The Methods of Cognitive Neuroscience. Sensory Systems and Perception: Auditory, Mechanical, and Chemical Senses 93

Transcription

1 Contents in Brief CHAPTER 1 Cognitive Neuroscience: Definitions, Themes, and Approaches 1 CHAPTER 2 The Methods of Cognitive Neuroscience CHAPTER 3 Sensory Systems and Perception: Vision 55 CHAPTER 4 CHAPTER 5 CHAPTER 6 Sensory Systems and Perception: Auditory, Mechanical, and Chemical Senses 93 Motor Systems: The Organization of Action Attention and Its Effects on Stimulus Processing CHAPTER 7 The Control of Attention 205 CHAPTER 8 Memory: Varieties and Mechanisms 243 CHAPTER 9 Declarative Memory 279 CHAPTER Emotion CHAPTER 1 1 Social Cognition 359 CHAPTER Language 393 CHAPTER Executive Functions 429 CHAPTER Decision Making 465 CHAPTER 1 5 Evolution and Development of Brain and Cognition 503 APPENDIX The Human Nervous System 539 Principles of cognitive neuroscience 2013 digitalisiert durch: IDS Basel Bern

2 Contents Cognitive Neuroscience: Definitions, and Approaches 1 Introduction 1 Cognition 2 Natural philosophy and early psychology 2 Behaviorism 2 Cognitive science 3 Neuroscience 5 Cognitive Neuroscience: The Neurobiological Approach to Cognition 9 Methods: Convergence and Complementarity 10 Conclusions 14 BOX CONVERGENCE THROUGH META-ANALYSIS 1 2 The Methods of Cognitive Neuroscience 17 Introduction 17 Brain Perturbations That Elucidate Cognitive Functions Perturbations imposed by stroke, trauma, or disease Pharmacological perturbations 21 Perturbation by intracranial brain stimulation 26 Perturbation by extracranial brain stimulation 26 Optogenetics 28 Measuring Neural Activity during Cognitive Processing 29 Direct electrophysiological recording from neurons 29 Electroencephalography (EEG) 31 Event-related potentials 34 Magnetoencephalography (MEG) 36 Positron emission tomography (PET) imaging 38

3 Vii Functional magnetic resonance imaging (functional or fmri) 39 Using to analyze activation patterns within a brain area 42 Using fmri to examine activity relationships between brain areas 44 Optical brain imaging 45 Assembling Evidence and Delineating Mechanisms 46 Associations and dissociations 46 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 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 64 Topography 64 Cortical magnification 66 Cortical modularity 66 Visual receptive fields 67 Visual Perception 69 Lightness and brightness 69 Color 73 Form 76 Distance and depth 79 Motion 83 Object recognition 85 Perceiving remembered images 88 INTRODUCTORY BOX PROSOPAGNOSIA 56 BOX 3A SYNESTHESIA 65 BOX 3B MEASURING PERCEPTION 70 BOX 3C THE INVERSE PROBLEM 78

4 Sensory Systems and Perception: Auditory, Mechanical, and Chemical Senses 93 Introduction 93 The Auditory System 93 Sound stimuli 93 The peripheral auditory system 96 The auditory cortices 100 The perception of sound Perceiving the location of sound sources The Systems The cutaneous/subcutaneous system The pain system The Modalities The olfactory system The taste system Trigeminal chemosensation Some Final Points about Sensory Systems Coding and labeled lines 124 The malleability of sensory circuitry Awareness of sensory stimuli The representation of sensory percepts INTRODUCTORY BOX THE REMARKABLE SUCCESS OF MEASURING LOUDNESS BOX 4B MUSIC AND EFFECTS COCHLEAR IMPLANTS 94 BOX 4C SOMATOSENSORY ILLUSIONS 2 BOX 4D PHANTOM LIMBS Motor Systems: The Organization of Action 131 Introduction 131 Motor Control Is Hierarchical Anatomical organization of motor systems Cortical Pathways for Motor Control Organization of the primary motor cortex Movement maps in the primary motor cortex Coding Movements by the Activity of Neuronal Populations Planning Movements 144 Selecting goals for action Motivational control of goal selection

