The Prefrontal Cortex as an Executive, Emotional, and Social Brain PDF
Preview The Prefrontal Cortex as an Executive, Emotional, and Social Brain
Masataka Watanabe Editor The Prefrontal Cortex as an Executive, Emotional, and Social Brain The Prefrontal Cortex as an Executive, Emotional, and Social Brain Masataka Watanabe Editor The Prefrontal Cortex as an Executive, Emotional, and Social Brain Editor MasatakaWatanabe DepartmentofPhysiologicalPsychology TokyoMetropolitanInstituteofMedicalScience Tokyo,Japan ISBN978-4-431-56506-2 ISBN978-4-431-56508-6 (eBook) DOI10.1007/978-4-431-56508-6 LibraryofCongressControlNumber:2017931562 ©SpringerJapanKK2017 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilarmethodologynowknownorhereafterdeveloped. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publicationdoesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexempt fromtherelevantprotectivelawsandregulationsandthereforefreeforgeneraluse. Thepublisher,theauthorsandtheeditorsaresafetoassumethattheadviceandinformationinthis book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained hereinor for anyerrors oromissionsthat may havebeenmade. Thepublisher remainsneutralwith regardtojurisdictionalclaimsinpublishedmapsandinstitutionalaffiliations. Printedonacid-freepaper ThisSpringerimprintispublishedbySpringerNature TheregisteredcompanyisSpringerJapanKK Theregisteredcompanyaddressis:ChiyodaFirstBldg.East,3-8-1Nishi-Kanda,Chiyoda-ku,Tokyo 101-0065,Japan Preface The prefrontal cortex (PFC) is phylogenetically most developed in humans, occupying about one-third of the cerebral cortex. It takes more than 20 years to fullydevelopandisthefirstregionofthebraintodecline.Untilrecently,thispartof the brain was called the “silent areas” or “uncommitted cortex”, because neither stimulation nor injury to this area induces noticeable sensorimotor signs. About 50yearsago,H-L.Teuberwroteachapterinthestillveryinfluentialbookonthe PFC The Frontal Granular Cortex and Behavior (JM Warren and K Akert, eds., New York, McGraw Hill, 1964) titled “The Riddle of Frontal Lobe Function in Man”(pp.410–444).Inhiswords:“Therecertainlyisnoothercerebralstructurein whichlesionscanproducesuchawiderangeofsymptoms,fromcruelalternations incharactertomildchangesinmoodthatseemtobecomeundetectableinayearor two after the lesion”. Twenty years later, PS Goldman-Rakic called PFC regions “uncharted provinces of the brain” (Trends in Neuroscience, 1984, 7:426–429), indicatinghowthisbrainareahadnotbeenfullyexplored. Atleastuntiltheendofthetwentiethcentury,althoughclinical,animallesions, andneurophysiologicalstudiesonthePFChadaccumulated,thePFCwasstillnot well charted and was full of riddles. Thanks to the development of neuroimaging techniques, such as positron emission tomography (PET), functional magnetic resonance imaging (fMRI), and diffusion tensor imaging (DTI), and also of noninvasive brain stimulation techniques, such as repetitive transcranial magnetic stimulation (rTMS) and transcranial direct current stimulation (tDCS), many attemptshavebeenmadetochartPFCregions,andseveralriddlesappeartohave beensolved. However, compared with other brain areas, the PFC is the area that is still the leastexploredandisstillfullofenigmas.Forexample,therehavelong been,and still are, hot discussions regarding whether the PFC has a unitary function or is functionally segregated, and when it is functionally segregated, how it is segregated. It has been well documented that the PFC plays the most important role for executive control. Also, it is well known that the PFC is important for emotional v vi Preface and motivational behavior. Recent studies also indicate that the PFC is actively involvedinsocialcognitionandsocialbehavior.However,itisstillnotwellknown how these functions are (if they are) segregated within the PFC. I have long been concernedwithinvestigatingcognitiveandmotivationaloperationsinthePFCby recording neuronal activities from task-performing monkeys. I have noticed that eachsinglePFCneuronisrelatedtoseveralaspectsoftaskevents,suchasstimulus, delay, response, and reward/no reward, as well as to several different kinds of functionssuchasperception,memory,andaction.Also,suchmulti-aspect-related neuronsareobservedacrossalmostallPFCregions,indicatingnoclearfunctional differentiation among different PFC regions. Human neuroimaging studies also have shown that several regions of the PFC are activated in relation to a certain function, and a certain PFC region is activated in relation to several functions. For example, in both neuroimaging and neurophysiological studies, motivation/ emotion-relatedneuralactivitiesareobservedinallthelateral,medial,andorbital