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Behavioral Neuroscience 9th Edition S. Mark Breedlove
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GrossElectricalActivityoftheBrainIsReadilyDetected
TheCuttingEdge■Optogenetics:UsingLighttoProbeBrain-BehaviorRelationships
VisualSummary
4 TheChemistryofBehavior NeurotransmittersandNeuropharmacology
TheBirthofaPharmaceuticalProblemChild
SynapticTransmissionInvolvesaComplexElectrochemicalProcess ManyNeurotransmittersHaveBeenIdentified NeurotransmitterSystemsFormaComplexArrayintheBrain
BOX4.1 PathwaysforNeurotransmitterSynthesis
TheEffectsofaDrugDependonItsSiteofActionandDose DrugsAffectEachStageofNeuralConductionandSynapticTransmission SomeNeuroactiveDrugsEasetheSymptomsofInjuryorPsychiatricIllness SomeNeuroactiveDrugsAreUsedtoAlterConsciousExperiences SubstanceAbuseandAddictionAreWorldwideSocialProblems
TheCuttingEdge■UncoveringtheInsula
VisualSummary
5 HormonesandtheBrain
CraftingaPersonalitythroughHormones
HormonesHaveManyActionsintheBody HormonesHaveaVarietyofCellularActions
BOX51 TechniquesofModernBehavioralEndocrinology EachEndocrineGlandSecretesSpecificHormones
TheCuttingEdge■BonesSecreteHormonestoRegulateAppetite HormonesRegulateSocialBehaviorsandViceVersa VisualSummary EvolutionandDevelopmentofthe NervousSystem
6 EvolutionoftheBrainandBehavior
WeAreNotSoDifferent,AreWe?
HowDidtheEnormousVarietyofSpeciesAriseonEarth? WhyShouldWeStudyOtherSpecies?
BOX61 WhyShouldWeStudyParticularSpecies?
BOX6.2 ToEachItsOwnSensoryWorld
AllVertebrateBrainsSharetheSameBasicStructures
TheEvolutionofVertebrateBrainsReflectsChangesinBehavior ManyFactorsLedtotheRapidEvolutionofaLargeCortexinPrimates
BOX6.3 EvolutionaryPsychology
EvolutionContinuesToday
TheCuttingEdge■AreHumansStillEvolving?
VisualSummary
7 Life-SpanDevelopmentoftheBrainandBehavior
OvercomingBlindness
NeuralDevelopmentIsGuidedbytheInteractionofGeneticandEnvironmental Factors
DevelopmentoftheNervousSystemCanBeDividedintoSixDistinctStages
BOX71 TransgenicandKnockoutMice
BOX7.2 DegenerationandRegenerationofNervousTissue
LifelongSynapseRearrangementIsGuidedbyExperience
ExperienceCanAlterGeneExpressiontoAffectBrainDevelopment
TheCuttingEdge■HarnessingGliatoReverseanInheritedBrainDisorder
TheBrainContinuestoChangeasWeGrowOlder
VisualSummary
PerceptionandAction 8
GeneralPrinciplesofSensoryProcessing,Touch,andPain
What’sHot?What’sNot?
SENSORYPROCESSING
SensoryReceptorOrgansDetectEnergyorSubstances
SensoryInformationProcessingIsSelectiveandAnalytical
BOX8.1 Synesthesia
TOUCH:MANYSENSATIONSBLENDEDTOGETHER
SkinIsaComplexOrganThatContainsaVarietyofSensoryReceptors
PAIN:ANUNPLEASANTBUTADAPTIVEEXPERIENCE
HumanPainCanBeMeasured
TheCuttingEdge■EvolvinganIndifferencetoToxins
PainCanBeDifficulttoControl
VisualSummary
9 Hearing,Balance,Taste,andSmell
NoEarforMusic
HEARING
PressureWavesintheAirArePerceivedasSound
BOX9.1 TheBasicsofSound
AuditorySignalsRunfromCochleatoCortex
PitchInformationIsEncodedinTwoComplementaryWays
BrainstemAuditorySystemsAreSpecializedforLocalizingSounds
TheAuditoryCortexProcessesComplexSounds
HearingLossIsaMajorDisorderoftheNervousSystem
VESTIBULARPERCEPTION AnInnerEarSystemSensesGravityandAcceleration
THECHEMICALSENSES:TASTEANDSMELL
ChemicalsinFoodsArePerceivedasTastes
TheCuttingEdge■MoreThanaMatterofTaste ChemicalsintheAirElicitOdorSensations
VisualSummary
10 Vision
FromEyetoBrain
WhenSeeingIsn’tSeeing
TheRetinaTransducesLightintoNeuronalActivity PropertiesoftheRetinaShapeManyAspectsofOurVision NeuralSignalsTravelfromtheRetinatoSeveralBrainRegions
BOX101 EyeswithLensesHaveEvolvedinSeveralPhyla NeuronsatDifferentLevelsoftheVisualSystemHaveVeryDifferentReceptive Fields
ColorVisionDependsonSpecialChannelsfromtheRetinalConesthrough CorticalAreaV4
BOX102 MostMammalianSpeciesHaveSomeColorVision
TheManyCorticalVisualAreasAreOrganizedintoTwoMajorStreams VisualNeuroscienceCanBeAppliedtoAlleviateSomeVisualDeficiencies
TheCuttingEdge■SeeingtheLight VisualSummary
11
MotorControlandPlasticity
WhatYouSeeIsWhatYouGet
TheBehavioralViewConsidersReflexesversusPlans NeuroscienceRevealsHierarchicalSystems
TheSpinalCordIsaCrucialLinkinControllingBodyMovement PathwaysfromtheBrainControlDifferentAspectsofMovements
BOX11.1 CorticalNeuronsCanGuideaRoboticArm ExtrapyramidalSystemsAlsoModulateMotorCommands
TheCuttingEdge■CerebellarGliaPlayaRoleinFineMotorCoordination BrainDisordersCanDisruptMovement
BOX11.2 Prion-LikeNeurodegenerationMayBeatWorkinParkinson’s VisualSummary
Sex
SEXUALBEHAVIOR
ReproductiveBehaviorCanBeDividedintoFourStages
TheNeuralCircuitryoftheBrainRegulatesReproductiveBehavior
TheCuttingEdge■SexualExperienceSolidifiesNeuralCircuitsforMating
TheHallmarkofHumanSexualBehaviorIsDiversity ForManyVertebrates,ParentalCareDeterminesOffspringSurvival
SEXUALDIFFERENTIATION
SexDeterminationandSexualDifferentiationOccurEarlyinDevelopment GonadalHormonesDirectSexualDifferentiationoftheBrainandBehavior
BOX121 TheParadoxicalSexualDifferentiationoftheSpottedHyena DoFetalHormonesMasculinizeHumanBehaviorsinAdulthood?
VisualSummary
13 Homeostasis
ActiveRegulationoftheInternalEnvironment
HarshRealityTV
HomeostasisMaintainsaConsistentInternalEnvironment:TheExampleof Thermoregulation
BOX13.1 PhysiologicalandBehavioralThermoregulationAreIntegrated FLUIDREGULATION
WaterShuttlesbetweenTwoBodyCompartments TwoInternalCuesTriggerThirst FOODANDENERGYREGULATION
NutrientRegulationHelpsPrepareforFutureNeeds AHypothalamicAppetiteControllerIntegratesMultipleHungerSignals
TheCuttingEdge■FriendswithBenefits
ObesityandEatingDisordersAreDifficulttoTreat
BOX13.2 BodyFatStoresAreTightlyRegulated,EvenafterSurgicalRemovalofFat VisualSummary
14 BiologicalRhythms,Sleep,andDreaming
WhenSleepGetsOutofControl
BIOLOGICALRHYTHMS
ManyAnimalsShowDailyRhythmsinActivity
TheHypothalamusHousesaCircadianClock SLEEPINGANDWAKING
HumanSleepExhibitsDifferentStages WhyandHowDidSleepEvolve?
