpositiveaffect
areconceptuallysimilartoAristotle’searlyideasofeudaimonia(goalstriving/meaningmaking)andhedonia(enjoyment),respectively.
Neuroscientificinvestigationsofconsummatoryandanticipatorybehaviorshave
suggestedthattheventralstriatum,specificallytheNAcc,mayplayakeyroleinboth
componentsofpositiveaffect(Dillonetal.,2008;Taha&Fields,2005).Theseprocesses
canbedistinguishedby(a)sub-regionswithinNAcc,(b)thespecificbiochemicalcircuits
involved,and(c)interconnectionsbetweenNAccandotherbrainregions.Specifically,
whereastheNAcccoreisorientedtowardanticipatorystates,theNAccshellisoriented
towardconsummatorystates.Thesedifferencesinactivationarefacilitatedbytherelease
ofdopamineandendogenousopioids,respectively.Uniqueneuralsubstratesfor
anticipatoryandconsummatorytypesofpositiveaffectmayalsoindicatephysiological
differences(Baldo&Kelley,2007):whereasanticipatorybehaviorsarelikelyassociated
withregionsinvolvedinprocessingexpectedrewardandingoal-directedbehavior,
consummatorybehaviorsmaybefacilitatedbyactivationinregionsinvolvedwiththe
apprehensionofapleasurablestimulusanditsprocessingforrewardvalue.Inthe
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Positive Emotion and the Brain
followingsections,wedescribetheuniquecontributionstobothanticipatoryand
consummatorytypesofpositiveaffect.
AnticipatoryProcesses
Someneuralprocessesarespecifictothepositiveaffectinducedbytheanticipationofa
futurepositiveevent.ActivationofdopamineinthecoreoftheNAccincreasesmotivated
responsesassociatedwiththeanticipatorytypeofpositiveaffect(Berridge&Robinson,
2003).Forexample,onestudydemonstratedthatlesionstodopaminergicregionsinrats
affectanticipatory,butnotconsummatory,behaviors(Baldo&Kelley,2007).Further,
thisincreasedactivationmaybespecifictotheanticipationofpositivebutnotnegative
outcomes(Knutson,Adams,Fong,&Hommer,2001).Anticipatorybehaviorisalso
associatedwithreciprocalprojectionsbetweenOFCandNAcc(Rolls,1999).Thisis
consistentwiththeaforementionedliteratureonrewardprediction.
(p.103) Humanshaveauniquementalabilitytosimulatehypotheticalfuturesthatare
notimacprofiatelyrelevanttotheirsurroundings.Mentalsimulationofpossiblefutures
andtheiroutcomes,allowsustoplanaheadandmaximizethelikelihoodofpositive
outcomes.Cognitiveelaborationabouthypotheticalfutureevents,asinthecaseof
prospection,hasbeendemonstratedtoengagehigher-ordercorticalregions.Indeed,
someresearchhassuggestedthatgenerationofpossiblefutureeventsengagestheleft
ventrolateralPFCandrightfrontopolarcortex(Addis,Wong,&Schacter,2007),areas
generallyinvolvedinmemoryretrievalandevaluationofinternallygeneratedinformation,
respectively(Christoff&Gabrieli,2000).Arecentmeta-analysissuggestedthatacore
networkofregionsmayunderlieprospectionalongwithmanycognitivestates(Spreng,
Mar,&Kim,2008).Theseareasincluderegionsintheacprofialtemporalandparietal
lobesaswellasthelateralPFC(seealsoCunningham,Haas,&Jahn,2011).Therefore,
anticipatoryprocessescanrangefromverybasic,imacprofiatetypesofanticipationto
moreabstract,long-rangetypes.
Furtherevidenceoftheneuroscientificbasisofanticipatorystatescomesfromemotion
researchgroundedinthereinforcementlearningliterature.Someemotionsrequirean
affectivetrajectorythroughtime—meaningthattheyemergeacrosstimeinresponseto
change(Kirkland&Cunningham,2012).Accordingtothisprocess,psychological
outcomesmayemergefromtheinteractionoftheevaluationsofone’scurrentstate,
predictionsforthefuture,andtheoutcomesthatoneexperiencesafterthesepredictions
(Cunningham&VanBavel,2009;Cunningham&Zelazo,2009;Kirkland&Cunningham,
2011).Wehaveelsewhere(Kirkland&Cunningham,2012)proposedthreeneural
circuitsthatareinvolvedinrepresentingvalenceacrosstime.Predictingfutureevents
involvesacprofialtemporalcortex,amygdala,andbasalganglia(includingNAcc),and
evaluatingoutcomesinvolvestheOFC(Cunningham&Zelazo,2007;Schultz,2000).
