PRINCIPLES OF Cognitive Neuroscience S ECOND DITION
PRINCIPLES OFCognitiveNeuroscienceSECOND EDITIONDale PurvesRoberto CabezaScott A. HuettelKevin S. LaBarMichael L. PlattMarty G. WoldorffContributorElizabeth M. BrannonCenter for Cognitive NeuroscienceDuke UniversitySinauer Associates, Inc. PublishersSunderland, MA U.S.A. Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd iii 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
Contents in BriefCHAPTER 1Cognitive Neuroscience: Definitions, Themes, and ApproachesCHAPTER 2The Methods of Cognitive Neuroscience17CHAPTER 3Sensory Systems and Perception: Vision55CHAPTER 4Sensory Systems and Perception: Auditory, Mechanical,and Chemical Senses 93CHAPTER 5Motor Systems: The Organization of ActionCHAPTER 6Attention and Its Effects on Stimulus ProcessingCHAPTER 7The Control of AttentionCHAPTER 8Memory: Varieties and MechanismsCHAPTER 9DeclarativeMemory1131167205243279CHAPTER 10Emotion319CHAPTER 11SocialCHAPTER 12LanguageCHAPTER 13Executive unctionsFCHAPTER 14DecisionCHAPTER 15Evolution and Development of Brain and CognitionCognition359393Making429465APPENDIX The Human Nervous System503539 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd v 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
Contents1Cognitive Neuroscience:Definitions, Themes, and Approaches 1IntroductionCognition12Natural philosophy and early psychologyBehaviorism 2Cognitive science 3Neuroscience25Cognitive Neuroscience:The Neurobiological Approach to Cognition9Methods: Convergence and ComplementarityConclusions1014 BOX 1A CONVERGENCE THROUGH META-ANALYSIS 122The Methods of CognitiveNeuroscience 17Introduction17Brain Perturbations That Elucidate Cognitive FunctionsPerturbations imposed by stroke, trauma, or diseasePharmacological perturbations 21Perturbation by intracranial brain stimulation 26Perturbation by extracranial brain stimulation 26Optogenetics 28Measuring Neural Activity during Cognitive ProcessingDirect electrophysiological recording from neuronsElectroencephalography (EEG) 31Event-related potentials (ERPs) 34Magnetoencephalography (MEG) 36Positron emission tomography (PET) imaging 3819192929 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd vi 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
CONTENTS viiFunctional magnetic resonance imaging (functional MRI or fMRI) 39Using fMRI to analyze activation patterns within a brain area 42Using fMRI to examine activity relationships between brain areas 44Optical brain imaging 45Assembling Evidence and Delineating Mechanisms46Associations and dissociations 46Multimethodological approaches 48 INTRODUCTORY BOX EARLY BRAIN MAPPINGIN HUMANS 18 BOX 2A AN INTRODUCTION TO STRUCTURALBRAIN IMAGING TECHNIQUES 22 BOX 2B IMAGING STRUCTURAL CONNECTIONSIN THE BRAIN 24 BOX 2C NEUROIMAGING GENOMICS 513Sensory Systemsand Perception: Vision 55IntroductionVisual Stimuli5555The Initiation of Vision56Subcortical Visual ProcessingCortical Visual Processing5961Other Key Characteristics of the Visual Cortex64Topography 64Cortical magnification 66Cortical modularity 66Visual receptive fields 67Visual Perception69Lightness and brightness 69Color 73Form 76Distance and depth 79Motion 83Object recognition 85Perceiving remembered images88 INTRODUCTORY BOX PROSOPAGNOSIA 56 BOX 3A SYNESTHESIA 65 BOX 3B MEASURING PERCEPTION 70 BOX 3C THE INVERSE PROBLEM 78 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd vii 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
viii CONTENTS4Sensory Systems and Perception: Auditory,Mechanical, and Chemical Senses 93Introduction93The Auditory System93Sound stimuli 93The peripheral auditory system 96The auditory cortices 100The perception of sound 100Perceiving the location of sound sourcesThe Mechanosensory Systems110The cutaneous/subcutaneous systemThe pain system 116The Chemosensory ModalitiesThe olfactory system 120The taste system 122Trigeminal chemosensation107110119124Some Final Points about Sensory Systems124Coding and labeled lines 124The malleability of sensory circuitry 124Awareness of sensory stimuli 126The representation of sensory percepts 127 INTRODUCTORY BOX THE REMARKABLE SUCCESS OFCOCHLEAR IMPLANTS 94 BOX 4A MEASURING LOUDNESS 101 BOX 4B MUSIC AND ITS EFFECTS 103 BOX 4C SOMATOSENSORY ILLUSIONS 112 BOX 4D PHANTOM LIMBS 1155Motor Systems:The Organization of Action 1317Introduction131Motor Control Is Hierarchical132Anatomical organization of motor systemsCortical Pathways for Motor Control133137Organization of the primary motor cortex 139Movement maps in the primary motor cortex 141Coding Movements by the Activity of Neuronal Populations 143Planning Movements144Selecting goals for action 146Motivational control of goal selection148 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd viii 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