BOX14.1 SleepDeprivationCanBeFatal AtLeastFourInteractingNeuralSystemsUnderlieSleep
TheCuttingEdge■CanIndividualNeuronsBe“Sleepy”?
SleepDisordersCanBeSerious,EvenLife-Threatening
VisualSummary
TheHazardsofFearlessness
BroadTheoriesofEmotionEmphasizeBodilyResponses
BOX15.1 LieDetector?
DidaCoreSetofEmotionsEvolveinHumansandOtherAnimals? SpecializedNeuralMechanismsMediatetheExperienceandExpressionof Emotions
TheCuttingEdge■SynapticChangesduringFearConditioning NeuralCircuitry,Hormones,andSynapticTransmittersMediateViolenceand Aggression
StressActivatesManyBodilyResponses
VisualSummary
16
Psychopathology
BiologicalBasisofBehavioralDisorders “TheVoice”
SchizophreniaIstheMajorNeurobiologicalChallengeinPsychiatry
BOX16.1 Long-TermEffectsofAntipsychoticDrugs
DepressionIstheMostPrevalentMoodDisorder
TheCuttingEdge■CanOurGenesTellUsWhichDrugstoUse?
BOX16.2 TheSeasontoBeDepressed?
ExtremeMoodCyclesDefineBipolarDisorder
ThereAreSeveralTypesofAnxietyDisorders
BOX16.3 Tics,Twitches,andSnorts:TheUnusualCharacterofTourette'sSyndrome
VisualSummary
PARTVI CognitiveNeuroscience 17 LearningandMemory
TrappedintheEternalNow
FUNCTIONALPERSPECTIVESONLEARNINGANDMEMORY
ThereAreSeveralKindsofLearningandMemory DifferentFormsofNondeclarativeMemoryInvolveDifferentBrainRegions SuccessiveProcessesCapture,Store,andRetrieveInformationintheBrain
BOX17.1 EmotionsandMemory
NEURALMECHANISMSOFMEMORYSTORAGE
MemoryStorageRequiresPhysicalChangesintheBrain SynapticPlasticityCanBeMeasuredinSimpleHippocampalCircuits
TheCuttingEdge■ArtificialActivationofanEngram IntheAdultBrain,NewlyBornNeuronsMayAidLearning
VisualSummary
OneThingataTime ATTENTION
AttentionSelectsStimuliforProcessing
BOX18.1 ReactionTimeResponses,fromInputtoOutput
TargetsofAttention:AttentionAlterstheFunctioningofManyBrainRegions
SourcesofAttention:ANetworkofBrainSitesCreatesandDirectsAttention
CONSCIOUSNESSANDEXECUTIVEFUNCTION
ConsciousnessIsaMysteriousProductoftheBrain
BOX18.2 PhineasGage
TheCuttingEdge■BuildingaBetterMindReader
VisualSummary
LanguageandLateralization
SilencingtheInnerVoice
BRAINASYMMETRYANDLATERALIZATIONOFFUNCTION
TheLeftandRightHemispheresAreDifferent
BOX19.1 TheWadaTest
Right-HemisphereDamageImpairsSpecificTypesofCognition Left-HemisphereDamageCanCauseAphasia
CompetingModelsDescribetheLeft-HemisphereLanguageSystem VERBALBEHAVIOR:SPEECHANDREADING
LanguageHasBothLearnedandUnlearnedComponents
BOX19.2 WilliamsSyndromeOffersCluesaboutLanguage
BOX19.3 VocalBehaviorinBirdsandOtherSpecies
ReadingSkillsAreDifficulttoAcquireandFrequentlyImpaired RECOVERYOFFUNCTIONAFTERBRAINDAMAGE
19.7
StabilizationandReorganizationAreCrucialforRecoveryofFunction
BOX19.4 TheAmazingResilienceofaChild’sBrain
TheCuttingEdge■ContactSportsCanBeCostly VisualSummary
Appendix
Glossary
References
AuthorIndex
SubjectIndex
Preface Twenty-fouryearsago,anewkindoftextbookwaspublishedforUniversitycoursesthatwereoftencalled“Brain andBehavior”Asthefieldevolved,thebook’stitlemetamorphosedfromBiologicalPsychology to Behavioral Neuroscience,butthesamedrivetoprovideadefinitiveandcomprehensivesurveyoftheneurosciencesliesatthe heartofallourefforts.Westrivetokeepthebookup-to-datewhilekeepingaconversationaltonetomakethis wealthofinformationnotjustaccessible,butfascinating Thebiggestchangeinthisneweditionisthedevelopmentof LearningObjectives foreachsegmentofthebook,withtheideathattelegraphingwhat’stocomewillfocus readers’attentionandfacilitatelearning Asyoufinisheachsectionoftext,itwouldbeagoodideatogobackand readtheassociatedLearningObjectivestoseewhetherinfactyouincorporatedthematerial.Ifnot,aquickreview ofthattextmaybeinorder
Asalways,therehavebeenplentyofnewfindingstoaddtothisedition Infact,theproblemwefaceiswhichof themany,manynewfindingstoleaveout thosethatarenotquiteessentialforasurveyofthefield Wearepretty pickyaboutwhatweadd,andstillitseemslikeadelugeofnewinformationandideas.Hundredsofnewpapersare citedinthisedition Ifthatsoundslikealot,letusgiveyouaperspectiveonhowmanynewpaperswereomitted Onournewsfeedsite(wwwbiopsychologycom/news/),1,299newlinkswereaddedin2018alone Thosearejust thefindingsthatwereimportantenoughtogettheattentionofmassmediareporters AswenoteinChapter1,over 40,000newarticlesindexedunder“neuroscience”appearedthatyearinPubMed Itwouldtakeathicktomejustto listthetitlesofthepapersfrom2018!
Whilebeingvery,veryselectiveinsamplingthisfloodoffindings,wehavemadesubstantialchangesinevery chapter.Forexample,inChapter3wehaveanewfigurecomparing“kissandrun”synapseswithmoretraditional modelsofsynaptictransmission Chapter5hasnewmaterialaboutahormonesecretedfrombonethatactsonthe hypothalamustoreduceappetite WetotallyreorganizedChapter7foramorestreamlinedapproachanddiscussthe growingdoubtsaboutwhetheramyloiddepositscauseAlzheimer’s Chapter9 needed a newfigure comparing transductioninthefivetastereceptors.Chapter13talksaboutyetanotherfactoraffectingappetite,glucagon-like peptide1 Chapter16nowdiscussesthelogicofpharmacogenomicstotreatdepression,whileChapter18was thoroughlyreorganizedandincludesmoreaboutexecutivefunction Honestly,wecouldgoonlikethisforevery chapter Clearlythisisanexcitingeraintheneurosciences AsLewisCarrolputit,“Wemustrunasfastaswecan justtostayinplace!”
