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

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PRINCIPLES OFCognitive Neuroscience SECOND EDITION Dale Purves Roberto Cabeza Scott A. Huettel Kevin S. LaBar Michael L. Platt Marty G. WoldorffContributor Elizabeth M. BrannonCenter for Cognitive Neuroscience Duke UniversitySinauer 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 539CogNeuroFrontmatter.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 1Introduction 1Cognition 2Natural philosophy and early psychology 2Behaviorism 2Cognitive science 3Neuroscience 5Cognitive Neuroscience: The Neurobiological Approach to Cognition 9Methods: Convergence and Complementarity 10Conclusions 14 BOX 1A CONVERGENCE THROUGH META-ANALYSIS 12ContentsThe Methods of Cognitive Neuroscience 17Introduction 17Brain 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 28Measuring Neural Activity during Cognitive Processing 29Direct electrophysiological recording from neurons 29Electroencephalography (EEG) 31Event-related potentials (ERPs) 34Magnetoencephalography (MEG) 36Positron emission tomography (PET) imaging 3812CogNeuroFrontmatter.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 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 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 51Sensory Systems and Perception: Vision 55Introduction 55Visual Stimuli 55The Initiation of Vision 56Subcortical Visual Processing 59Cortical Visual Processing 61Other Key Characteristics of the Visual Cortex 64Topography 64Cortical magnification 66Cortical modularity 66Visual receptive fields 67Visual 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 783CogNeuroFrontmatter.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 CONTENTSSensory Systems and Perception: Auditory, Mechanical, and Chemical Senses 93Introduction 93The Auditory System 93Sound stimuli 93The peripheral auditory system 96The auditory cortices 100The perception of sound 100Perceiving the location of sound sources 107The Mechanosensory Systems 110The cutaneous/subcutaneous system 110The pain system 116The Chemosensory Modalities 119The olfactory system 120The taste system 122Trigeminal chemosensation 124Some 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 115Motor Systems: The Organization of Action 1317Introduction 131Motor Control Is Hierarchical 132Anatomical organization of motor systems 133Cortical Pathways for Motor Control 137Organization of the primary motor cortex 139Movement maps in the primary motor cortex 141Coding Movements by the Activity of Neuronal Populations 143Planning Movements 144Selecting goals for action 146Motivational control of goal selection 14845CogNeuroFrontmatter.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 ixSequential Movements and the Supplementary Motor Area 149Sensory-Motor Coordination 151Initiation of Movement by the Basal Ganglia 152Basal Ganglia and Cognition 156Error Correction and Motor Coordination by the Cerebellum 159Cerebellar 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 153Attention and Its Effects on Stimulus Processing 167Introduction 167The Concept of Attention 169Global states, arousal, and attention 169The selective nature of attention 169Behavioral Studies of Attention Capacity and Selection 170The level at which selection occurs 170Endogenously versus exogenously driven selective attention 172Neuroscience Approaches to Studying Attention 174Studying the neural effects of attention on stimulus processing 175Studying the control of attention in the brain 175Neural Effects of Attention on Stimulus Processing: Auditory Spatial Attention 176Electrophysiological 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 180Neural 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 visual spatial 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 Stimulus Attributes 1956CogNeuroFrontmatter.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 CONTENTSThe Control of Attention 205Introduction 205Clinical Evidence for Brain Regions Involved in Attentional Control 206Control 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 216Control of Exogenously Induced Changes in Attention 218Attentional shifts triggered by sudden stimulus onsets 218Attentional reorienting activates a ventral frontoparietal system 219Visual Search 220Behavioral studies of visual search 220Theoretical models of visual search 221Neural processes underlying visual search 223Attentional Control as a System of Interacting Brain Areas 224Interactions between Components of the Attentional System 226Generality of Attentional Control Systems 228Attention, 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 2297The 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 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 190CogNeuroFrontmatter.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 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 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 2718Declarative Memory 279Introduction 279Basic 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 284The Nature of Medial Temporal Lobe Representations 285Theories of hippocampal memory function 286Differences between medial temporal lobe subregions 2909CogNeuroFrontmatter.