PRINCIPLES OF Cognitive Neuroscience ... - Sinauer OF Cognitive Neuroscience ... 2012 Sinauer Associates, Inc. ... Principles of Cognitive Neuroscience, Second Edition

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  • PRINCIPLES OF

    Cognitive Neuroscience SECOND EDITION

    Dale Purves

    Roberto Cabeza

    Scott A. Huettel

    Kevin S. LaBar

    Michael L. Platt

    Marty G. Woldorff

    Contributor Elizabeth M. Brannon

    Center for Cognitive Neuroscience Duke University

    Sinauer Associates, Inc. PublishersSunderland, MA U.S.A.

    CogNeuroFrontmatter.indd iii 9/14/12 10:46 AM2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

    Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher.

  • Contents in Brief

    CHAPTER 1 Cognitive Neuroscience: Definitions, Themes, and Approaches 1

    CHAPTER 2 The Methods of Cognitive Neuroscience 17

    CHAPTER 3 Sensory Systems and Perception: Vision 55

    CHAPTER 4 Sensory Systems and Perception: Auditory, Mechanical, and Chemical Senses 93

    CHAPTER 5 Motor Systems: The Organization of Action 131

    CHAPTER 6 Attention and Its Effects on Stimulus Processing 167

    CHAPTER 7 The Control of Attention 205

    CHAPTER 8 Memory: Varieties and Mechanisms 243

    CHAPTER 9 Declarative Memory 279

    CHAPTER 10 Emotion 319

    CHAPTER 11 Social Cognition 359

    CHAPTER 12 Language 393

    CHAPTER 13 Executive Functions 429

    CHAPTER 14 Decision Making 465

    CHAPTER 15 Evolution and Development of Brain and Cognition 503

    APPENDIX The Human Nervous System 539

    CogNeuroFrontmatter.indd v 9/14/12 10:46 AM2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

    Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher.

  • Cognitive Neuroscience: Definitions, Themes, and Approaches 1

    Introduction 1

    Cognition 2Natural philosophy and early psychology 2Behaviorism 2Cognitive science 3

    Neuroscience 5

    Cognitive Neuroscience: The Neurobiological Approach to Cognition 9

    Methods: Convergence and Complementarity 10

    Conclusions 14

    BOX 1A CONVERGENCE THROUGH META-ANALYSIS 12

    Contents

    The Methods of Cognitive Neuroscience 17

    Introduction 17

    Brain Perturbations That Elucidate Cognitive Functions 19Perturbations imposed by stroke, trauma, or disease 19Pharmacological perturbations 21Perturbation by intracranial brain stimulation 26Perturbation by extracranial brain stimulation 26Optogenetics 28

    Measuring Neural Activity during Cognitive Processing 29Direct electrophysiological recording from neurons 29Electroencephalography (EEG) 31Event-related potentials (ERPs) 34Magnetoencephalography (MEG) 36Positron emission tomography (PET) imaging 38

    1

    2

    CogNeuroFrontmatter.indd vi 9/14/12 10:46 AM2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

    Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher.

  • CONTENTS vii

    Functional 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 45

    Assembling Evidence and Delineating Mechanisms 46Associations and dissociations 46Multimethodological approaches 48

    INTRODUCTORY BOX EARLY BRAIN MAPPING IN HUMANS 18

    BOX 2A AN INTRODUCTION TO STRUCTURAL BRAIN IMAGING TECHNIQUES 22

    BOX 2B IMAGING STRUCTURAL CONNECTIONS IN THE BRAIN 24

    BOX 2C NEUROIMAGING GENOMICS 51

    Sensory Systems and Perception: Vision 55

    Introduction 55

    Visual Stimuli 55

    The Initiation of Vision 56

    Subcortical Visual Processing 59

    Cortical Visual Processing 61

    Other Key Characteristics of the Visual Cortex 64Topography 64Cortical magnification 66Cortical modularity 66Visual receptive fields 67

    Visual Perception 69Lightness and brightness 69Color 73Form 76Distance and depth 79Motion 83Object recognition 85Perceiving remembered images 88

    INTRODUCTORY BOX PROSOPAGNOSIA 56

    BOX 3A SYNESTHESIA 65

    BOX 3B MEASURING PERCEPTION 70

    BOX 3C THE INVERSE PROBLEM 78

    3

    CogNeuroFrontmatter.indd vii 9/14/12 10:46 AM2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

    Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher.