We’ve also kept severalverypopular features fromprevious editions:The Cutting Edge appears ineach chapter,whereweexploresomeofthemostexcitingexamplesofrecentresearch,andeachchapterendswitha VisualSummary,whereyoucanseegraphicremindersasyoureviewtheprinciplefindingsthatwejustpresented TheseVisualSummariesreallyshineonline,wherewithjustaclickyoucanreviewfigures,animations,andquizzesto helpintegratethematerial Wealsocontinuetoopeneachchapterwithagrippingvignette,relatingsomeone’sreallifeexperiencesthatwillbebetterunderstoodasthecontentofthechapterunfolds,andweagainreplacedseveralof thesevignettesasmorerecenteventsbringtothesurfacemanyoftheimportantissuesinbehavioralneuroscience Likewisewe’veretainedthemarginalglossarythatmakesiteasytofindthedefinitionsthatunlockthematerial,as wellastwofeaturestoletyouburrowinonaparticularsubject:theonlinesupplementscalledAStepFurthercited throughoutthetext,andtheRecommendedReadingatthecloseofeachchapter
Youmightthinkthatapproachingthequarter-centurymarkwe’dbejadedaboutimprovingandrevisingour presentations,butwestillloveit,perhapsbecausethedynamicandexcitingpaceofneuroscienceresearchshowsno signofabatingsoon Asalways,wewelcomeallfeedback,praiseorcriticism,cutsoradditions,fromourreaders Youcanemailusdirectlyatbehavneuro@gmailcom
Acknowledgments We continue to feelso luckyto work withthe inestimable teamatSinauerAssociates, nowa partofOxford UniversityPress,whosedeep skillsand generousguidancetransformourhundredsoffiles,thousandsofemail attachmentsandsometimesscrambledemailsintoyetanotherbeautifulbook.Again,wefeelsogratefultobenefit fromtheexperienceandexquisitetasteofothers Inparticular,thebookcouldnotexistwithoutthecontributionsof SeniorAcquisitionEditorSyd Carroll, ProductionEditorAlisonHornbeck, ProductionManagerJoanGemme, BookDesignerandProductionSpecialistAnnetteRapier,andMediaandSupplementsEditorZanCarterandher crew.WealsofondlybidadieutotherecentlyretiredChrisSmall,ProductionManagerforallourpreviouseditions. Wehopeyou’reenjoyingyourself,Chris,buthowcouldyouabandonus!?Acadreofcommandosdelveddeepin thearchivestodealwithcopyrightsandpermissions,sowesaluteyouMicheleBeckta,MarkSiddall,andTracy Marton We’dalsoliketothankourcopyeditorLouDoucette,andourlongtimeartstudio,DragonflyMedia,who bringamazingskillandcommitmenttomakeuslookgood.
WemustalsothankthefounderofSinauerAssociates,AndySinauer,forhisunwaveringsupportovertheyears, withatouchofsadnessuponhisretirement WearesoproudtobeapartofAndy’stremendouslegacy,begunall thoseyearsagowithFrom Neuron to Brain,creatinggorgeousbooksthatmakeeventhemostcomplextopics accessibleandenjoyable.
Bythis pointinthe evolutionofthe book, we have benefited fromthe wisdomand advice ofhundreds of colleagueswhohavegenerouslyservedasreviewersofpasteditions Althoughwedon’thavethespacetolistthem all,wewanttoacknowledgethatinmanywaysthebookyouareholdingistheproductofawholecommunityof neuroscientists Inthis,theNinthEdition,thefollowingcolleagueshaveprovidedinvaluablecritiqueandcommentary:
SusanBachus,UniversityofMaryland,BaltimoreCounty
SusanBarron,UniversityofKentucky
ChristopherBeeman,CentralWashingtonUniversity
JinBo,EasternMichiganUniversity
DavidBrodbeck,AlgomaUniversity
ElizabethCaldwell,UniversityofNewHampshire
JamesCherry,BostonUniversity
MichaelCohen,LoyolaUniversityChicago
PaulJ Currie,ReedCollege
PatrickCushen,MurrayStateUniversity
DeanaDavalos,ColoradoStateUniversity
DarraghPDevine,UniversityofFlorida
ChristopherWDrapeau,ValparaisoUniversity
KelliA Duncan,VassarCollege
RaymondH.Dye,Jr.,LoyolaUniversityChicago
TaffetaElliott,NewMexicoInstituteofMiningandTechnology
AlisonA Fedio,ArgosyUniversity,NorthernVirginia
SaraB Festini,UniversityofTampa
CynthiaMichelleFinley,CollegeofMarin
JonathanFranz,SUNYEmpireStateCollege
KorenGanas,UniversityofIllinois
SophieGeorge,DixieStateUniversity
AaronGodlaski,CentreCollege
BrianJ Hock,AustinPeayStateUniversity
JenniferIngemi,NortheasternUniversity
MaryEllenKelly,HaverfordCollege
SusanKennedy,DenisonUniversity
MichaelKerchner,WashingtonCollege
SaritaLagalwar,SkidmoreCollege
StephenLippi,GeorgeMasonUniversity
MarioL.Mata,CaliforniaStateUniversity,LosAngeles
AlexandraRoach,UniversityofSouthCarolina,Aiken
RussellRomeo,BarnardCollegeofColumbiaUniversity
TimothyRoth,FranklinandMarshallCollege
EmmaSarro,DominicanCollege
PeterA Serrano,HunterCollege,CityUniversityofNewYork
FredricShaffer,TrumanStateUniversity
KatieAnnSkogsberg,CentreCollege
LucyJ.Troup,UniversityoftheWestofScotland
AdrianaUruena-Agnes,St PetersburgCollege
JenniferWilhelm,CollegeofCharleston
JanR Wessel,UniversityofIowa
SusanZup,UniversityofMassachusettsBoston
Finally,wethankallthosetirelesscolleaguestryingtounderstandtheneuralbasisofbehavior,withtechniquesthat wouldhaveseemedlikesorceryonlyafewyearsago,andwhosharetheirhard-wonfindingswithusall.
S MarcBreedlove NeilVWatson
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TheBehavioralNeuroscienceCompanionWebsitecontainsarangeofstudyandreviewresourcestohelpstudents masterthematerialpresentedineachchapterofthetextbook Accesstothesiteisincludedwitheachnewcopyof thetextbook(seeinsidefrontcover) Thesiteincludesthefollowingresources:
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Introduction ScopeandOutlook MachineorHuman? InthenearfuturedepictedintheHBOseriesWestworld,peoplevisitathemeparksetintheOldWest,withsteam locomotives,saloons,andbrothels,populatedwithandroids,called“hosts,”toentertainhumans Themechanicalhosts providetheirguestswithanything,fromcasualbantertogunfights,harmlessflirtingtokinkysex,theonlyrestriction beingthattherobotsarenevertoharmthehumans Theandroidhostsaresolifelikeinappearanceandbehaviorthat visitorsmayhaveahardtimedistinguishingwhethersomeoneisafellowguestorarobot Tomaketheandroids’ simulationofhumanscomplete,theyaregivenbackstories,falsememoriesofalifebeforetheirappearanceforeach newbatchofguests.Importantly,noneoftheandroidsknowthattheyaremechanicalbeingsratherthanhumans.It’s probablynotmuchofaspoilertosaythatseveralplotlinesintheserieshingeonandroidsslowlydiscoveringtheirtrue nature,movingfromshockandshamethattheyaremeremachines,toopenlyrebellingfromthenotionthattheyareto beused,andabused,asmereplaythingsforthehumans
Wearen’ttoldtoomuchabouthowtheandroid“brains”inWestworld work,because,ofcourse,suchtechnology remainsfaroutsideourgrasp,sothewriters,reducedtomerespeculation,remainrathervague Butapparentlythe knowledgeandpersonalityforanyparticularandroidliesina“controlunit,”agolf-ball-sizedevicethatcanbeextracted fromtheheadofonehostandimplantedintotheheadofanother,interchangeablebody Presumably,ifwehadenough knowledgeandsurgicalskill,wecouldremoveyourbrainfromyourheadandconnectituptotheheadofsomeother body Wouldyoustillbeyou?Evenifweputyourbrainintoabodyoftheoppositesex?Cometothinkofit,areyou entirelysurethereisabraininyourhead,andnotoneofthosecontrolunits?