CONTENTS ixSequential Movements and the Supplementary Motor Area 149Sensory-Motor Coordination151Initiation of Movement by the Basal GangliaBasal Ganglia and Cognition152156Error Correction and Motor Coordination by theCerebellum 159Cerebellar Contributions to Cognitive Behavior161 INTRODUCTORY BOX APRAXIA 132 BOX 5A REFLEXES, CENTRAL PATTERN GENERATORS,AND RHYTHMIC BEHAVIORS 135 BOX 5B MOTOR CONTROL OF FACIAL EXPRESSIONS 138 BOX 5C MOTOR SYSTEMS AND INTERVAL TIMING 1536Attention and Its Effectson Stimulus Processing 167Introduction167The Concept of Attention169Global states, arousal, and attention 169The selective nature of attention 169Behavioral Studies of Attention Capacity and SelectionThe level at which selection occurs 170Endogenously versus exogenously driven selective attentionNeuroscience Approaches to Studying Attention170172174Studying the neural effects of attention on stimulus processingStudying the control of attention in the brain 175175Neural Effects of Attention on Stimulus Processing:Auditory Spatial Attention 176Electrophysiological studies of the effects of auditory spatialattention 176Neuroimaging studies of the effects of auditory spatial attentionAnimal studies of the effects of auditory spatial attention 180The effects of auditory spatial attention on auditory featureprocessing 180178Neural Effects of Attention on Stimulus Processing:Visual Spatial Attention 182Electrophysiological studies of the effects of visual spatial attention 182Neuroimaging studies of the effects of visual spatial attention 184Combining electrophysiological and neuroimaging studies of visualspatial attention 189Animal studies of the effects of visual spatial attention 189The effects of visual spatial attention on visual feature processing 194Neural Effects of Attending to Nonspatial StimulusAttributes 195 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd ix 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
x CONTENTSThe neural effects of attention to nonspatial auditory features 195The neural effects of attention to nonspatial visual features 196The effects of visual attention to objects 198Neural Effects of Attention across Sensory Modalities200 INTRODUCTORY BOX THE COCKTAIL PARTY EFFECT 168 BOX 6A THE ATTENTIONAL BLINK AND LATEATTENTIONAL SELECTION 186 BOX 6B ATTENTION-RELATED “REENTRANT”ACTIVITY 1907The Control of Attention 205Introduction 205Clinical Evidence for Brain Regions Involved in AttentionalControl 206Control of Voluntary Attention210Activation in frontal and parietal cortex during endogenous attentionaltasks 210Delineating the role of the frontoparietal network in the controlof attention 210Ascertaining the temporal flow of brain activations underlyingattentional control 212Single-neuron recordings in frontal and parietal cortex duringattentional control 214Preparatory activation of sensory cortices during attentional control 216Control of Exogenously Induced Changes in Attention218Attentional shifts triggered by sudden stimulus onsets 218Attentional reorienting activates a ventral frontoparietal systemVisual Search219220Behavioral studies of visual search 220Theoretical models of visual search 221Neural processes underlying visual search223Attentional Control as a System of Interacting Brain Areas 224Interactions between Components of the AttentionalSystem 226Generality of Attentional Control Systems228Attention, Levels of Arousal, and Consciousness231Sleep and wakefulness 232Consciousness 234Neural correlates of consciousness in normal subjects 234Neural correlates of consciousness in pathological conditions237 INTRODUCTORY BOX HEMISPATIAL NEGLECTSYNDROME 206 BOX 7A THE DEFAULT-MODE NETWORK 229 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd x 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