  • viii CONTENTS

    Sensory Systems and Perception: Auditory, Mechanical, and Chemical Senses 93

    Introduction 93

    The Auditory System 93Sound stimuli 93The peripheral auditory system 96The auditory cortices 100The perception of sound 100Perceiving the location of sound sources 107

    The Mechanosensory Systems 110The cutaneous/subcutaneous system 110The pain system 116

    The Chemosensory Modalities 119The olfactory system 120The taste system 122Trigeminal chemosensation 124

    Some Final Points about Sensory Systems 124Coding and labeled lines 124The malleability of sensory circuitry 124Awareness of sensory stimuli 126The representation of sensory percepts 127

    INTRODUCTORY BOX THE REMARKABLE SUCCESS OF COCHLEAR IMPLANTS 94

    BOX 4A MEASURING LOUDNESS 101

    BOX 4B MUSIC AND ITS EFFECTS 103

    BOX 4C SOMATOSENSORY ILLUSIONS 112

    BOX 4D PHANTOM LIMBS 115

    Motor Systems: The Organization of Action 1317

    Introduction 131

    Motor Control Is Hierarchical 132Anatomical organization of motor systems 133

    Cortical Pathways for Motor Control 137Organization of the primary motor cortex 139Movement maps in the primary motor cortex 141

    Coding Movements by the Activity of Neuronal Populations 143

    Planning Movements 144Selecting goals for action 146Motivational control of goal selection 148

    4

    5

    CogNeuroFrontmatter.indd viii 9/14/12 10:46 AM2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

    Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher.

  • CONTENTS ix

    Sequential Movements and the Supplementary Motor Area 149

    Sensory-Motor Coordination 151

    Initiation of Movement by the Basal Ganglia 152

    Basal Ganglia and Cognition 156

    Error Correction and Motor Coordination by the Cerebellum 159

    Cerebellar Contributions to Cognitive Behavior 161

    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 153

    Attention and Its Effects on Stimulus Processing 167

    Introduction 167

    The Concept of Attention 169Global states, arousal, and attention 169The selective nature of attention 169

    Behavioral Studies of Attention Capacity and Selection 170The level at which selection occurs 170Endogenously versus exogenously driven selective attention 172

    Neuroscience Approaches to Studying Attention 174Studying the neural effects of attention on stimulus processing 175Studying the control of attention in the brain 175

    Neural Effects of Attention on Stimulus Processing: Auditory Spatial Attention 176

    Electrophysiological studies of the effects of auditory spatial attention 176Neuroimaging studies of the effects of auditory spatial attention 178Animal studies of the effects of auditory spatial attention 180The effects of auditory spatial attention on auditory feature processing 180

    Neural Effects of Attention on Stimulus Processing: Visual Spatial Attention 182

    Electrophysiological studies of the effects of visual spatial attention 182Neuroimaging studies of the effects of visual spatial attention 184Combining electrophysiological and neuroimaging studies of visual spatial attention 189Animal studies of the effects of visual spatial attention 189The effects of visual spatial attention on visual feature processing 194

    Neural Effects of Attending to Nonspatial Stimulus Attributes 195

    6

    CogNeuroFrontmatter.indd ix 9/14/12 10:46 AM2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

    Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher.

  • x CONTENTS

    The Control of Attention 205

    Introduction 205

    Clinical Evidence for Brain Regions Involved in Attentional Control 206

    Control of Voluntary Attention 210Activation in frontal and parietal cortex during endogenous attentional tasks 210Delineating the role of the frontoparietal network in the control of attention 210Ascertaining the temporal flow of brain activations underlying attentional control 212Single-neuron recordings in frontal and parietal cortex during attentional control 214Preparatory activation of sensory cortices during attentional control 216

    Control of Exogenously Induced Changes in Attention 218Attentional shifts triggered by sudden stimulus onsets 218Attentional reorienting activates a ventral frontoparietal system 219

    Visual Search 220Behavioral studies of visual search 220Theoretical models of visual search 221Neural processes underlying visual search 223

    Attentional Control as a System of Interacting Brain Areas 224

    Interactions between Components of the Attentional System 226

    Generality of Attentional Control Systems 228

    Attention, Levels of Arousal, and Consciousness 231Sleep and wakefulness 232Consciousness 234Neural correlates of consciousness in normal subjects 234Neural correlates of consciousness in pathological conditions 237

    INTRODUCTORY BOX HEMISPATIAL NEGLECT SYNDROME 206

    BOX 7A THE DEFAULT-MODE NETWORK 229

    7

    The neural effects of attention to nonspatial auditory features 195The neural effects of attention to nonspatial visual features 196The effects of visual attention to objects 198

    Neural Effects of Attention across Sensory Modalities 200

    INTRODUCTORY BOX THE COCKTAIL PARTY EFFECT 168

    BOX 6A THE ATTENTIONAL BLINK AND LATE ATTENTIONAL SELECTION 186

    BOX 6B ATTENTION-RELATED REENTRANT ACTIVITY 190

    CogNeuroFrontmatter.indd x 9/14/12 10:46 AM2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

    Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher.