Ouraiminthisbookistohelpyoulearnwhatisknownsofarabouthowbrainswork,andabouthowmuchmore wehaveyettolearn Wewillexplorethemanywaysinwhichthestructuresandactionsofthebrainproducemind andbehavior.Butthatisonlyhalfofourtask.Wearealsointerestedinthewaysinwhichbehaviorandexperience modifythe structures and actions ofthe brain One ofthe mostimportantlessons we wantto conveyis that interactionsbetweenbrainand behaviorare reciprocal Thebraincontrolsbehaviorand, inturn, behaviorand experiencealterthebrain
Wehopetogiveaninterestingaccountofthemainideasandresearchinbehavioralneuroscience,whichisof greatpopularaswellasscientificinterest Mostimportant,wetrytocommunicateourowninterestandexcitement aboutthemysteriesofmindandbody
BrainExplorer
1.1.3
TheBrainIsFullofSurprises LearningObjectives Afterreadingthissection,youshouldbeableto: Namethemaintypeofcellsfoundinthebrain,andnametheconnectionsbetweenthem Listthenamesofsomeofthemanyfieldsofstudyrelatedtobehavioralneuroscience Describefivedifferentperspectivestakeninunderstandingthebiologyofbehavior.
I usedtothink thatthebrainwasthemostwonderfulorganinmybody ThenI realizedwhowastellingmethis EmoPhilips (Americancomedian)
Ofcourseweshouldalwaysconsiderthesourcewhenevaluatinganidea,butevenso,thebrainindeedseemslikea prettywonderfulorgan For one thing, brains produced the entire extent ofhumanknowledge, everythingwe understandabouttheuniverse,howeverlimitedthatmaybe Brainsalsoproducedeverywrittendescriptionofthat hard-wonknowledge(includingthisbookyouholdinyourhands),aswellaseveryworkofvisualart,fromdoodles tothesweepingfrescosontheceilingoftheSistineChapel
Mostofushaveahardtimegraspingtheideaofabillionofanything,butyourheadcontainsanestimated86 billionnervecells,orneurons(fromtheGreekwordfor“nerve”or“cord”)(Herculano-Houzel,2012) Eachneuron contactsmanyothercellsatpointscalledsynapses,sotherearetrillionsofthosebetweenyourears.Aspecialized extensionofneurons,calledan axon,ismicroscopicallyslender,yetitmaybeseveralfeetlong We’lllearnthat axonsproduceelectricalimpulsesthattravelhundredsofmilesperhour Figure11offersalistofjustafewofthe thingswewilllearnaboutthehumanbraininthecourseofthisbook Allthishardwareisn’tjustforshow itallows youtotakeinalltheinformationinthatfigureinlessthanaminute.
1.1 YourBrainbytheNumbers Thecerebralcortexistheoutermostportionofthebrain
Whatisbehavioralneuroscience? Notreatyortradeunionagreementdefinestheboundariesofbehavioralneuroscience Thefirstpeopletostudythe relationshipsbetweenbrainandbehaviorregardedthemselvesasphilosophers,andtheirfindingscontributedtothe birthsofbiologyandpsychologyThosedisciplinesmergedinthetwentiethcenturytoformbiologicalpsychology, thefieldthatrelatesbehaviortobodilyprocesses.Withthemodernexplosionofneuroscience,thestudyofthe brain,thisresearchhasevolvedtothepointthatbehavioralneuroscience offersamoreaccuratedescription Whichevernameisused,themaingoalofthisfield isto understand theneuroscienceunderlyingbehaviorand experience
Behavioralneuroscience is a field that includes many players who come fromquite different backgrounds: psychologists,biologists,physiologists,engineers,neurologists,psychiatrists,andmanyothers Thus,therearemany careeropportunities,inbothuniversitiesandprivateindustry,forpeoplewithinterestsinthisfield(Hitt,2007) Figure 12 maps the relations of behavioral neuroscience to these many other disciplines Clearly, the behavioral neuroscienceumbrellaopensverywide.
1.2 What’sinaName? Inthisgraphicalrepresentationoftherelationshipsamongbehavioralneuroscienceandother scientificdisciplines,fieldstowardthecenterofthemapareclosesttobehavioralneuroscienceintheirhistory,outlook, aims,and/ormethods
Fiveviewpointsexplorethebiologyofbehavior Inourefforttounderstandtheneurosciencebasesofbehavior,weuseseveraldifferentperspectives.Becauseeach oneyieldsinformationthatcomplementstheothers,thecombinationofperspectivesisespeciallypowerful Wewill discussfivemajorperspectives:
1. 2. 3. 4. 5.
Describingbehavior
Observingthedevelopmentofbehavioranditsbiologicalcharacteristicsoverthelife-span
Studyingthebiologicalmechanismsofbehavior
Studyingapplicationsofbehavioralneuroscience forexample,itsapplicationtodysfunctionsofhumanbehavior
Studyingtheevolutionofbehavior
Theseperspectivesarediscussedinthesectionsthatfollow,andTable1.1illustrateshoweachperspectivecanbe appliedtothreekindsofbehavior
TABLE1.1
FiveResearchPerspectivesAppliedtoThreeKindsofBehavior
Researchperspective Sexualbehavior Learningandmemory Languageand communication
DESCRIPTION
Structural
Functional
Whatarethemainpatternsof reproductivebehaviorand sexdifferencesin behavior?
Howdospecializedpatterns ofbehaviorcontributeto matingandtocareof young?
ONTOGENY(development) Howdoreproductiveand secondarysex
Inwhatmainwaysdoes behaviorchangeasa consequenceofexperience forexample, conditioning?
Howdocertainbehaviors leadtorewardsor avoidanceofpunishment?
Howarethesoundsof speechpatterned?
Whatbehaviorisinvolvedin makingstatementsor askingquestions?
Howdolearningandmemory changeaswegrowolder? Whatchangesinthebrain whenachildlearnsto
MECHANISMS
APPLICATIONS
EVOLUTION
characteristicsdevelop overthelife-span? speak?
Whatneuralcircuitsand hormonesareinvolvedin reproductivebehavior?
Lowdosesoftestosterone restorelibidoinsome postmenopausalwomen
Howdoesmatingdependon hormonesindifferent species?
Whatanatomicaland chemicalchangesinthe brainholdmemories?
Genetherapyandbehavioral therapyimprovememoryin somesenilepatients
Whatbrainregionsare particularlyinvolvedin language?
Speechtherapy,in conjunctionwith amphetaminetreatment, speedslanguagerecovery followingstroke
Howdodifferentspecies compareinkindsandspeed oflearning?
Howdidthehumanspeech apparatusevolve?