Critically,neuralcommunicationamongtherelevantcircuitsallowsforcross-situational
comparisons,allowingustomapoutourparticularplaceintime.Thus,anticipatorystates,
facilitatedbypredictionsforthefuture,involveactivationinandcomparisonsbetween
areassuchasamygdala,NAcc,andmPFC.Thisperspectivealsoallowsforaricher
understandingofthetemporalcontextinwhichapositiveaffectiveeventunfolds,
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Positive Emotion and the Brain
includingcommunicationbetweenlower-andhigher-orderprocessingregions.
ConsummatoryProcesses
Anothertypeofpositiveaffectisinducedbyconsummatorystates,whicharefacilitated
byhedonicresponsestoactivationofopioidsintheshelloftheNAcc(Kelleyetal.,2002;
Peciña&Berridge,2000;seeBerridge&Kringelbach,2008,forareview).Anetworkof
neuralsubstratesthataremodulatedbyopioidtransmission(Smith&Berridge,2007)
includingtheinterconnectionsbetweentheNAccshellandtheventralpallidum(Aldridge
&Berridge,2010)enhanceconsummatory(or“liking”)reactionstoavarietyof
pleasurablestimuli(Berridge&Kringelbach,2008).
Althoughmuchresearchonpleasureandthebrainhasfocusedonanimals,researchon
someuniquelyhumanexperiencescanbringgreaterdepthtoourunderstandingof
consummatoryprocessing.Savoring,theprocessofdirectingone’sattentiontopositive
experiences,isanemotionregulationstrategythatcanbeusedtomaintainorenhance
positiveaffectandpositiveemotions(Bryant,Chadwick,&Kluwe,2011).Savoringcovers
arangeofprocesses,fromappreciatingone’scurrentcircumstancestobeingimmersed
inthem;however,theneurobiologyofsavoringisnotyetwellunderstood.Another
relevantconceptisflow,theexperienceoftask(p.104) immersionassociatedwith
intenseengagementandlackofattentiontootherinformationsuchasthepassageoftime
(Csikszentmihalyi,1990).Weberandcolleagues(2009)haveelaboratedonthisdefinition,
proposingflowasa“discrete,energeticallyoptimized,andgratifyingexperienceresulting
fromcognitivesynchronizationofattentionalandrewardnetworks”(p.397)when
individualsareengagedinacomplextaskatwhichtheyaresufficientlyskilledtoperform
well.Specifically,theysuggestthatpeopleexperiencingflowshowenhancedfunctional
connectivitybetweenareasinvolvedinattention(includingfrontalandparietalcortices,
frontaleyefields,andsuperiorcolliculus)andreward(limbicsystem),supportinga
subjectiveexperienceinwhichhighlevelsofengagementareexperiencedasgratifying.
Neuralprocessesalsosupportthesubjectivedescriptionofflowaseffortless;for
example,oneprojectdemonstratedthatarchersexperiencingaflowstateshowed
activityinareassupportingwell-coordinated,learnedmotoractivityratherthanareas
responsibleforplanningcomplexmotormovement(Ferrell,Beach,Szeverenyi,Krch,&
Fernhall,2006),highlightingtheautomaticityofbehaviorsinaflowstate.Flowmaybeone
typeofconsummatoryprocessbecauseitistheproductofabsorptioninthemoment
withoutreflectiononthefuture.
MultipleFacetsofHappiness
Althoughwehavethusfardiscussedtheneuroscienceofpositiveaffect,considering
happinessasamultifacetedconceptmayhelpbuildamodeloftheheterogeneityof
positiveemotion.Indeed,ratherthanbeinganaturalkind(i.e.,somethingthatexists
independentoftheobserver;Barrett,2006),“happiness”maybeametacognitivelabel
thatpeopleusetointerprettheirongoingsubjectiveexperience.Totheextentthat
peopleusedifferenttypesofinformationasthebasisfortheirinterpretation,different
interpretationsandconclusionsregardinghappinesscanbereached.Critically,thetype
ofinformationthatpeopleuseasthebasisforjudging“happiness”mayvarybysituation
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Positive Emotion and the Brain
and/orindividual.Differentsituationsorpeoplemayhavedifferentcriteriaforthetypeof
informationthatisrelevantand/ornecessarytolabelone’scurrentstateashappiness.
