Brain Arousal and Information Theory Brain Arousal and Information Theory Neural and Genetic Mechanisms Donald Pfaff Harvard University Press Cambridge,MassachusettsandLondon,England 2006 Copyright©2006bythePresidentandFellowsofHarvardCollege Allrightsreserved. PrintedintheUnitedStatesofAmerica LibraryofCongressCataloging-in-PublicationData Pfaff,DonaldW.,1939– Brainarousalandinformationtheory:neuralandgeneticmechanisms/ DonaldPfaff. p.;cm. Includesbibliographicalreferencesandindex. ISBN0-674-01920-2(cloth:alk.paper) 1.Arousal(Physiology) 2.Informationtheory. 3.Neurogenetics. 4.Neurophysiology. I.Title. [DNLM:1.Arousal—physiology. 2.NerveNet—physiology. 3.Neurobiology. WL103P523b2006] QP405.P432006 153—dc22 2005046338 Contents PrefaceandDedication vii 1 TowardaUniversalTheoryofBrainArousal 1 WhatIstoBeExplained?EthologyandtheMechanismsofArousal 2 OperationalDefinitionofArousal 4 AQuantitativeApproachtoPhysicalMeasurementofGeneralizedArousal 6 TheNeurobiologyofArousalConstitutesanInterestingApplicationof InformationTheory 13 ClaimsforThisChapterandIntroductiontoChaptersFollowing 24 2 AnatomyIsNotDestiny,butaLittleNeuroanatomyHelps 26 MultiplicityandRedundancyofAscendingArousalPathwaysPreventFailure 26 Primitive“MasterCells”intheBrainstemProvideaNeuroanatomic CorethatTheoreticallyMatchestheBehavioralData 42 Long-DistanceLinesTuningLocalModules 48 SummaryandHypotheticalImplicationsforHumanBehavior 51 3 ArousalIsSignaledbyElectricalDischargesinaSystemofNerveCells 55 TravelingUptheBrainstem 56 SpecialCases:OlfactionandVision 65 InformationalContentGovernsAmplitudeofResponseinNeurons RelatedtoArousal 66 CerebralCortex,theEEG 68 ElectricalActivityinThreeArousal-RelatedBiologicalSystems 68 4 AutonomicNervousSystemChangesSupportingArousal; theUnityoftheBody 71 PatternsofAutonomicResponses 72 Reformulations 77 AHighInformationSystemShowsCoordinationsansCorrelation 78 SupportingHormone-DependentBehaviors 81 Summary 82 5 GenesWhoseNeurochemicalProductsSupportArousal 83 GenesAssociatedwithClassicalSystems 83 GenesNewlyRecognized 90 ConceptsandQuestions 95 Summary 98 6 HeightenedStatesofArousal:SexComparedtoFear 99 SexBehavior’sCNSMechanismsRequireArousal 100 GeneralizedArousalAffectsSpecificArousalsandViceVersa 107 ContrastSexandFear 115 ApplicabilityofInformationTheory 119 LibidoandStressinHumans 120 Summary 124 7 MajorSystemsQuestionsaboutBrainArousalNetworks 125 WhatAreUniversalOperatingFeaturesofArousalSystems? 126 HowDoWeMeettheRequirementforRapidChangesofCNSState? 129 SensitivityandAlacrityofResponse,YetStability?How? 131 DoesAutomataTheoryApply? 132 QuestionsintheTimeDomain 136 QuestionsaboutSpatialProperties 137 Thermodynamics,InformationTheory,andQuestionsfortheCNS 138 HowDoesaSineWaveImpactaSawtooth? 140 UnityfromDiversity? 141 8 SummaryandPracticalImportance:FromBiologicalMechanismsto HealthApplications 143 MainPoints 143 ApplicationstoHumanConditions 147 WorksCited 155 Acknowledgments 197 Index 199 vi Contents Preface and Dedication ThisbookisdedicatedtoFredPlum,M.D.,UniversityProfessorEmeritusof Neurology and former Chair of Neurology at Cornell University School of Medicine.Heroicallyhedevotedhiseffortstotheunderstandingandamelio- rationofdisordersofhumanCNSarousal.Hewasgeneroustomeinarrang- ing Wednesday lunches with brilliant younger members of his department, Nicholas Schiff, M.D. (Director, Laboratory of Cognitive Neuromodula- tionandAssistantProfessorofNeurologyandNeuroscience),KeithPurpura, Ph.D. (Associate Professor of Neuroscience), Jonathan Victor, M.D., Ph.D. (Professor of Neurology), and with his colleague then at Cornell, Michael Posner,Ph.D.Talkingwiththem,IrealizedthatIcouldexpandmyownlab’s workfromexplainingthemechanismsofsimpleinstinctivebehaviorsandspe- cific forms of motivation to more general arousal states. But I had not read aboutthelattersincecross-registeringfromM.I.T.toHarvardMedicalSchool morethan30yearsbefore.The“pilgrimages”frommyRockefellerlabacross the street to Fred Plum’s conference room supplied much of the background andcourageneededtoattemptthisbook. IalsowishtothankParParekh,ofTheRockefellerUniversity,forhisex- pert management of the illustrations and tables and extensive textual criti- cisms. Thebookisintendedtoproposenewideasformycolleaguesinthisfield, butnottosubstituteforanAnnualReviewsarticleinitsscientificdetail.There- fore, it is heavily referenced to the original literature and to pertinent review papersforproperscholarlysupport.Ihavetriedtowriteitinanopenandclear fashion, so that readers with college educations in science could understand the main ideas, and appreciate my excitement as a neurobiologist working in thisfieldatthistime. Brain Arousal and Information Theory
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