principles of cognitive neuroscience s econd … control is hierarchical 132 anatomical organization...
Post on 10-Jun-2018
214 Views
Preview:
TRANSCRIPT
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 AM©2012 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 AM©2012 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 AM©2012 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 AM©2012 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 AM©2012 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 AM©2012 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 AM©2012 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 AM©2012 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 Klüver-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 AM©2012 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 mental-state attribution 383Theory of mind in children and apes 384Empathy, sympathy, and prosocial behavior 386
Social Competition 388Social rank and stress 388Power motivation and dominance contests 390
■ INTRODUCTORY BOX AUTISM 360
■ BOX 11A MEASURING IMPLICIT AND EXPLICIT RACIAL ATTITUDES 377
■ BOX 11B SOCIAL BONDS AND KINSHIP 389
11
Interoception and the somatic marker hypothesis 340In search of categories of emotional experience 343
Interactions with Other Cognitive Functions 345Emotional influences on perception and attention 345Emotional influences on memory consolidation 350
Regulation of Emotion 354
■ INTRODUCTORY BOX THE NEUROSCIENCE AND NEUROETHICS OF POSTTRAU- MATIC STRESS DISORDER 320
■ BOX 10A PSYCHOPHYSIOLOGY AND THE BRAIN-BODY LINK 326
■ BOX 10B STRESS AND THE HYPOTHALAMIC-PITUITARY- ADRENAL AXIS 352
CogNeuroFrontmatter.indd xiii 9/14/12 10:46 AM©2012 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.
xiv ■ CONTENTS
Language 393
Introduction 393
Speech 393Producing speech 393Comprehending speech 396Interpreting speech sounds 396Sentences, grammar, and syntax 397The importance of context 398
Acquiring Speech and Language 400Learning a vocabulary 400The shaping of phonemes and phones 400A critical period for language acquisition 401Mechanisms of language learning 402Effects of language deprivation 402
Theories of Language 403Is there a “universal grammar”? 404Connectionist theory 404
The Neural Bases of Language 406Neural bases for producing speech and language 406Neural bases for comprehending language 407Additional evidence from neurosurgery 408Contributions of the right hemisphere to language 413
Noninvasive Studies of Language Organization 414Evidence that the neural basis of language is fundamentally symbolic 418
Genetic Determination of Language Functions 420
Is Human Language Unique? 420
The Origins of Human Language 424
■ INTRODUCTORY BOX DYSLEXIA 394
■ BOX 12A REPRESENTING SPEECH SOUNDS IN WRITTEN FORM 398
■ BOX 12B LANGUAGE, HANDEDNESS, AND CEREBRAL DOMINANCE 411
■ BOX 12C REPRESENTING NUMBER 421
■ BOX 12D LEARNED VOCAL COMMUNICATION IN NON- HUMAN SPECIES 422
12
CogNeuroFrontmatter.indd xiv 9/14/12 10:46 AM©2012 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 ■ xv
Executive Functions 429
Introduction 429
A Taxonomy of Executive Function 429
Prefrontal Cortex: A Key Contributor to Executive Function 432
Organization and connectivity of the prefrontal cortex 434Consequences of damage to the prefrontal cortex 436
Establishing and Modifying Behavioral Rules 437Initiating 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 world 445Hierarchical models for executive function 449
Control: Matching Behavior to Context 452Conflict monitoring 453Challenges to the conflict-monitoring model 454Functional organization of dorsomedial prefrontal cortex 456
Working Memory: Maintaining Information and Rules over Time 458
Neural substrates of working memory 460
■ INTRODUCTORY BOX ENVIRONMENTAL DEPENDENCY SYNDROME 430
■ BOX 13A COMPARATIVE ANATOMY OF THE PREFRONTAL CORTEX 432
■ BOX 13B THE NEUROBIOLOGY OF INTELLIGENCE 446
■ BOX 13C REASONING 450
13
Decision Making 465
Introduction 465
Decision Making: From Rational Choice to Behavioral Economics 467
Reward and Utility 470Dopamine: Pleasure or motivation? 470Reward prediction error 473Responses to negative outcomes 476
Uncertainty: Risk, Ambiguity, and Delay 479Risk and ambiguity 479Delay: Discounting future rewards 481
14
CogNeuroFrontmatter.indd xv 9/14/12 10:46 AM©2012 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.
xvi ■ CONTENTS
Social Context 483Social rewards 484Social cooperation 485Social punishment 488
Integration: Combining and Comparing Information to Reach a Decision 489
Perceptual decision making 489Value-based decision making 491
Heuristics in Decision Making 495
Future Directions 497
■ INTRODUCTORY BOX ADDICTION TO GAMBLING 466
■ BOX 14A LEARNING VALUES AND FORMING HABITS 476
■ BOX 14B MODELING SIMPLE DECISIONS 492
■ BOX 14C NEUROMARKETING 498
Evolution and Development of Brain and Cognition 503
Introduction 503
Early Thinking about the Evolution and Development of Cognition 505
Early Brain Development 508Neuronal differentiation and myelination 508The development of neural connections 510
Linking Brain and Cognitive Development 511Brain size and the evolution of cognition 514Relative brain size and cerebral complexity 516
Evolution of Brain Development 520
Evolutionary Specializations of Brain and Behavior 522Evolution and development of learning and memory 523Evolution and development of quantitative cognition 526Evolution and development of social cognition 529Evolution and development of language 535
■ INTRODUCTORY BOX SAVANT SYNDROME 504
■ BOX 15A DARWIN AND THE BRAIN 507
■ BOX 15B BRAIN DIFFERENCES IN MODERN HUMANS: IMPLICATIONS FOR COGNITION 516
■ BOX 15C EVOLUTION OF HUMAN BRAIN AND COGNITION EVIDENT IN THE FOSSIL RECORD 518
15
CogNeuroFrontmatter.indd xvi 9/19/12 10:02 AM©2012 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 ■ xvii
APPENDIXThe Human Nervous System 539
Cellular Components of the Nervous System 539
Nerve Cells and Their Signaling Functions 540
Functional Organization of the Human Nervous System 543Neural circuits 543Neural systems 545
Structural Organization of the Human Nervous System 547
Major Subdivisions of the Central Nervous System 549The brainstem 550The spinal cord 550Surface features of the brain 552Internal features of the brain 557The ventricular system 561
The Brain’s Blood Supply 561
■ BOX A1 INTRACELLULAR RECORDING FROM NERVE CELLS 546
■ BOX A2 ORGANIZATION OF THE CEREBRAL CORTEX 553
■ BOX A3 ANATOMICAL TERMINOLOGY 558
Glossary 565
Illustration Credits 583
Index 591
CogNeuroFrontmatter.indd xvii 9/14/12 10:46 AM©2012 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.
top related