Individualdifferencesmayalsoaffectthislabelingprocess;somepeoplemayberelatively
moreinclinedtolabeltheircurrentstateashappinesswhennothingcatastrophichas
happened,whereasothersmayfocusmoreongainingdesiredends.Specifically,two
generalsubsystemsexistthatindividualscanpoll(indifferentways,atdifferenttimes,
andindifferentamounts)togetacurrentreadontheiranswertothequestion:“How
happyamI?”TheremainderofthisChapteroutlinesthisintegrativeframework.
Peoplehavetwobasicsourcesofpsychologicalinput—externalandinternaltotheself—
thatcorrespondtotwoprocessingstyles.Whereasperceptualprocessingreliesoninput
fromtheexternalworld,reflectiveprocessingoccursindependentofsensorystimulation
andreliesmoreoninternalsensationsandprocesses.Twogeneralsubsystems
correspondingtotheseprocessingstylescanbeusedtoinferhappiness.One
subsystem,sensorypleasure,arisesfromexternalstimulation(i.e.,stimulationofbasic
sensoryreceptors)andisassociatedwithmoreperceptualprocessing.Theother
subsystem,goalorientation,arisesfrominternalstimulation(i.e.,strivingtowardor
achievementofagoalsetbyoneself)andisassociatedwithmorereflectiveprocessing.
Typically,thesesubsystemsarenotexperiencedasindependent;cognitiveprocessing
mixesperceptualandreflectiveprocessingsoadeptlythatsubjectiveexperienceis
holistic(Johnson&Hirst,1991).Bothsubsystemscanbeexaminedfromthe
perspectivesofreward/(p.105) punishmentorientationandmotivational/hedonic
orientation;peoplemayusethisinformationindifferentwaystoreachdifferent
conclusionsabouttheirhappiness.Positiveaffectisthereforere-representedthroughout
theneuroaxis,witheachlevelbecomingprogressivelymorecomplex.
OriginsofPositiveAffect:PerceptualMechanisms
Oneformofhappinesscanstemfromperceptualprocessing.Sensorypleasureisatype
ofpositiveaffectinducedbyexternalstimulation—engagementofone’ssensory
receptors—thatisassociatedwithrelativelymoreperceptualprocessing.Critically,
thoughperceptualprocessescanoccurthroughbothtop-downandbottomup
processing,theyaredependentontheexternalworld.
ThelinkbetweenphysicalpleasureandhappinessdatestoAristotle’sconceptof
hedonia—enjoymentofpleasurablephysicalfeelings.Aristotletookitasagiventhatboth
humansandnonhumananimalsareinclinedtopursuepleasure,andthatpleasureserves
tocompletetheactivitiesthatitaccompanies(forexample,listeningtomusicwouldbea
morecompleteactivityifaccompaniedbypleasurethanifdonewithoutpleasure).Recall
ourearlierdiscussionofsavoring—theprocessofmaintainingandincreasingpositive
affectthroughattentiontopositivestimuli.BryantandVeroff(2007)distinguishbetween
savoringexperiencesinvolvingcognitivereflection,orintrospectionaboutone’s
subjectiveexperience,fromthoseinvolvingexperientialabsorption,astateofsavoringin
whichoneminimizesintrospectioninfavorofperceptualimmersion.Thelatterstrategy
maybeparticularlyeffectiveformaximizingsensorypleasure.Thenatureofsensory
pleasureprocessingiscontingenttosomedegreeonthesourceofthepleasure.Some
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Positive Emotion and the Brain
pleasuresinvolvereceivingrewards,whereasothersinvolveavoidingpunishments.
Sensoryrewardscancomefromsourcesasdiverseasfood,sex,wine,music,andart.
Convergingneurobiologicalevidencesuggeststhatfollowingmodality-specificprocessing
intheearlieststagesofprocessing(e.g.,visual,auditory,ormotorcortex),information
becomesintegratedathigherlevelsofprocessingsuchthatavarietyoftypesofsensory
pleasuresharingsimilarvaluecodinginthebrainareprocessedinthesameregions
(Grabenhorst&Rolls,2011),beginninginthelimbicsystemandcontinuingthroughthe
prefrontalcortex.Specifically,theamygdalaandNAccarebothheavilyinvolvedinthe
processingofrewardingsensorystimuliacrossmodalitiesasdiverseastaste,viewing
art,andhearingmusic(e.g.,Blood&Zatorre,2001;O’Dohertyetal.,2001;Rollsetal.,
2003;Suzukietal.,2008).Rewardinginformationisfurtherintegratedintheprefrontal
cortex,particularlyacprofialOFC,acrossseveralsensorymodalities(Grabenhorst&
Rolls,2011;Kawabata&Zeki,2004).Throughdenseinterconnectionsbetweenacprofial
OFCandregionsofthelimbicsystemsuchastheamygdala,sensoryinformationis
sharedandinterpretedacrosslevelsofprocessing.Thesharedneuralsubstratesamong
avarietyofsensorystimulireflecttheinteractionsbetweentheperceptionofsensory
informationandjudgmentsofthemeaningofthatinformation.