CONTENTS8 xiMemory:Varieties and Mechanisms 243Introduction 243Memory Phases, Processes, Systems, and Tasks 245Dissociating Memory Systems 248Working memory versus declarative memory 248Declarative versus nondeclarative memory 249Nondeclarative Memory 251Priming 252Perceptual priming 253Conceptual priming 255Semantic priming 256Repetition enhancement 257Skill Learning 258Motor skill learning 259Perceptual skill learning 261Cognitive skill learning 263Conditioning 266Cellular Mechanisms of Memory 270Habituation and sensitization 272Long-term potentiation and depression 273Linking LTP to memory performance 274Learning-related changes in synaptic morphology275 INTRODUCTORY BOX THE CASE OF H.M. 244 BOX 8A INVESTIGATING DECLARATIVE MEMORY INNON-HUMAN ANIMALS 246 BOX 8B MEDIAL TEMPORAL LOBE CONTRIBUTIONSBEYOND DECLARATIVE MEMORY 251 BOX 8C CONNECTIONIST MODELS 2719Declarative MemoryIntroduction279279Basic Concepts and Assumptions 281A taxonomy of declarative memory 281A simple neurological model of encoding, storage, and retrievalUsing the model to explain the effects of brain damage 284The Nature of Medial Temporal Lobe RepresentationsTheories of hippocampal memory function 286Differences between medial temporal lobe subregions283285290 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd xi 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
xii CONTENTSCortical Regions Storing Semantic and Episodic MemoryRepresentations 293The organization of semantic knowledge in the cortex 293The reactivation of cortical regions for recent episodic memories297Contributions of the Prefrontal Cortex to Encoding andRetrieval 300Functional neuroimaging of episodic encoding 300Functional neuroimaging of episodic retrieval 303Effects of frontal lobe lesions 306Contributions of the Posterior Parietal Cortex to Encoding andRetrieval 308The role of posterior parietal cortex during retrieval 308The role of posterior parietal cortex during encoding 311Memory Consolidation312Synaptic versus system consolidation 312Theories of system consolidation in declarative memoryConsolidation, reactivation, and sleep 314313 INTRODUCTORY BOX DEVELOPMENTAL AMNESIA 280 BOX 9A ORGANIZATION OF THE MEDIAL TEMPORALLOBE MEMORY SYSTEM 286 BOX 9B FUNCTIONAL NEUROIMAGING METHODS TOSTUDY EPISODIC MEMORY 302 BOX 9C ERP STUDIES OF EPISODIC RETRIEVAL 30910Emotion 319Introduction319What Is Emotion?321Psychological Classification of EmotionsCategorical theories 322Dimensional theories 323Component process theories322325Early Neurobiological Theories of Emotion325The James-Lange feedback theory 329The Cannon-Bard diencephalic theory 329The Papez circuit and Klüver-Bucy syndrome 331The limbic system theory and its challenges 332Contemporary Approaches to Studying the Neurobiology ofEmotion 334Hemispheric-asymmetry hypotheses 334Vertical integration models: Fear acquisition 337Vertical integration models: Fear modification 340 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd xii 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
CONTENTS xiiiInteroception and the somatic marker hypothesis 340In search of categories of emotional experience 343Interactions with Other Cognitive Functions345Emotional influences on perception and attention 345Emotional influences on memory consolidation 350Regulation of Emotion354 INTRODUCTORY BOX THE NEUROSCIENCE ANDNEUROETHICS OF POSTTRAUMATIC STRESS DISORDER 320 BOX 10A PSYCHOPHYSIOLOGY AND THE BRAIN-BODYLINK 326 BOX 10B STRESS AND THE HYPOTHALAMIC-PITUITARYADRENAL AXIS 35211Social Cognition 359IntroductionThe Self359361Self-reflection 363Embodiment 365Perception of Social Cues Evident in the Face and BodyFace perception 367Perception of biological motion 370Interpersonal attention and action directionSocial Categorization367372374Perception of social category information 374Stereotypes and automatic racial biases 375Monitoring and controlling racial bias 376Impression formation and trust 379Understanding the Actions and Emotions of Others380Mirror neurons 381Perspective taking and mental-state attribution 383Theory of mind in children and apes 384Empathy, sympathy, and prosocial behavior 386Social Competition388Social rank and stress 388Power motivation and dominance contests390 INTRODUCTORY BOX AUTISM 360 BOX 11A MEASURING IMPLICIT AND EXPLICIT RACIALATTITUDES 377 BOX 11B SOCIAL BONDS AND KINSHIP 389 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd xiii 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