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 CONTENTSCortical Regions Storing Semantic and Episodic Memory Representations 293The organization of semantic knowledge in the cortex 293The reactivation of cortical regions for recent episodic memories 297Contributions of the Prefrontal Cortex to Encoding and Retrieval 300Functional neuroimaging of episodic encoding 300Functional neuroimaging of episodic retrieval 303Effects of frontal lobe lesions 306Contributions of the Posterior Parietal Cortex to Encoding and Retrieval 308The role of posterior parietal cortex during retrieval 308The role of posterior parietal cortex during encoding 311Memory 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 309Emotion 319Introduction 319What Is Emotion? 321Psychological Classification of Emotions 322Categorical theories 322Dimensional theories 323Component process theories 325Early 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 332Contemporary Approaches to Studying the Neurobiology of Emotion 334Hemispheric-asymmetry hypotheses 334Vertical integration models: Fear acquisition 337Vertical integration models: Fear modification 34010CogNeuroFrontmatter.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 xiiiSocial Cognition 359Introduction 359The Self 361Self-reflection 363Embodiment 365Perception of Social Cues Evident in the Face and Body 367Face perception 367Perception of biological motion 370Interpersonal attention and action direction 372Social Categorization 374Perception of social category information 374Stereotypes and automatic racial biases 375Monitoring and controlling racial bias 376Impression formation and trust 379Understanding 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 386Social 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 38911Interoception and the somatic marker hypothesis 340In search of categories of emotional experience 343Interactions with Other Cognitive Functions 345Emotional influences on perception and attention 345Emotional influences on memory consolidation 350Regulation 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 352CogNeuroFrontmatter.indd xiii 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.xiv CONTENTSLanguage 393Introduction 393Speech 393Producing speech 393Comprehending speech 396Interpreting speech sounds 396Sentences, grammar, and syntax 397The importance of context 398Acquiring 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 402Theories of Language 403Is there a universal grammar? 404Connectionist theory 404The 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 413Noninvasive Studies of Language Organization 414Evidence that the neural basis of language is fundamentally symbolic 418Genetic Determination of Language Functions 420Is Human Language Unique? 420The 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 42212CogNeuroFrontmatter.indd xiv 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 xvExecutive Functions 429Introduction 429A Taxonomy of Executive Function 429Prefrontal Cortex: A Key Contributor to Executive Function 432Organization and connectivity of the prefrontal cortex 434Consequences of damage to the prefrontal cortex 436Establishing 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 449Control: Matching Behavior to Context 452Conflict monitoring 453Challenges to the conflict-monitoring model 454Functional organization of dorsomedial prefrontal cortex 456Working Memory: Maintaining Information and Rules over Time 458Neural 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 45013Decision Making 465Introduction 465Decision Making: From Rational Choice to Behavioral Economics 467Reward and Utility 470Dopamine: Pleasure or motivation? 470Reward prediction error 473Responses to negative outcomes 476Uncertainty: Risk, Ambiguity, and Delay 479Risk and ambiguity 479Delay: Discounting future rewards 48114CogNeuroFrontmatter.indd xv 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.xvi CONTENTSSocial Context 483Social rewards 484Social cooperation 485Social punishment 488Integration: Combining and Comparing Information to Reach a Decision 489Perceptual decision making 489Value-based decision making 491Heuristics in Decision Making 495Future 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 498Evolution and Development of Brain and Cognition 503Introduction 503Early Thinking about the Evolution and Development of Cognition 505Early Brain Development 508Neuronal differentiation and myelination 508The development of neural connections 510Linking Brain and Cognitive Development 511Brain size and the evolution of cognition 514Relative brain size and cerebral complexity 516Evolution of Brain Development 520Evolutionary 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 51815CogNeuroFrontmatter.indd xvi 9/19/12 10:02 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 xviiAPPENDIXThe Human Nervous System 539Cellular Components of the Nervous System 539Nerve Cells and Their Signaling Functions 540Functional Organization of the Human Nervous System 543Neural circuits 543Neural systems 545Structural Organization of the Human Nervous System 547Major Subdivisions of the Central Nervous System 549The brainstem 550The spinal cord 550Surface features of the brain 552Internal features of the brain 557The ventricular system 561The Brains Blood Supply 561 BOX A1 INTRACELLULAR RECORDING FROM NERVE CELLS 546 BOX A2 ORGANIZATION OF THE CEREBRAL CORTEX 553 BOX A3 ANATOMICAL TERMINOLOGY 558Glossary 565Illustration Credits 583Index 591CogNeuroFrontmatter.indd xvii 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.

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