  • CONTENTS xi

    Memory: Varieties and Mechanisms 243

    Introduction 243

    Memory Phases, Processes, Systems, and Tasks 245

    Dissociating Memory Systems 248Working memory versus declarative memory 248Declarative versus nondeclarative memory 249

    Nondeclarative Memory 251

    Priming 252Perceptual priming 253Conceptual priming 255Semantic priming 256Repetition enhancement 257

    Skill Learning 258Motor skill learning 259Perceptual skill learning 261Cognitive skill learning 263

    Conditioning 266

    Cellular Mechanisms of Memory 270Habituation and sensitization 272Long-term potentiation and depression 273Linking LTP to memory performance 274Learning-related changes in synaptic morphology 275

    INTRODUCTORY BOX THE CASE OF H.M. 244

    BOX 8A INVESTIGATING DECLARATIVE MEMORY IN NON-HUMAN ANIMALS 246

    BOX 8B MEDIAL TEMPORAL LOBE CONTRIBUTIONS BEYOND DECLARATIVE MEMORY 251

    BOX 8C CONNECTIONIST MODELS 271

    8

    Declarative Memory 279

    Introduction 279

    Basic Concepts and Assumptions 281A taxonomy of declarative memory 281A simple neurological model of encoding, storage, and retrieval 283Using the model to explain the effects of brain damage 284

    The Nature of Medial Temporal Lobe Representations 285Theories of hippocampal memory function 286Differences between medial temporal lobe subregions 290

    9

    CogNeuroFrontmatter.indd xi 9/14/12 10:46 AM2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

    Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher.

  • xii CONTENTS

    Cortical Regions Storing Semantic and Episodic Memory Representations 293

    The organization of semantic knowledge in the cortex 293The reactivation of cortical regions for recent episodic memories 297

    Contributions of the Prefrontal Cortex to Encoding and Retrieval 300

    Functional neuroimaging of episodic encoding 300Functional neuroimaging of episodic retrieval 303Effects of frontal lobe lesions 306

    Contributions of the Posterior Parietal Cortex to Encoding and Retrieval 308

    The role of posterior parietal cortex during retrieval 308The role of posterior parietal cortex during encoding 311

    Memory Consolidation 312Synaptic versus system consolidation 312Theories of system consolidation in declarative memory 313Consolidation, reactivation, and sleep 314

    INTRODUCTORY BOX DEVELOPMENTAL AMNESIA 280

    BOX 9A ORGANIZATION OF THE MEDIAL TEMPORAL LOBE MEMORY SYSTEM 286

    BOX 9B FUNCTIONAL NEUROIMAGING METHODS TO STUDY EPISODIC MEMORY 302

    BOX 9C ERP STUDIES OF EPISODIC RETRIEVAL 309

    Emotion 319

    Introduction 319

    What Is Emotion? 321

    Psychological Classification of Emotions 322Categorical theories 322Dimensional theories 323Component process theories 325

    Early Neurobiological Theories of Emotion 325The James-Lange feedback theory 329The Cannon-Bard diencephalic theory 329The Papez circuit and Klver-Bucy syndrome 331The limbic system theory and its challenges 332

    Contemporary Approaches to Studying the Neurobiology of Emotion 334

    Hemispheric-asymmetry hypotheses 334Vertical integration models: Fear acquisition 337Vertical integration models: Fear modification 340

    10

    CogNeuroFrontmatter.indd xii 9/14/12 10:46 AM2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

    Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher.

  • CONTENTS xiii

    Social Cognition 359

    Introduction 359

    The Self 361Self-reflection 363Embodiment 365

    Perception of Social Cues Evident in the Face and Body 367Face perception 367Perception of biological motion 370Interpersonal attention and action direction 372

    Social Categorization 374Perception of social category information 374Stereotypes and automatic racial biases 375Monitoring and controlling racial bias 376Impression formation and trust 379

    Understanding the Actions and Emotions of Others 380Mirror neurons 381Perspective taking and me...

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