However,focusonpunishmentdoesnotnecessarilyhavetocauseunhappiness.A
secondtypeofperceptualpleasuremaycomefromavoidanceofharm,achievedby
maintenanceofbaselineaffectandone’scurrentphysiologicalstate.Particularlyfor
prevention-focusedpeople(Higgins,1997),thistypeofmaintenanceisimportantand
negativestimuliaremoreprominentastheysignalthreatstobaseline.Avoidance
motivationmanifestsinadesireforthemaintenanceofhomeostasis,alow-arousal,lowvariabilitystatethatsignalscomfortandsafety.Cannon(1929)fraacprofhomeostasisas
theprocessofmaintainingequilibriumratherthana(p.106) staticstate;this
maintenanceinvolvesacontinuousseriesofsmalladjustmentsinresponseto
disturbancesbytheoutsideenvironment.“Asorganismsbecomemoreindependent,”he
suggested,“theydosobypreservinguniformtheirowninnerworldinspiteofshiftsof
outercircumstances”(p.400);thisuniformityisaccomplishedthroughcompensatory
responsestothoseshifts.Inherentinthissuggestionistheideathatflexibilityin
respondingtotheworldfacilitatesmoreefficientstrategies.Subsequentresearchhas
suggestedthatreturningtohomeostasisthroughengagementwithasensorystimulus
canbeexperiencedaspleasurable,withthemagnitudeofthepleasureproportionalto
theabilityofthestimulustoreturnthebodytohomeostasis.Forexample,ahotdrinkor
blanketmaybeexperiencedaspleasurablebysomeonefeelingcold(Cabanac,1971).At
aneurobiologicallevel,researchhasimplicatedtheamygdala,anteriorcingulatecortex,
andextrastriatecortexasactivetonegativestimuliduringpreventionfocus
(Cunningham,Raye,&Johnson,2005).
OriginsofPositiveAffect:ReflectiveMechanisms
Asecondformofhappinesscanarisefromreflectiveprocessing.Goal-orientedpositive
affectisinducedbyinternalstimulation—inputintothesystemthatcomesfromtheself
ratherfromanexternal,sensorystimulus.Reflectiveprocessingoccursindependentof
sensorystimulationandisfocusedoninternalprocesses,includingself-generatedgoals.
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Weconsiderreflectiveprocessingtoencompassthepursuitorachievementofanyselfgeneratedgoal.Thesecanincludeconcretegoals,suchasseekingfoodorshelter;they
canalsoincludemoreabstractgoals,suchassocializing,acquiringknowledge,seeking
novelty,andsoon.Theycanalsoincludepunishment-orprevention-focusedgoals,such
asharmavoidance.Whereasgoalpursuitismorerelatedtoanticipatoryprocessing,goal
achievementismorerelatedtoconsummatoryprocessing.Theseprocessesdifferin
termsofthetimepointonwhichoneisfocused:whereaspursuitisprediction-oriented,
future-focused,andassociatedwithenthusiasmandengagement,achievementis
present-focusedandassociatedwithsatisfactionandsavoring.
TheearlyrootsofreflectivepositiveaffectareinatheorybyAbrahamMaslow(1943),
whosuggestedthatpeople’sneedsareorganizedhierarchicallyandthattheyare
motivatedtofulfillmorebasicphysiologicalneeds(e.g.,hunger,shelter)beforemovingon
toaddresshigher-levelpsychologicalandsocialneeds(e.g.,relationships,knowledge
acquisition).Physiological,security,social,andesteemneedsaredeficiencyneeds,
meaningthattheyariseduetodeprivation.Thehighestlevelofthehierarchycomprises
growthneeds,whichstemfromthedesireforpersonalgrowthratherthandeficiency.
Maslowbelievedthatfulfillmentofneedsmotivatesmostbehaviors.Thus,beyond
strivingforsimpleformsofsensorystimulation,manypeoplestrivetoachievegoalsthat
theysetforthemselves.Goal-orientedpositiveaffectiscomplementarywithAristotle’s
eudaimonichappinessdiscussedabove,whichisoftenconsideredbyphilosophersas
beingamore“complete”formofhappinessinthatitsatisfiesthesocialworldandone’s
moralvirtuesratherthanthe“simpler”(i.e.,sensory)pleasures.