Behaviorcanbedescribedaccordingtodifferentcriteria Untilwedescribewhatwewanttostudy,wecannotaccomplishmuch Dependingonourgoals,wemaydescribe behaviorintermsofdetailedactsorprocesses,orintermsofresultsorfunctions Ananalyticaldescriptionofarm movementsmightrecordthesuccessivepositionsofthelimborthecontractionofdifferentmuscles Afunctional behavioraldescription,bycontrast,wouldstatewhetherthelimbwasbeingusedinwalkingorrunning,textingor sexting To be usefulforscientific study, a descriptionmustbe precise and revealthe essentialfeaturesofthe behavior,usingaccuratelydefinedtermsandunits
Thebodyandbehaviordevelopoverthelife-span Ontogenyistheprocessbywhichanindividualchangesinthecourseofitslifetime thatis,growsupandgrows old Observingthewayinwhichaparticularbehaviorchangesduringontogenymaygiveuscluestoitsfunctionsand mechanisms.Forexample,weknowthatlearningabilityinmonkeysincreasesoverthefirstyearsoflife.Therefore, wecanspeculatethatprolongedmaturationofbraincircuitsisrequiredforcomplexlearningtasks Inrodents,the abilitytoformlong-termmemorieslagssomewhatbehindthematurationoflearningability So,youngrodentslearn wellbutforgetmore quicklythanolderones, suggestingthatlearningand memoryinvolve differentprocesses Studyingthedevelopmentofreproductivecapacityandofdifferencesinbehaviorbetweenthesexes,alongwith changesinbodystructuresandprocesses,throwslightonbodymechanismsunderlyingsexualbehaviors
Biologicalmechanismsunderlieallbehavior To learnaboutthe mechanismsofanindividual’sbehavior, we studyhowhisorher present bodyworks To understand the underlying mechanisms ofbehavior, we must regard the organism(with alldue respect) as a “machine,”madeupofbillionsofneurons.Wemustask,Howisthisthingconstructedtobeabletodoallthat? Thesearesometimesdescribedas proximate questions questionsaboutthephysicalinteractionsthatcontrola particular behavior How cells inyour eye respond differentlyto light ofdifferent wavelengths is a proximate question Ontheotherhand,whycolorvision,onceitarose,benefitedourancestorsisanevolutionaryquestion
Our major aimin behavioralneuroscience is to examine body mechanisms that make particular behaviors possible Inthecaseoflearningandmemory,forexample,wewouldliketoknowthesequenceofelectricaland biochemicalprocessesthatoccurwhenwelearnsomethingandretrieveitfrommemoryWhatpartsofthenervous systemareinvolvedinthatprocess?Inthecaseofreproductivebehavior,wealsowanttounderstandtheneuronal andhormonalprocessesthatunderliematingbehaviors.
Researchcanbetranslatedtoaddresshumanproblems
Likeothersciences,behavioralneuroscienceisalsodedicatedtoimprovingthehumancondition Numeroushuman diseasesinvolvemalfunctioningofthebrain Manyofthesearealreadybeingalleviatedasaresultofresearchinthe neurosciences,andtheprospectsforcontinuingadvancesaregood Attemptstoapplyknowledgealsobenefitbasic research.Forexample,thestudyofmemorydisordersinhumanshaspushedinvestigatorstoextendourknowledge ofthebrainregionsinvolvedindifferentkindsofmemory(seeChapter17)
Wecomparespeciestolearnhowthebrainandbehaviorhaveevolved Natureisconservative Onceparticularfeaturesofthebodyorbehaviorevolve,theymaybemaintainedformillions ofyearsandmaybeseeninanimalsthatotherwiseappearverydifferent Forexample,theelectricalmessagesused bynervecells(seeChapter3)areessentiallythesameinajellyfish,acockroach,andahumanbeing Someofthe chemicalcompounds thattransmitmessages throughthe bloodstream(hormones) are also the same indiverse animals(seeChapter5) Speciessharetheseconservedcharacteristicsbecausethefeaturesfirstaroseinashared ancestor(Box11onthenextpage) Butmeresimilarityofafeaturebetweenspeciesdoesnotguaranteethatthe featurecamefromacommonancestralspecies.Similarsolutionstoaproblemmayhaveevolvedindependentlyin differentclassesofanimals
Howdosimilaritiesanddifferencesamongpeopleandanimalsfitintobehavioralneuroscience?Eachpersonisinsome wayslikeallotherpeople,insomewayslikesomeotherpeople,andinsomewayslikenootherperson Asthefigure shows,wecanextendthisobservationtothemuchbroaderrangeofanimallife Eachpersonisinsomewayslikeall otheranimals(eg,needingtoingestcomplexorganicnutrients),insomewayslikeallothervertebrates(eg,havinga spinalcolumn),insomewayslikeallothermammals(e.g.,nursingouryoung),andinsomewayslikeallother primates(eg,havingahandwithanopposablethumbandarelativelylarge,complexbrain)
Whetherknowledgegainedaboutaprocessinanotherspeciesappliestohumansdependsonwhetherwearelike thatspeciesinregardtothatprocess Thefundamentalresearchonthemechanismsofinheritanceinthebacterium Escherichiacoliprovedsowidelyapplicablethatsomemolecularbiologistsproclaimed,“Whatistrueof E coliistrue oftheelephant”Toaremarkableextent,thatstatementistrue,buttherearealsosomeimportantdifferencesinthe geneticmechanismsof E coliandmammals
Withrespecttoeachbiologicalproperty,researchersmustdeterminehowanimalsareidenticalandhowtheyare different.Whenweseekanimalmodelsforstudyinghumanbehaviororbiologicalprocesses,wemustaskthe followingquestion:Doestheproposedanimalmodelreallyhavesomethingsincommonwiththeprocessatworkin humans(Seoketal,2013)?Wewillseemanycasesinwhichitdoes
Evenwithinthesamespecies,however,individualsdifferfromoneanother:catfromcat,bluejayfrombluejay,and personfromperson Behavioralneuroscienceseekstounderstandindividualdifferencesaswellassimilarities Therefore,thewayinwhicheachpersonisabletoprocessinformationandstorethememoriesoftheseexperiencesis anotherpartofourstory
Charles Darwin’s theory of evolution through naturalselection is centralto allmodern biology Fromthis perspectiveemergetworatherdifferentemphases:(1)the continuity ofbehaviorandbiologicalprocessesamong speciesthatreflectssharedancestryand(2)thespecies-specific differences inbehaviorand biologythathave evolvedasadaptationstodifferentenvironments
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1.2.4
1.2.5
ThreeApproachesRelateBrainandBehavior LearningObjectives Afterreadingthissection,youshouldbeableto:
Differentiatebetweentheindependentanddependentvariablesinscientificexperiments
Namethetypeofresearchinwhichapartofthebrainismanipulatedtoobserveeffectsonbehavior,and offerexamples.
Namethetypeofresearchinwhichbehaviororexperienceismanipulatedtoobserveeffectsonthebrain, andofferexamples.
Describecorrelationalresearchaboutthebrainandbehavior,andofferexamples.
Explainwhythebrainmustbecapableofchangingitsstructure,andnamethetermtodescribethat changeability.
Behavioralneuroscientistsusethreeapproachestounderstandtherelationshipbetweenbrainandbehavior:somatic intervention,behavioralintervention,andcorrelation.Inthemostcommonapproach,somaticintervention(Figure 13A),wealterastructureorfunctionofthebrainorbodytoseehowthisalterationchangesbehavior Here, somaticinterventionistheindependentvariable,andthebehavioraleffectisthedependentvariable;thatis,the resultingbehaviordependsonhowthebrainhasbeenaltered Forexample,inresponsetomildelectricalstimulation ofonepartofherbrain,notonlydidonepatientlaugh,butshefoundwhatevershehappenedtobelookingat amusing(Friedetal,1998)
13 ThreeMainApproachestoStudyingtheNeuroscienceofBehavior (A)Insomaticintervention,investigators changethebodystructureorchemistryofananimalinsomewayandobserveandmeasureanyresultingbehavioral effects.(B)Conversely,inbehavioralintervention,researcherschangeananimal’sbehaviororitsenvironmentandtryto ascertainwhetherthechangeresultsinphysiologicaloranatomicalchanges (C)Measurementsofbothkindsofvariables allowresearcherstoarriveatcorrelationsbetweensomaticchangesandbehavioralchanges (D)Eachapproachenriches andinformstheothers
Inlaterchapterswedescribemanykindsofsomaticinterventionwithbothhumansandotheranimals,asinthe followingexamples:
Ahormoneisadministeredtosomeanimalsbutnottoothers;variousbehaviorsofthetwogroupsarelater compared
Apartofthebrainisstimulatedelectrically,orbyuseoflighttostimulateonlyaparticularclassofneurons,and behavioraleffectsareobserved.