5 ix Sequential Movements and the Supplementary Motor Area Sensory-Motor Coordination 151 Initiation of Movement by the Basal Ganglia Basal Ganglia and Cognition 156 Error Correction and Motor Coordination by the Cerebellum 159 Cerebellar Contributions to Cognitive Behavior INTRODUCTORY BOX APRAXIA BOX 5A REFLEXES, CENTRAL PATTERN GENERATORS, AND RHYTHMIC BEHAVIORS 1 35 BOX 5B MOTOR CONTROL OF FACIAL EXPRESSIONS BOX 5C MOTOR SYSTEMS AND INTERVAL TIMING 1 53 Introduction 167 The Concept of Attention Global states, arousal, and attention The selective nature of attention Behavioral Studies of Attention Capacity and Selection 170 The level at which selection occurs Endogenously versus exogenously driven selective attention Neuroscience Approaches to Studying Attention 174 Studying the neural effects of attention on stimulus processing Studying the control of attention in the brain Neural Effects of Attention on Stimulus Processing: Auditory Spatial Attention 176 Electrophysiological studies of the effects of auditory spatial attention 176 Neuroimaging studies of the effects of auditory spatial attention Animal studies of the effects of auditory spatial attention The effects of auditory spatial attention on auditory feature processing Neural Effects of Attention on Stimulus Processing: Visual Spatial Attention 182 Electrophysiological studies of the effects of visual spatial attention Neuroimaging studies of the effects of visual spatial attention Combining electrophysiological and neuroimaging studies of visual spatial attention Animal studies of the effects of visual spatial attention The effects of visual spatial attention on visual feature processing Neural Effects of Attending to Nonspatial Stimulus Attributes 195

6 The neural effects of attention to nonspatial auditory features The neural effects of attention to nonspatial visual features The effects of visual attention to objects Neural Effects of Attention across Sensory Modalities 200 INTRODUCTORY BOX THE COCKTAIL PARTY EFFECT BOX 6A THE ATTENTIONAL BLINK AND LATE ATTENTIONAL SELECTION 1 86 BOX 6B ATTENTION-RELATED "REENTRANT" ACTIVITY Clinical Evidence for Brain Regions Involved in Attentional Control 206 Control of Voluntary Attention Activation in frontal and parietal cortex during endogenous attentional tasks Delineating the role of the frontoparietal network in the control of attention Ascertaining the temporal flow of brain activations underlying attentional control Single-neuron recordings in frontal and parietal cortex during attentional control Preparatory activation of sensory cortices during attentional control Control of Exogenously Induced Changes in Attention Attentional shifts triggered by sudden stimulus onsets Attentional reorienting activates a ventral frontoparietal system Visual Search 220 Behavioral studies of visual search 220 Theoretical models of visual search 221 Neural 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 231 Sleep and wakefulness 232 Consciousness 234 Neural correlates of consciousness in normal subjects 234 Neural correlates of consciousness in pathological conditions 237 INTRODUCTORY BOX HEMISPATIAL NEGLECT SYNDROME 206 BOX 7A THE DEFAULT-MODE NETWORK 229

7 Xi Memory: Varieties and Mechanisms 243 Introduction 243 Memory Phases, Processes, Systems, and Tasks 245 Dissociating Memory Systems 248 Working memory versus declarative memory 248 Declarative versus nondeclarative memory 249 Nondeclarative Memory 251 Priming 252 Perceptual priming 253 Conceptual priming 255 Semantic priming 256 Repetition enhancement 257 Skill Learning 258 Motor skill learning 259 Perceptual skill learning 261 Cognitive skill learning 263 Conditioning 266 Cellular Mechanisms of Memory 270 Habituation and sensitization 272 Long-term potentiation and depression 273 Linking LTP to memory performance 274 Learning-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 Declarative Memory 279 Introduction 279 Basic Concepts and Assumptions 281 A taxonomy of declarative memory 281 A simple neurological model of encoding, storage, and retrieval 283 Using the model to explain the effects of brain damage 284 The Nature of Medial Temporal Lobe Representations 285 ' Theories of hippocampal memory function 286 Differences between medial temporal lobe subregions 290

8 Xii Cortical Regions Storing Semantic and Episodic Memory Representations 293 The organization of semantic knowledge in the cortex 293 The reactivation of cortical regions for recent episodic memories 297 Contributions of the Prefrontal Cortex to Encoding and Retrieval 300 Functional neuroimaging of episodic encoding 300 Functional neuroimaging of episodic retrieval 303 Effects of frontal lobe lesions 306 Contributions of the Posterior Parietal Cortex to Encoding and Retrieval 308 The role of posterior parietal cortex during retrieval 308 The role of posterior parietal cortex during encoding Memory Consolidation 312 Synaptic versus system consolidation Theories of system consolidation in declarative memory Consolidation, reactivation, and sleep INTRODUCTORY BOX DEVELOPMENTAL 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 Introduction 319 What Is Emotion? 321 Psychological Classification of Emotions Categorical theories 322 Dimensional theories 323 Component process theories 325 Early Neurobiological Theories of Emotion 325 The James-Lange feedback theory 329 The Cannon-Bard diencephalic theory 329 The Papez circuit and syndrome 331 The limbic system theory and its challenges 332 Contemporary Approaches to Studying the Neurobiology of Emotion 334 Hemispheric-asymmetry hypotheses 334 Vertical integration models: Fear acquisition 337 Vertical integration models: Fear modification 340