xiv CONTENTS12Language 393IntroductionSpeech393393Producing speech 393Comprehending speech 396Interpreting speech sounds 396Sentences, grammar, and syntax 397The importance of context 398Acquiring Speech and Language400Learning a vocabulary 400The shaping of phonemes and phones 400A critical period for language acquisition 401Mechanisms of language learning 402Effects of language deprivation 402Theories of Language403Is there a “universal grammar”?Connectionist theory 404404The Neural Bases of Language406Neural bases for producing speech and language 406Neural bases for comprehending language 407Additional evidence from neurosurgery 408Contributions of the right hemisphere to language 413Noninvasive Studies of Language Organization 414Evidence that the neural basis of language is fundamentallysymbolic 418Genetic Determination of Language FunctionsIs Human Language Unique?420420The Origins of Human Language424 INTRODUCTORY BOX DYSLEXIA 394 BOX 12A REPRESENTING SPEECH SOUNDS IN WRITTENFORM 398 BOX 12B LANGUAGE, HANDEDNESS, AND CEREBRALDOMINANCE 411 BOX 12C REPRESENTING NUMBER 421 BOX 12D LEARNED VOCAL COMMUNICATION IN NONHUMAN SPECIES 422 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd xiv 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
CONTENTS13 xvExecutive Functions 429Introduction429A Taxonomy of Executive Function429Prefrontal Cortex: A Key Contributor to ExecutiveFunction 432Organization and connectivity of the prefrontal cortex 434Consequences of damage to the prefrontal cortex 436Establishing and Modifying Behavioral Rules437Initiating rules for behavior 438Inhibiting rules for behavior 441Inhibiting socially inappropriate behaviors 443Shifting among rules for behavior 444Relating rules to create higher-order models of the worldHierarchical models for executive function 449Control: Matching Behavior to Context445452Conflict monitoring 453Challenges to the conflict-monitoring model 454Functional organization of dorsomedial prefrontal cortex456Working Memory: Maintaining Information and Rules overTime 458Neural substrates of working memory460 INTRODUCTORY BOX ENVIRONMENTAL DEPENDENCYSYNDROME 430 BOX 13A COMPARATIVE ANATOMY OF THE PREFRONTALCORTEX 432 BOX 13B THE NEUROBIOLOGY OF INTELLIGENCE 446 BOX 13C REASONING 45014Decision Making 465Introduction465Decision Making: From Rational Choice to BehavioralEconomics 467Reward and Utility470Dopamine: Pleasure or motivation? 470Reward prediction error 473Responses to negative outcomes 476Uncertainty: Risk, Ambiguity, and DelayRisk and ambiguity 479Delay: Discounting future rewards479481 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.CogNeuroFrontmatter.indd xv 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.9/14/12 10:46 AM
xvi CONTENTSSocial Context483Social rewards 484Social cooperation 485Social punishment 488Integration: Combining and Comparing Information to Reacha Decision 489Perceptual decision making 489Value-based decision making 491Heuristics in Decision MakingFuture Directions495497 INTRODUCTORY BOX ADDICTION TO GAMBLING 466 BOX 14A LEARNING VALUES AND FORMING HABITS 476 BOX 14B MODELING SIMPLE DECISIONS 492 BOX 14C NEUROMARKETING 49815Evolution and Development of Brainand Cognition 503Introduction 503Early Thinking about the Evolution and Development ofCognition 505Early Brain Development508Neuronal differentiation and myelination 508The development of neural connections 510Linking Brain and Cognitive Development511Brain size and the evolution of cognition 514Relative brain size and cerebral complexity 516Evolution of Brain Development520Evolutionary Specializations of Brain and Behavior522Evolution and development of learnin
Functional magnetic resonance imaging (functional MRI or fMRI) 39 Using fMRI to analyze activation patterns within a brain area 42 Using fMRI to examine activity relationships between brain areas 44 Optical brain imaging 45 . Principl
Cognitive neuroscience is the branch of neuroscience that seeks to understand the mechanisms of the nervous system that are directly related to cognitive (mental) processes. These mechanisms are thought to reside in the brain. Because cognition refers to functions of the mind, we must begin our study of cognitive neuroscience by first examining .
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