Aconnectionbetweentwopartsofthenervoussystemiscut,andchangesinbehavioraremeasured
Theapproachoppositetosomaticinterventionispsychologicalorbehavioralintervention(Figure13B) Inthis approach,thescientistintervenesinthebehaviororexperienceofanorganismandlooksforresultingchangesin bodystructureorfunction Here,behavioristheindependentvariable,andchangeinthebodyisthedependent variable Amongtheexamplesthatwewillconsiderinlaterchaptersarethefollowing: Puttingtwoadultsofoppositesextogethermayleadtoincreasedsecretionofcertainhormones.
Exposingapersonoranimaltoavisualstimulusprovokeschangesinelectricalactivityandbloodflowinpartsof thebrain
Trainingofanimalsinamazeisaccompaniedbyelectrical,biochemical,andanatomicalchangesinpartsoftheir brains.
Thethirdapproachtobrain-behaviorrelations,correlation(Figure13C),consistsoffindingtheextenttowhich
agivenbodymeasurevarieswithagivenbehavioralmeasure Laterwewillexaminethefollowingquestions,among others:
Arepeoplewithlargebrainsmoreintelligentthanpeoplewithsmallerbrains(atopicwe’lltakeuplaterinthis chapter)?
Areindividualdifferencesinsexualbehaviorcorrelatedwithlevelsofcertainhormonesintheindividuals? Istheseverityofschizophreniacorrelatedwiththemagnitudeofchangesinbrainstructure?
Suchcorrelationsshouldnotbetakenasproofofcausalrelationship Foronething,evenifacausalrelation exists,thecorrelationdoesnotrevealitsdirection thatis,whichvariableisindependentandwhichisdependent. Foranother,twofactorsmightbecorrelatedonlybecauseathird,unknownfactoraffectsthetwofactorsmeasured Ifyouandyourstudypartnergetsimilarscoresonanexam,that’snotbecauseyourperformancecausedhertoget thescoreshedid,orviceversa Whatacorrelationdoessuggestisthatthetwovariablesarelinkedinsomeway directlyorindirectly.Suchacorrelationoftenstimulatesinvestigatorstoformulatehypothesesandtotestthemby somaticorbehavioralintervention Onlybymovingontosuchinterventionapproachescanweestablishwhetherone variableiscausingchangesintheother
CombiningthesethreeapproachesyieldsthecirclediagramofFigure13D,incorporatingthebasicapproachesto studyingrelationships betweenbodilyprocesses and behavior. It also emphasizes the theme that the relations betweenbrainandbehaviorarereciprocal:eachaffectstheotherinanongoingcycleofbodilyandbehavioral interactions Wewillseeexamplesofthisreciprocalrelationshipthroughoutthebook
Neuroplasticity:behaviorcanchangethebrain Theideathatthereisareciprocalrelationshipbetweenbrainandbehaviorhasembeddedwithinitaconceptthatis, formostpeople,startling Whenwesaythatbehaviorandexperienceaffectthebrain,wemeanthatthey,literally, physicallyalterthebrain ThebrainofachildgrowingupinaFrench-speakinghouseholdassemblesitselfintoa configurationdifferentfromthatofthebrainofachildwhohearsonlyEnglish That’swhythefirstchild,asanadult, understands Frencheffortlesslywhile the second does not. Inthis case we cannottellyouwhatthe structural differencesareexactly,butwedoknowonepartofthebrainthatisbeingalteredbythesedifferentexperiences(see Chapter19)
Numerousexamples,almostallinanimalsubjects,showthatexperiencecanaffectthenumberorsizeofneurons, or the number or size ofconnections betweenneurons. This abilityofthe brain, bothindevelopment and in adulthood,tobechangedbytheenvironmentandbyexperienceiscalledneuroplasticity(orneuralplasticity,or simplyplasticity)
Todaywhenwehearthewordplastic,wethinkoftheclassofmaterialsfoundinsomanymodernproducts But originally,plasticmeant“flexible,malleable”(fromtheGreekplassein,“tomoldorform”),andthemodernmaterials werenamed plastics becausetheycanbemoldedintonearlyanyshape In1890,WilliamJames(1842–1910) describedplasticityasthepossessionofastructureweakenoughtoyieldtoaninfluencebutstrongenoughnotto yieldallatonce
Intheensuingyears,researchhasshownthatthebrainisevenmoreplastic,moreyielding,thanJamessuspected. Forexample,partsofneuronsknownasdendriticspines(seeChapter2)appeartobeinconstantmotion,changing shapeinthecourseofseconds Wewillseemanyexamplesinwhichexperiencealtersthestructureand/orfunction ofthebrain:InChapter5,you’llreadthathearingababycrycausesthemother’sbraintosecreteahormone In Chapter7,we’llseethatvisualexperienceinkittensdirectstheformationofconnectionsinthebrain.InChapter12, we’lldiscusshowamotherrat’sgroomingofherpupsaffectsthesurvivalofspinalcordneurons AndChapter17 talksabouthowaseasluglearningataskchangestheconnectionsbetweentwoparticularneurons
MediaClip11:Neuroplasticity
Behavioralneuroscienceandsocialpsychologyarerelated Theplasticityofthehumanbrainhasaremarkableconsequence:otherindividualscanaffectthephysicalstructureof yourbrain!Indeed,thewholepointofcomingtoalecturehallistohavetheinstructorusewordsandfigurestoalter yourbrainsothatyoucanretrievethatinformationinthefuture(inotherwords,teachyousomething) Manyof thesealterationsinyourbrainlastonlyuntilyoutakeanexam,buteveryonceinawhiletheinstructormaytellyou somethingthatyou’llrememberfortherestofyourlife Mostaspectsofoursocialbehaviorarelearned fromthe languagewespeaktotheclotheswewearandthekindsoffoodweeat sothemechanismsoflearningand memory(seeChapter17)areimportantforunderstandingsocialbehavior
Foranexamplefromananimalmodel,considerthefactthatratsspendalotoftimeinvestigatingthesmells aroundthem,includingthosecomingfromotherrats Cookeetal (2000)tookyoungmalerats,justweanedfrom theirmother,andraisedthemintwodifferentways:eitheraloneinseparatecages,orwithothermalesingroupcages sotheycouldengageinplay(includingalotofsniffingofeachother’sbutts) Examinationoftheseanimalsasadults foundonlyonebraindifferencebetweenthegroups:aregionofthebrainknowntoprocessodorswassmallerinthe isolatedmalesthaninthemalesraisedwithplaymates(Figure14) Wasitthelackofplay(N S Gordonetal, 2003), the lack ofodors to investigate, or the stress ofisolationthatmade the regionsmaller? Whatever the mechanism, social experience affects this brain structure. In Chapter 17 we’ll see more examples of social experiencealteringthebrain
1.4 TheRoleofPlayinBrainDevelopment Abrainregioninvolvedinprocessingodors(theposterodorsalportionof
themedialamygdala)wassmallerinmaleratshousedindividuallythaninmaleshousedtogetherandallowedtoplay Other nearbyregionswereidenticalinthetwogroups (AfterB M Cookeetal,2000 Behav BrainRes117:107–113)
Here’sanexampleofhowsocialinfluencescanaffecthumanbrainfunction Whenpeoplewereaskedtoputa handintomoderatelyhotwater(47°C),partofthebrainbecameactive,presumablybecauseofthediscomfort involved(Rainvilleetal,1997) Butpeoplewhowereledtobelievethewaterwouldbe very hothadamore activatedbrainthandidthoseledtobelievethediscomfortwouldbeminimal(Figure1.5),eventhoughthewater wasthesametemperatureforeveryone Thesociallyinducedpsychologicalexpectationaffectedthemagnitudeof thebrainresponse,eventhoughthephysicalstimuluswasexactlythesame (Bytheway,thepeoplewiththemore activatedbrainsalsoreportedfeelingmorepain)
1.5 PicturesofPain Peopletoldtoexpectonlymilddiscomfortfromputtingahandinto47°Cwater(left)showedless activationinaparticularbrainregion(theanteriorcingulatecortex)thandidpeopleexpectingmorediscomfort(right)from wateroftheverysametemperature Areasofhighactivationareindicatedbyorange,red,andwhite
Inmostcases,biologicalandsocialfactorscontinuallyinteractandaffecteachotherinanongoingseriesofevents asbehaviorunfolds Forexample,thelevelofthehormonetestosteroneincirculationcanaffectdominancebehavior andaggression(seeChapter15).Thedominancemaybeexhibitedinagreatvarietyofsocialsettings,rangingfrom playingchesstophysicalaggression Inhumansandotherprimates,theleveloftestosteronecorrelatespositively withthedegreeofdominanceandwiththeamountofaggressionexhibited Winningacontest,whetheragameof chessoraboxingmatch,raisestheleveloftestosterone;losingacontestlowersthelevel Thus,atanymomentthe leveloftestosterone is determined, inpart, byrecent dominant-submissive socialexperience, and the levelof testosteronedetermines,inpart,thedegreeofdominanceandaggressioninthefuture Ofcourse,socialandcultural factorsalsohelpdeterminethefrequencyofaggression;cross-culturaldifferencesinratesofaggressionexistthat cannotbecorrelatedwithhormonelevels,andwaysofexpressingaggressionanddominanceareinfluencedby socioculturalfactors.