9 Xiii Interoception and the somatic marker hypothesis 340 In search of categories of emotional experience 343 Interactions with Other Cognitive Functions 345 Emotional influences on perception and attention 345 Emotional 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 STRESS AND THE HYPOTHALAMIC-PITUITARY- 352 Introduction 359 Self 361 Self-reflection 363 Embodiment 365 Perception of Social Cues Evident in the Face and Body 367 Face perception 367 Perception of biological motion 370 Interpersonal attention and action direction 372 Social Categorization 374 Perception of social category information 374 Stereotypes and automatic racial biases 375 Monitoring and controlling racial bias 376 Impression formation and trust 379 Understanding the Actions and Emotions of Others 380 Mirror neurons 381 Perspective taking and mental-state attribution 383 Theory of mind in children and apes 384 Empathy, sympathy, and prosocial behavior 386 Social Competition 388 Social rank and stress 388 Power motivation and dominance contests 390 INTRODUCTORY BOX AUTISM 360 BOX 11A MEASURING IMPLICIT AND EXPLICIT RACIAL ATTITUDES 377 BOX SOCIAL BONDS AND KINSHIP 389

10 xiv Introduction 393 Speech 393 Producing speech 393 Comprehending speech 396 Interpreting speech sounds 396 Sentences, grammar, and syntax 397 The importance of context 398 Acquiring Speech and Language 400 Learning a vocabulary 400 The shaping of phonemes and phones 400 A critical period for language acquisition 401 Mechanisms of language learning 402 Effects of language deprivation 402 Theories of Language 403 there a "universal grammar"? 404 Connectionist theory 404 The Neural Bases of Language 406 Neural bases for producing speech and language 406 Neural bases for comprehending language 407 Additional evidence from neurosurgery 408 Contributions of the right hemisphere to language Noninvasive Studies of Language Organization 414 Evidence that the neural basis of language is fundamentally symbolic 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 BOX 12C REPRESENTING NUMBER 421 BOX LEARNED VOCAL COMMUNICATION HUMAN SPECIES 422

11 XV Introduction 429 A Taxonomy of Executive Function 429 Prefrontal Cortex: A Key Contributor to Executive Function 432 Organization and connectivity of the prefrontal cortex 434 Consequences of damage to the prefrontal cortex 436 Establishing and Modifying Behavioral Rules 437 Initiating rules for behavior 438 Inhibiting rules for behavior 441 Inhibiting socially inappropriate behaviors 443 Shifting among rules for behavior 444 Relating rules to create higher-order models of the world 445 Hierarchical models for executive function 449 Control: Matching Behavior to Context 452 Conflict monitoring 453 Challenges to the conflict-monitoring model 454 Functional 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 Decision Making 465 Introduction 465 Decision Making: From Rational Choice to Behavioral Economics 467 Reward and Utility 470 Dopamine: Pleasure or motivation? 470 Reward prediction error 473» Responses to negative outcomes 476 Uncertainty: Risk, Ambiguity, and Delay 479 Risk and ambiguity 479 Delay: Discounting future rewards 481

12 xvi Social Context 483 Social rewards 484 Social cooperation 485 Social punishment 488 Integration: Combining and Comparing Information to Reach a Decision 489 Perceptual decision making 489 Value-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 508 Neuronal differentiation and myelination 508 The development of neural connections Linking Brain and Cognitive Development Brain size and the evolution of cognition Relative brain size and cerebral complexity Evolution of Brain Development 520 Evolutionary Specializations of Brain and Behavior 522 Evolution and development of learning and memory 523 Evolution and development of quantitative cognition 526 Evolution and development of social cognition 529 Evolution 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

13 XVii APPENDIX The 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 543 Neural circuits 543 Neural systems 545 Structural Organization of the Human Nervous System 547 Major Subdivisions of the Central Nervous System 549 The brainstem 550 The spinal cord 550 Surface features of the brain 552 Internal features of the brain 557 The 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