ConsistentwithMaslow’s(1943)ideaofhierarchy,thesegoal-motivatedreflective
processesbuildonthebasicprocessesrelevanttotheperceptualsection.Theessenceof
reflectivepositiveaffectisinitslackofrelianceonexternalstimuli.Ofcourse,agoalmight
befocusedontheattainmentoravoidanceofanexternalstimulus,suchasachievinga
goodgrade,say,ornotgettingsickaftereatingshellfish.However,thegoalmust
precedethestimulus,andthus(p.107) involvesmovementthroughtime(and
anticipatory/consummatoryprocesses)inadditiontosimpleapproach/avoidmotivations.
ConsistentwithCarverandScheier’s(1990)selfcontrolfeedbackloop,goalpursuit
involvesaconstantprocessofcomparingone’scurrentprogressonthegoaltothe
desiredendstateandprovidingfeedbackdesignedtoreducediscrepanciesbetweenthe
twostates.
HierarchicalOrganization
Thesesubsystemsaremeanttobeauseful,heuristicwayoforganizingand
understandingthemanyroutestopositiveaffectandhappiness;itisnotnecessarythat
onlyoneroutebeusedatatime.Althoughthesubsystemshaveuniqueneurobiology,
theylikelyinteractwithoneanothertoproduceasingleexperience.Afundamentaltenet
ofthisframeworkisthatthesesubsystemsareorganizedhierarchically,suchthat
multipleneuralsystemsinteracttorepresentinformationandproducebehaviorsat
increasinglevelsofcomplexity.Whiletheearliestincominginformationmaybesimple
cues,atsubsequenttimepointsinprocessingthesystemcanhandleincreasinglycomplex
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Positive Emotion and the Brain
andsubtleinformation.Morecomplexprocessingcaninformsubsequentrepresentations
ofsimplerprocessesinadynamicandrecursivefashion.Thus,bothsensoryandgoalrelevantinformationcanbeprocessedinmultiplewaysandatmultiplestagesof
interaction.Inthisway,thegestaltimpressionofhappinesscanbeanemergent
interpretationofsubjectiveexperiencebasedonmoresimplecomputationsina
hierarchicalcognitivesystem.Althoughthisdescriptioniscompatiblewithcurrent
neuroscientificbeliefsaboutthecognitiveandneuralarchitectureofthehumanmind,the
ideaofhierarchicalorganizationdatestothe1800s,whenEnglishneurologistJohn
HughlingsJacksondiscussedthecontinuousevolutionarylayeringandre-representation
ofmotorsystemsatprogressivelyhigherlevelsoftheneuroaxis(Jackson,1958).Jackson
suggestedthatbrainregionsareorganizedalongacontinuumwhereinlower-order
regionsareresponsibleforprocessesthataresimple,specializedforspecificfunctions,
organized,andreflexive,whereashigher(frontal)regionsareresponsibleforprocesses
thatarecomplex,general,lessorganized,andvoluntary.Thehighestregionsare
simultaneouslythemostcomplexandtheleaststereotyped,allowingforgreater
processingflexibility.Forexample,whereasspinalandbrainstemreflexesorganize
primitivemovementsandbalance,higher-levelmotorsystemsallowforflexibilityand
graceinmovement.Theanalogycanbemadetodevelopmentalstages:veryyoung
childrenfocusonmasteryoftheroughskillsneededforstanding,walking,andgrasping,
whereasadolescentsarecapableofavarietyofcomplexandsubtlemovements.
Crucially,higher-orderoperationsareinforacprofbyprocessingatlowerlevels(e.g.,
gracefuldanceposesrequiremasteryofbalance),suchthatprocessingincreasesin
complexityandflexibilityasafunctionofbothlower-andhigher-orderareas.Thesame
principlecanbeappliedmoregenerallytoavarietyofprocesses;Jacksonwasanearly
proponentoftheideathatthefrontallobesareresponsibleforavarietyofconscious
cognitiveoperationsrangingfrommemorytoemotion,allofwhichhavetheirrootsin
lower-orderprocesses.