Perhaps nothing distinguishes neuroscience from the other sciences more clearly than this fascination with neuroplasticityandtheroleofexperience.Neuroscientistshaveapervasiveinterestinhowexperiencephysically altersthebrainandthereforeaffectsfuturebehaviorWewilltouchonthisthemeineverychapterofthisbook
StudyQuestions [PleaseNote:YoumusthaveanInternetconnectiontoviewthiscontent]
1.3.4
LearningObjectives Afterreadingthissection,youshouldbeableto:
Nameanddescribethescientificapproachofexplainingmechanismsatsimplerandsimplerlevels
Giveasurveyofimportantongoingquestionsabouttherelationshipbetweenthebrainandbehavior
Offerestimatesoftheextentofneurologicalandpsychiatricdisorders.
Explaintheimportanceofresearchwithanimalsforneuroscience,anddiscusstheethicsofsuchresearch.
Scientificexplanationsofsystemsorstructuresorfunctionsusuallyinvolvebreakingthemdownintosmallerparts,as awayofunderstandingthem Thisapproachisknownasreductionism Inprinciple,itispossibletoreduceeach explanatoryseriesdowntothemolecularoratomiclevel,thoughforpracticalreasonsthisextentofreductionismis rare Forexample,mostchemistsdealwithlarge,complexmoleculesandthelawsthatgovernthem;seldomdothey seekexplanationsintermsofsubatomicquarksandbosons
Understandingbehavioroftenrequiresseverallevelsofbiologicalanalysis Theunitsofeachlevelofanalysisare simpler instructure and organizationthanthose ofthe levelabove. The levels ofanalysis range fromsocial interactionstothebrain,continuingtosuccessivelylesscomplexunitsuntilwearriveatsinglenervecellsandtheir evensimpler,molecularconstituents
Naturally,inallfieldsdifferentproblemsarecarriedtodifferentlevelsofanalysis,andfruitfulworkisoftenbeing donesimultaneouslybydifferentworkersatseverallevels(Figure1.6).Thus,intheirresearchonvisualperception, cognitiveneuroscientistsadvanceanalyticaldescriptionsofbehavior Theytrytodeterminehowtheeyesmovewhile lookingatavisualpattern,orhowthecontrastamongpartsofthepatterndeterminesitsvisibilityMeanwhile,other behavioralneuroscientistsstudythedifferencesinvisualabilitiesamongspeciesandtrytodeterminetheadaptive significanceofthesedifferences.Forexample,howisthepresence(orabsence)ofcolorvisionrelatedtothelifeofa species?Atthesametime,otherinvestigatorstraceoutbrainstructuresandnetworksinvolvedindifferentkindsof visualdiscrimination Stillotherscientiststrytoascertaintheelectricalandchemicaleventsthatoccurinthebrain duringvision
1.6 LevelsofAnalysisinBehavioralNeuroscience Thescopeofbehavioralneurosciencerangesfromthelevelofthe individualinteractingwithothers,tothelevelofthemolecule Dependingonthequestionathand,investigatorsuse differenttechniquestofocusonthesemanylevels,butalwayswithaneyetowardhowtheirfindingsapplytobehavior
Wewillencountermanydiversebrainandbehaviortopics Herearesomeexamplesofresearchtopicsconsideredinthisbook:
Howdoesthebraingrow,maintain,andrepairitselfoverthelife-span(seeChapter7),andhowarethese capacitiesrelatedtothegrowthanddevelopmentofthemindandbehaviorfromthewombtothetomb?
Howdoesthenervoussystemcapture,process,andrepresentinformationabouttheenvironment?Forexample, sometimesbraindamagecausesapersontolosetheabilitytoidentifyotherpeople’sfaces(seeChapter18); whatdoesthattellusabouthowthebrainrecognizesfaces?
Howdoessexualorientationdevelop?Somebrainregionsaredifferentinstraightversusgaymen(seeFigure 1226);what,ifanything,dothosedifferencestellusaboutthedevelopmentofhumansexualorientation? Somepeoplesufferdamagetothebrainandafterwardseemalarminglyunafraidindangeroussituationsand unabletojudgethefearfulnessofotherpeople;whatpartsofthebrainaredamagedtocausesuchchanges(see
Figure1516)?
Howdoesthebrainmanagetochangeduringlearning(seeChapter17),andhowarememoriesretrieved? Whatbrainsitesandactivitiesunderliefeelingsandemotionalexpression?Areparticularpartsofthebrainactive inromanticlove,forexample(Figure17A)?
Whyaredifferentbrainregionsactiveduringdifferentlanguagetasks(Figure17B)?
1.7 “TellMeWhereIsFancyBred,OrintheHeartOrintheHead?” (A)Thepartsofthebrainhighlightedhere becomeespeciallyactivewhenapersonthinksabouthisorherromanticpartner (B)Differentbrainregionsareactivated whenpeopleperformfourdifferentlanguagetasks ThetechniquesusedtogeneratesuchimagesaredescribedinChapter 2
Therelationshipbetweenthebrainandbehavioris,ontheonehand,verymysteriousbecauseitisdifficultto understandhowaphysicaldevice,thebrain,couldberesponsibleforoursubjectiveexperiencesoffear,love,and awe Yetdespitethismystery,wealluseourbrainseveryday Perhapsitisthe“everydaymiracle”aspectofthe topicthathasgeneratedsomuchfolkwisdomaboutthebrain.Thinkofitas“neuromythology.”