PFC regions. And, for example, in the lateral PFC, neural activity is observed in relation to all the cognition, emotion, and social behavior. Furthermore, I have observed in our neurophysiological studies that many neurons are concerned not only with, for example, cognition and motivation, but also with the integration of cognitionandmotivation. Although there is a consensus that there is some functional segregation within thePFCwiththelateralregionmoreconcernedwithexecutivecontrol,theorbital region more concerned with motivation/emotion, and the medial region more concerned with self and social cognition/behavior, the segregation is hardly absolute, and only a kind of “gradient” in functional differentiation is observed acrossPFCregions.Indeed,recentstudiesindicateananterior-posteriorgradientin information processing where the more abstract the representation is, the more anteriortheregionthatisactiveinthePFC.Takingtheseresultsintoconsideration, itappearstobemoreappropriatetosupposethatthePFChasaunitaryfunctionas theintegratorofexecutive,emotional,andsocialfunctionswheredifferentregions areconcernedwiththeintegrationofdifferentkindsoffunctionsinanoverlapping manner. In this book, thus, I have focused on this aspect of “integration” of different kinds of function within the PFC. In Part I, “Functional Organization of the PrefrontalCortexinHumanandNonhumanPrimates”,howthePFCisfunctionally organized rather than segregated is described. In Chap. 1, Seo et al. describe how primate PFC regions including the frontopolar and dorsomedial regions are functionally differentiated/organized in relation to strategic decision making, and indicate that each of the PFC regions makes multiple contributions to improving the strategies of decision makers through experience. In Chap. 2, Tanaka et al. describe how different regions of the primate PFC differentially contribute to the performance of an analog of the Wisconsin Card Sorting task, introducing the uniqueroleofthefrontalpoleinexploratorybehavior,andarguingthattheoverall performance of the PFC goes beyond a mere sum of each subarea’s elementary function.InChap.3,Postlechallengesthecommonlyacceptedidearegardingthe working memory function of the PFC, indicating that the PFC is not so much Preface vii concerned with maintaining the information in working memory but is more concerned with attention and behavioral inhibition, by reviewing monkey neurophysiological studies as well as introducing the author’s fMRI and TMS studies. In this chapter, the author proposes a new idea regarding the temporal organizationofthePFCinrelationtoworkingmemorytaskperformance. In Part II, “The Prefrontal Cortex as an Integrator of Executive and Emotional Function”,how thePFC is involved inthe integration ofexecutiveandemotional function is described. In Chap. 4, Barbas et al. show that PFC regions associated withemotionsandcognitionarestronglylinkedandinfluenceeachotheraccording to principles based on the structural organization of the cortex, stressing the importanceoftheconnectionoftheposteriororbitofrontalcortex/anteriorcingulate cortex with the lateral PFC, whose connection is essential for the integration of emotionandcognition.InChap.5,KodamaandWatanabeintroducetheirprimate microdialysisstudiesandshowhowtheneurotransmittersdopamineandglutamate interactintheprimatePFCfortheintegrationofcognition(workingmemory)and motivation (reward). In Chap. 6, Schultz introduces human neuroimaging and monkeyneurophysiologicalstudiesregardinghow risk(whichbothcognitionand motivation are required to process) is represented in the PFC, and indicates the existenceoftheneuralactivitythatsignalsrewardriskdistinctfromrewardvalue. In Chap. 7, Dixon et al. describe how goals at different temporal scales are representedinahierarchicalanterior-posteriorgradientinthehumanPFC,referring also to the functional significance of the default mode of brain activity in goal- directedbehavior. In Part III, “The Prefrontal Cortex as an Integrator of Executive and Social Function”,howthePFCisrelatedtotheintegrationofexecutiveandsocialfunction is introduced. In Chap. 8, Isoda describes neuronal activity of the primate medial PFCobservedwhiletwomonkeysweremonitoringeachother’sactions.Theauthor indicates that the medial PFC plays pivotal roles in differentiating between one’s ownactionsandothers’actionsandinmonitoringthecorrectnessofothers’actions for adaptive social decision. In Chap. 9, Hosokawa describes neuronal activity of the primate lateral PFC during a face-to-face computer video game, introducing PFC neurons’ differential coding of the reward/no-reward depending on the