Amorerecentperspectiveistheiterativereprocessingmodel,whichisbasedoncurrent
advancesincognitiveneuroscience(Cunningham&Zelazo,2007).Accordingtothis
model,theevaluativesystemreliesuponmultiplecomponentprocessesthatwork
togethertofacilitatejudgmentsabouttheworld.Stimulisuchaspeople,objects,or
eventsinitiateaniterativesequenceofevaluativeprocessesthroughwhichthestimuli
areinterpretedandre-interpretedinthroughthelensofincreasinglycomplex
representations.Onesetofprocessesisconsistently(p.108) involvedinevaluation,
whereasotherprocessesarerecruitedatdifferentpointsinthecycle.Whereas
evaluationsbasedonfewiterationsofthecyclearerelativelyautomatic(i.e.,reflexiveand
occurwithoutconsciouscontrol),evaluationsbasedonmanyiterationsarerelatively
controlled(i.e.,reflectiveandsubjecttoconsciouschange).Theevaluativesystemis
organizedsuchthatlower-orderprocessescontinuetoprovideaffectiveinformation
evenasadditionalhigher-orderprocessesarerecruited,meaningthatbothlower-and
higher-orderprocesseshavethepotentialtointeractdynamicallytocreateacomplex
representation.
Appliedtothesubsystemsinvolvedintheexperienceofhappiness,theearliestresponses
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Positive Emotion and the Brain
inbothofthesesystemsareimacprofiateandrequirerelativelylessconsciousthought;
laterresponses,bycontrast,willbemorecomplex,andsubtle(e.g.,Cunningham&
Zelazo,2007).AsJackson(1958)alluded,allprocessesinvolvelayersofinteractionwith
otherprocesses,assuringthatnosinglepsychologicalprocesscanbefullyindependent.
Informationmaybesharedacrossthesubsystems,resultinginthepotentialforagiven
stimulustoinfluencetheindividualthroughmultiplesubsystems.Evenwhenastate
beginsasaproductofonesubsystem,itislikelythatadditionalprocessingwillfacilitate
congruentactivationwithinadditionalsubsystemsastheresponsebuildsincomplexity
overtime.Forexample,somesensoryinput,suchastheexperienceofasunnyday,can
beconsideredperceptualatearlystagesofprocessing(warmthonone’sskinis
physicallypleasurable)andreflectiveatlaterstagesofprocessing(onehasbeenlonging
foranicedayafterseveralmonthsofwinter).Further,somebrainareasarerecruited
acrossmultipleprocessessuchastheOFC,forexample,whichhasdemonstratedthe
abilitytocomparedifferenttypesofperceptualstimuli(Montague&Berns,2002)aswell
astocomparerealvs.imaginedstimuli(Cunningham,Johnson,&Waggoner,2011).Itis
likelythatafullydevelopedresponsethatresultsinsubjectivelabelingrequiresinputat
multiplelevels.
Althoughthepresentdiscussionhascenteredontemporaryexperiencesofhappiness,
differentformsoftraithappinesscanarisefromrecurrentstateexperiences.Thatis,
someindividualsmaybemorelikelytodrawupononesubsystemoveranotherasa
basisforconclusionsabouthappiness.Thesesystematicpredispositionscanbetheresult
ofavarietyoffactors,suchasindividualdifferencesingenes,neurotransmitterlevels
andrelevantneurobiologicalsystems.Forexample,individualswithmoresensitive
sensoryreceptors(e.g.,goodvisionorperfectpitch)maybemorelikelytodrawupon
perceptual/sensoryfeedbackasabasisfortheirhappiness.Inthisway,differentspecific
kindsofhappinesscanarise,anddifferenttypesoftrait-happypeoplecanexist.
CognitiveFlexibility
Usinginformationprimarilyfromonesubsystemasabasisforhappinessislesslikelyto
beconsistentlyrewardinginanygivensituationthanstrategicallychoosingthe
subsystemthatismostlikelytoprovidepositivefeedbackinthatsituation.Althoughtraithappypeopleshoulddemonstratethesamecognitivepatternsasstate-happypeople
(e.g.,rewardsensitivity),traithappinessisprobablynotsimplyconstanthighstate
happiness.Rather,themostadaptivestrategymaybetheabilitytoflexiblychoose
subsystemstosample,aswellasre-construestimuliwithinthosesubsystems.This
strategywouldbemostlikelytoproducepositiveoutcomesacrossmanydifferent
situationswhileallowingforappropriaterecognitionofandresponsetonegative
situations.Sometimes,dealingwithnegativityinthe(p.109) moment(e.g.,preventing
somethingbadfromhappening)helpstoensurefuturewell-being.Themostadaptive
formoftraithappiness—lessreliantononesubsystemandlikelytobedisplayedacross
thewidestvarietyofsituations—arisesfromthisabilitytobecognitivelyflexiblewhen
appropriate.