Sometimesthesepopularideasaboutthebrainareinlinewithourcurrentknowledge,butinmanycasesweknow theyarefalse Forexample,thenotionthatwenormallyuseonly10%ofourbrainiscommonplace asurveyof teachersfoundthatnearlyhalfofthemagreedwiththisnotion(Howard-Jones,2014) butitispatentnonsense Brainscansmakeitclearthattheentirebrainisactivatedbyevenfairlymundanetasks.Indeed,althoughtheareasof activationshowninFigure17appearrathersmallanddiscrete,wewillshowinBox23thatexperimentersmust work veryhard to create images that separate activationrelated to a particular task fromthe background of widespread,ongoingbrainactivity
WeofferalistofothercommonlyheldbeliefsaboutthebrainandbehavioronthewebsiteinASTEPFURTHER 11:NEUROMYTHOLOGY:FACTSORFABLES?Throughoutthebookweoffersuchopportunitiesforyouto exploreagiventopicinmoredetailonthewebsite,bn9ecom
Behavioralneurosciencecontributestoourunderstandingofhumandisorders
Oneofthegreatpromisesofneuroscienceisthatitcanhelpusunderstandbraindisordersanddevisetreatment strategies Likeanyothercomplexmechanism,thebrainissubjecttoavarietyofmalfunctionsandbreakdowns Peopleafflictedbydisordersofthebrainarenotanexoticfew aEuropeansurveyestimatedthatatleast38%of thepopulationwouldsufferfromamentaldisorderatsomepointinatypicalyear(Wittchenetal,2011) Atleast onepersoninfivearoundtheworldcurrentlyhasneurologicaland/orpsychiatricdisordersthatvaryinseverityfrom mildchallengestocompletedisabilityFigure18AshowstheestimatednumbersofUS residentsafflictedbysome ofthemainneurologicaldisorders Figure18Bgivesestimatesofthenumbersofadultsworldwidewithcertain psychiatricdisorders ThepercentageofUS adultssufferingfrommentalillnessmaybeincreasing(Twenge,2015; Twengeetal.,2010).
1.8 TheTollofBrainDisorders EstimatednumbersofpeopleintheUnitedStateswithneurologicaldisorders(A)and numberofpeopleworldwidewithpsychiatricdisorders(B) (PartAafterC L Goochetal,2017 AnnNeurol81:479–484;BafterH RitchieandM Roser,2019 "MentalHealth" PublishedonlineatOurWorldInDataorg Retrievedfrom: https://ourworldindataorg/mental-health Underlyingdataavailablefromhttp://ghdxhealthdataorg/gbd-results-tool)
Thetollofthesedisordersisenormous,intermsofbothindividualsufferingandsocialcosts(Demyttenaereetal, 2004).TheNationalAdvisoryMentalHealthCouncilhasestimatedthatdirectandindirectcostsofbehavioraland braindisordersamountto$400billionayearintheUnitedStatesalone Forexample,thecostfortreatmentof dementia(severelydisorderedthinking)exceedsthecostsoftreatingcancerandheartdiseasecombined TheWorld HealthOrganization(2004)estimatesthatover15%ofalldiseaseburden,intermsoflostproductivity,isdueto mentaldisorders. The highcost insufferingand expense has compelled researchers to tryto understand the mechanismsinvolvedinthesedisordersandtotrytoalleviateorevenpreventthem
Inthisquest,thedistinctionbetweenclinicalandlaboratoryapproachesbeginstofadeawayForexample,when cliniciansencounterapairoftwins,oneofwhomhasschizophreniawhiletheotherseemshealthy,thediscoveryof structuraldifferences intheir brains (Figure 1.9) immediatelyraises questions for laboratoryscientists:Did the structuraldifferences arise before the symptoms ofschizophrenia, or the other way around? Were the brain differencespresentatbirth,ordidtheyariseduringpuberty?Doesmedicationthatreducessymptomsaffectbrain structure?Whengenesassociatedwithschizophreniainpeopleareintroducedintomice,willthatchangethemouse brains(seeFigure16.7)?Thislastquestionisjustoneinstanceofwhenworkingwithanimalsisessential,anissuewe addressnext
19 IdenticalTwinsbutNonidenticalBrainsandBehavior Intheseimagesofthebrainsofidenticaltwins,thefluidfilledcerebralventriclesareprominentasdark“butterfly”shapes Thebrainofthetwinwithschizophrenia(A)hasthe enlargedcerebralventriclesthatsomeresearchersbelievearecharacteristicofthisdisorder.Theothertwindoesnothave schizophrenia;hisbrain(B)clearlyhassmallerventricles
Animalresearchmakesvitalcontributions Becausewewilldrawonanimalresearchthroughoutthisbook,wewanttocommentonsomeoftheethicalissuesof experimentationonanimals Humanbeings’involvementandconcernwithotherspeciespredatesrecordedhistory Earlyhumanshadtostudyanimalbehaviorandphysiologyinordertoescapesomespeciesandhuntothers To studybiologicalbasesofbehaviorinevitablyrequiresresearchonanimalsofotherspeciesaswellasonhuman beings Psychologystudentsusuallyunderestimatethecontributionsofanimalresearchbecausethemostwidelyused introductorypsychologytextbooksoftenpresentmajorfindingsfromanimalresearchasiftheywereobtainedwith humanparticipants(DomjanandPurdy,1995) Becauseoftheimportanceofcarefullyregulatedanimalresearchforbothhumanandanimalhealthandwell-being,
the NationalResearch Council(NRC Committee on Animals as Monitors of EnvironmentalHazards, 1991) undertookastudyonthemanyusesofanimalsinresearch.Thestudynotedthat93%ofthemammalsusedin researcharelaboratory-rearedrodents ItalsoreportedthatmostAmericansbelievethatanimalresearchshould continue Ofcourse,researchershaveanobligationtominimizethediscomfortoftheiranimalsubjects,andironically enough,animalresearchhasprovideduswiththedrugsandtechniquestomakemostresearchpainlessforthe animalsubjects(SunsteinandNussbaum,2004).
Nevertheless,averyactiveminorityofpeoplebelievethatresearchwithanimals,evenifitdoesleadtolasting benefits,isunethical Forexample,inhis1975book AnimalLiberation,PeterSingerassertsthatresearchwith animalscanbejustifiedonlyifitactuallyproducesbenefits Thetrick,ofcourse,ishowtopredictwhichexperiment willleadtoabreakthrough.ThusSingerrefusestosaythatanimalexperimentationisneverjustified(Neale,2006). In the meantime, animalrights groups have vandalized labs, burned down buildings, and exploded bombs in laboratories(ConnandParker,2008) In2008,animalrightsextremistssetofffirebombsatthehomesoftwo scientistsinSanta Cruz, California One scientist’sfamily, includingtwo youngchildren, had to flee theirhome through a second-story window (Figure 1.10) (Paddock and La Ganga, 2008). These personal attacks on individualsappeartobeintendedtointimidateandfrightenscientists(D Grimm,2014),andtheyhavealready houndedatleastoneresearcheroutofthefield(NatureNeuroscience,2015)
110 CarFirebombedbyAnimalRightsActivists Theextremiststargetedthecarsandhomesoftwoscientistswho workwithanimalsattheUniversityofCaliforniainSantaCruzin2008 Thenextyear,thecarofaresearcheratUCLA wastorched
Perhapsinafuturewhererobotscanbemadethatlookandactlikehumans,methodswillbeavailabletoclearly seealltheprocessesatworkinaliving,workinghumanbrain.Inthemeantime,there’snosubstituteforresearch withanimalsubjects Everychapter inthis book is teemingwithinformationthat was gathered fromhumane experimentswithanimals
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