presence/absence of the competitor and on the animacy of the opponent. In Chap. 10, Yaoi et al. describe their fMRI studies indicating how self-recognition isrealizedinthemedialPFC.Theyproposethatbyaccessinginternalrepresenta- tion of self and others, the medial PFC would support both our mental self and a wide variety of social activities by managing internal representation and make us socialbeings.InChap.11,SadatointroducesfMRIbrain-hyperscanningmethodsin studying real-time interaction between two persons and indicates that the shared attentioninducesinter-individualneuralsynchronizationintherightinferiorPFC. PartIV“DefaultModeofBrainActivityandthePrefrontalCortex”isconcerned withthe“defaultmodeofbrainactivity”whichisobservedwhilethesubjectisina resting state. The PFC is considered to be concerned with active processing of cognitive, motivational, and social information. However, recent studies indicate thattherearePFCregionsthatareactiveduringtherestingstate.Furthermore,there viii Preface are reports indicating the co-activation of the so-called executive PFC region and defaultPFCregioninrelationtocertainmentaloperations.InChap.12,Watanabe introducesthedefaultmodeofbrainactivityobservedinthemonkeyanddiscusses thefunctionalsignificanceoftheco-activationoftheexecutiveanddefaultregions inrelationtotheintegrationofcognitiveandmotivationaloperations.InChap.13, Koshinodescribestheco-activationofhumandefaultmodeandexecutivenetwork regions in relation to cognitive task performance, stressing that the relationship between the two networks changes dynamically during different phases within atask. The PFC is investigated by using many kinds of methodology. In this book, a varietyofmethodsininvestigatingtheintegrativefunctionofthePFCaredescribed: neuroanatomy(Chap.4),humanneuroimaging(fMRI,EEG)(Chaps.3,6,7,10,11, and13),humanstimulation(Chap.3),monkeyneuropsychology(Chap.2),monkey neurophysiology(Chaps.1,2,3,6,8,and9),monkeyneuroimaging(Chap.12),and monkeyneurochemistry(Chap.5). I hope readers of this book will obtain useful ideas about how the PFC is currently studied with a variety of methods, and how the PFC is concerned with the integration of cognitive, emotional/motivational, and social information for a betterwayoflife. Iexpressmygreatthankstoallauthorsforcontributingexcitingchapters.Ialso wishtothankDr.YasutakaOkazakiandMs.MomokoAsawaofSpringerJapanfor theirpatienthelpandproficientediting. Tokyo,Japan MasatakaWatanabe Contents PartI FunctionalOrganizationofthePrefrontalCortexinHuman andNonhumanPrimates 1 NeuralCorrelatesofStrategicDecision-MakinginthePrimate PrefrontalCortex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 HyojungSeo,SoyounKim,XinyingCai,HiroshiAbe, ChristopherH.Donahue,andDaeyeolLee 2 FunctionalDivisionAmongPrefrontalCorticalAreasinan AnalogofWisconsinCardSortingTest. . . . . . . . . . . . . . . . . . . . . 17 KeijiTanaka,MarkJ.Buckley,andFarshadA.Mansouri 3 WorkingMemoryFunctionsofthePrefrontalCortex. . . . . . . . . . 39 BradleyR.Postle PartII ThePrefrontalCortexasanIntegratorofExecutive andEmotionalFunction 4 PrefrontalCortexIntegrationofEmotionandCognition. . . . . . . . 51 HelenBarbasandMiguelA´ngelGarc´ıa-Cabezas 5 InteractionofDopamineandGlutamateReleaseinthePrimate PrefrontalCortexinRelationtoWorkingMemoryandReward. . . 77 TohruKodamaandMasatakaWatanabe 6 NeuronalRiskProcessinginHumanandMonkeyPrefrontal Cortex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 WolframSchultz 7 HierarchicalOrganizationofFrontoparietalControlNetworks UnderlyingGoal-DirectedBehavior. . . . . . . . . . . . . . . . . . . . . . . . 133 MathewL.Dixon,ManeshGirn,andKalinaChristoff ix x Contents PartIII ThePrefrontalCortexasanIntegratorofExecutive andSocialFunction 8 Self–OtherDifferentiationandMonitoringOthers’Actions intheMedialPrefrontalCortexoftheMonkey. . . . . . . . . . . . . . . 151 MasakiIsoda 9 NeuralCorrelatesofCompetitioninthePrimatePrefrontal Cortex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169 TakayukiHosokawa 10 Self-RecognitionProcessintheHumanPrefrontalCortex. . . . . . . 187 KenYaoi,MarikoOsaka,andNaoyukiOsaka 11 SharedAttentionandInterindividualNeuralSynchronization intheHumanRightInferiorFrontalCortex. . . . . . . . . . . . . . . . . 207 NorihiroSadato PartIV DefaultModeofBrainActivityandthePrefrontalCortex 12 DefaultModeofBrainActivityObservedintheLateral,Medial, andOrbitalPrefrontalCortexintheMonkey. . . .. . . . . .. . . . . .. 229 MasatakaWatanabe 13 CoactivationofDefaultModeNetworkandExecutiveNetwork RegionsintheHumanBrain. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247 HideyaKoshino
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