Cognitiveflexibilityreferstotheabilitytochangethoughtsorinterpretationsselectively
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Positive Emotion and the Brain
inresponsetoenvironmentalstimuliwiththegoalofperceiving,processing,and
respondingtosituationsindifferentways(Scott,1962).Theabilitytoflexiblyinterpret
andreinterpretinformationinone’senvironmentmayarisenaturallyfromapositive
affectivestate(e.g.,Ashby,Isen,&Turken,1999;Murray,Sujan,Hirt,&Sujan,1990),
and,reciprocally,oneimportantoutcomeofflexibilitymaybeincreasedpositiveaffect.
Givenourunpredictableandever-changingworld,demandingorimposingtoomuch
structureisnotadaptive.Rather,optimalinteractionwithone’senvironmentrequiresthe
abilitytodetectandrespondappropriatelytochange,bringingone’sexpectationsinline
withthecurrentstateoftheworldandreducingdiscrepancies.Totheextentthatoneis
cognitivelyflexible,thiswillbeeasiertodo.Justasnegativeemotionssignalaproblemin
theenvironmenttobeaddressed,positiveemotionssignalanabsenceofproblem—asign
thatalliswell.Whereasone’sgoalismoreclear-cutwithnegativeemotions(e.g.,escape
fromordealdirectlywithaclearthreat),agreatervarietyofpotentialbehavioroptions
areavailableforpositiveemotions.Onemayfocusonmaintainingone’spositivestate,
enjoyingone’ssurroundings,and/orseekingoutnewopportunities.Dependingonthe
situation,someoptionsmaybemoreappropriatethanotherswithinthislargepotential
repertoire.Atothertimes,emotionregulationthroughreconstruingstimuliintoone
subsystemoveranother(e.g.,reinterpretingfailureatagoalasareturntohomeostasis)
maybethebeststrategy.Thus,maximizingone’spotentialmaybedependentonthe
abilitytobeflexible.
Toillustratetheimportanceofcognitiveflexibilityintraithappiness,considertheexample
ofinflexibilityinpeoplesufferingfromdepressionwhoexperienceanhedonia—theinability
toexperiencepleasurefromrewardingstimuli.Theseindividualsmaynotfinditusefulto
relyonperceptual,consummatorypleasuresasasourceofhappiness.Rather,iftheyare
toexperiencehappiness,itmaybemorelikelytocomefrompreventionfocus,oriented
towardmaintenanceofabaselinestateandavoidanceofdeviationsfromthatstate.Inthe
contextofregulatoryfocus,positiveoutcomeslikelyarisefromtheabilitytodetectthe
appropriatenessofaparticularmotivationalfocusgiventheneedsofthespecificcontext,
ortoswitchmotivationalorientationswhenchangesoccurinthecontext,andthereforeto
alsomaintainvigilanceinapreventionfocusindangerousormaladaptivesituations.Tothe
extentthatpeopledrawonthe“wrong”subsystem(forthemandtheirparticular
orientation),theycanremainunhappy.Thus,individualswhosufferfrompsychological
dysfunctionmaynotbeusingtheappropriatestrategiestomaximizehappiness,including
knowingwhentoswitchtoadifferentsubsystem.
Whileincreasedpositiveaffectcanbeanoutcomeofcognitiveflexibility,anextreme
degreeofflexibilityinwhichallinformation,evenobjectivelynegativeevents,are
reconstruedaspositivedoesnotnecessarilyindicatehappinessorwell-being.For
example,peoplewithbipolardisordertendtoviewallstimuli(i.e.,positive,neutral,and
evennegative)asindiscriminatelypositive(Gruber,Johnson,Oveis,&Keltner,2008),
whichcanleadtoinappropriateandevendestructiveoutcomes.Thekeytohappinessis
probablybeingstrategicallyflexible,suchas“seeingthebrightside”ofambiguous
situations,withintheconstraintsofwhatiscontextuallyappropriate,includingsensitivity
tonegativeinformationwhenitisrelevant.Inotherwords,happypeoplehave
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Positive Emotion and the Brain
motivationalbiases,buttheyarenotblinders.Cognitiveflexibilitycantherefore(p.110)
bethoughtofasacontinuumwithanoptimallevelforadaptivefunctioning.Consistent
withthisframework,somerecentworksuggeststhathappinessisassociatedwith
enhancedamygdalaactivationtopositivestimuliwithoutreducedamygdalaactivationto
negativestimuli(Cunningham&Kirkland,2013).Thismaybebecausehappierpeople
displaygreateramygdalaflexibility—respondingtonegativeinformationwhenvigilanceis
necessary,andrespondingtopositiveinformationwhentherearenonegativestoattend
andopportunitiesabound.
Conclusion
Understandingtheneurobiologicalmechanismsunderlyingpositiveemotionand
happinesscanprovideinsightintohowpeopleinterpretandnavigatetheircomplexsocial
worlds.Inthischapter,wereviewedtheneuroscientificliteratureonpositiveemotion
andhappinessfromtwodistincttheoreticalperspectives,includingrewardsensitivityand
motivational/hedonicbehavior.Themajorareasinvolvedinthefirstperspective(reward
sensitivity)helptodetectstimuli,tagthemasrelevant,andguidebehaviortoward
rewardsand,conversely,awayfrompunishments.Projectionsfromventraltegmental
areatoamygdala,andfromamygdalatonucleusaccumbensandhippocampus,are
involvedindetectingrewardandarefacilitatedbythetransmissionofdopamine.
Informationfromamygdalaandnucleusaccumbensisfurtherintegratedinthe
orbitofrontalcortex,whichisresponsibleforguidingbehavior.Throughdensereciprocal
connectionsbetweenorbitofrontalcortexandtheamygdalaandnucleusaccumbens,
informationcanbesharedandinterpretedacrosslevelsofprocessing.
Bothmotivational(anticipatory)andhedonic(consummatory)behaviorsinvolvethe
nucleusaccumbensaswell.Thecoreofthenucleusaccumbensisorientedtoward
anticipatorystatesandfacilitatedbythereleaseofdopamine.Anticipatorystatesthat
involvegreatercognitiveelaboration,suchasprospection,alsoinvolvehigher-order
regionssuchastheprefrontalcortex.Bycontrast,theshellofthenucleusaccumbensis
orientedtowardconsummatorystatesandfacilitatedbythereleaseofendogenous
opioids.Interconnectionsbetweenthenucleusaccumbensshellandtheventralpallidum
servetoenhancetheseconsummatoryresponses.
Thisresearchhasclinicalimplicationsforimprovinghappiness.Giventheevidence
suggestingthathappinessisfacilitatedbycognitiveflexibility,cliniciansmaybeableto
improvesubjectivewell-beingintheirpatientsthroughinterventionsdesignedto
increasethisflexibility.Learningstrategiesforchangingthoughtsorinterpretationsin
responsetoenvironmentalstimulicouldbevaluabletodepressedpatientsandothers
whostrugglewithpsychologicaldysfunction.Cognitivebehavioraltherapy,whichfocuses
onchangingmaladaptivethoughtsandemotionstomoreadaptiveones(Hofmann,2011),
takesanapproachthatiscomplementarywiththisperspective.Itwouldbeinterestingto
findoutwhetherimprovingone’soverallabilitytobeflexible,ratherthanchanging
specificnegativethoughts,wouldhaveadditionalbenefits.
Anotheravenueforfutureresearchmightfocusonlinkinghappiness(es)tophysical
healththroughincreasedunderstandingofthebiologicalmechanisms.Forexample,ifone
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Positive Emotion and the Brain
routetohappinessinvolvesmaintenanceofhomeostasisthroughtheprocessofreducing
variabilityintheautonomicnervoussystem,thiscanexplainwhysomepeoplerespondso
negativelytoenvironmentalevents(e.g.,givingaspeech)asstressorsthatincrease
physiologicalarousal,whileothersrespondtothesameeventsbybeingenergizedand
performingbetter.Frequentexposure(p.111) totheseeventsmightresultinmore
harmfulhealthoutcomesforsomeindividualsthanothers.Agreaterunderstandingof
themechanismspredictingthesedifferentoutcomesmightleadtowaysinwhichphysical
healthcouldbeimproved,takingintoaccountanindividual’suniqueneedsandresponse
patterns.
Happinessisconstructedandreflectedthroughouttheneuroaxis,andthese
“happinesses”arelinkedtodifferentaspectsofpsychologicalfunctioning.Positiveaffect
canmanifestandbeinterpretedinseveralways,leadingtodifferentsubjectivesensesof
happiness.Whereassomepeoplederivehappinessfromsensoryexperiencessuchas
musicorfood,othersaremoreappreciativeofachievements,andstillothersare
happiestwhennothingisgoingwrong.Thus,althoughtheterm“happiness”isconstant
acrossindividuals,itmayreflectavarietyofsubjectiveexperiences.
Notes
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Notes:
(1).Thenucleusaccumbens(NAcc)isastructurelocatedintheventralstriatum,which
ispartofalargerareacalledthebasalganglia;wesometimesrefertothestriatumand
othertimestothebasalganglia,bothofwhichincludethenucleusaccumbens.
(2).Peoplefocusedonthepastmayalsobeinaconsummatorystateiftheyare
reliving/savoringpastexperiences.
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