Essentials of Cognitive Neuroscience

Bradley R. Postle is Professor of Psychology and Psychiatry at the University of Wisconsin–Madison. He has taught cognitive neuroscience, cognitive psychology, and systems neuroscience at the undergraduate and graduate level for two decades. He has published over 100 scientific journal articles and chapters on the cognitive neuroscience of memory. He is Associate Editor of the Journal of Cognitive Neuroscience and eNeuro, and has previously served in this capacity at the journals NeuroImage and Cortex.

Preface xvii

Acknowledgments xix

Walkthrough of Pedagogical Features xxi

Companion Website xxii

Section I: The Neurobiology of Thinking 1

1 Introduction and History 4

Key Themes 4

A Brief (and Selective) History 6

Construct validity in models of cognition 6

Localization of function vs. mass action 7

The first scientifically rigorous demonstrations of localization of function 9

What is a Brain and What Does It Do? 12

Looking Ahead to the Development of Cognitive Neuroscience 13

End-of-Chapter Questions 14

References 14

Other Sources Used 14

Further Reading 15

2 The Brain 16

Key Themes 16

Pep Talk 18

Gross Anatomy 18

The cerebral cortex 21

The Neuron 23

Electrical and chemical properties of the neuron 23

Oscillatory Fluctuations in the Membrane Potential 28

Neurons are never truly “at rest” 28

Oscillatory synchrony 29

Complicated, and Complex 31

End-of-Chapter Questions 32

References 32

Other Sources Used 33

Further Reading 33

3 Methods for Cognitive Neuroscience 34

Key Themes 34

Behavior, Structure, Function, and Models 36

Behavior 36

Neuropsychology, neurophysiology, and the limits of inference 36

Different kinds of neuropsychology address different kinds of questions 37

How does behavior relate to mental functions? 38

Methods for lesioning targeted areas of the brain 39

Nonlocalized trauma 39

Transcranial Neurostimulation 40

The importance of specificity (again) 41

Transcranial magnetic stimulation 43

Anatomy and Cellular Physiology 47

Techniques that exploit the cell biology of the neuron 48

Electrophysiology 51

Invasive recording with microelectrodes: action potentials and local field potentials 51

Electrocorticography 53

Electroencephalography 53

Magnetoencephalography 55

Invasive Neurostimulation 55

Electrical microstimulation 55

Optogenetics 55

Analysis of Time-Varying Signals 56

Event-related analyses 56

Magnetic Resonance Imaging 61

Physics and engineering bases 61

MRI methods for in vivo anatomical imaging 64

Functional magnetic resonance imaging 65

Functional connectivity 70

Resting state functional correlations 70

Magnetic Resonance Spectroscopy 73

Tomography 73

X-ray computed tomography 73

Positron emission tomography 73

Near-Infrared Spectroscopy 76

Some Considerations For Experimental Design 76

Computational Models and Analytic Approaches 78

Neural network modeling 78

Network science and graph theory 82

End-of-Chapter Questions 84

References 85

Other Sources Used 86

Further Reading 86

Section II: Sensation, Perception, Attention, and Action 87

4 Sensation and Perception of Visual Signals 90

Key Themes 90

The Dominant Sense in Primates 92

Organization of the Visual System 92

The visual field 92

The retina and the LGN of the thalamus 92

The retinotopic organization of primary visual cortex 93

The receptive field 95

Information Processing in Primary Visual Cortex – Bottom-Up Feature Detection 96

The V1 neuron as feature detector 96

Columns, hypercolumns, and pinwheels 99

Information Processing in Primary Visual Cortex – Interactivity 100

Feedforward and feedback projections of V1 100

The relation between visual processing and the brain’s physiological state 104

Where Does Sensation End? Where Does Perception Begin? 106

End-of-Chapter Questions 106

References 107

Other Sources Used 107

Further Reading 108

5 Audition and Somatosensation 109

Key Themes 109

Apologia 111

Audition 111

Auditory sensation 111

Auditory perception 115

Adieu to audition 119

Somatosensation 119

Transduction of mechanical and thermal energy, and of pain 119

Somatotopy 122

Somatosensory plasticity 126

Phantom limbs and phantom pain 129

Proprioception 131

Adieu to sensation 131

End-of-Chapter Questions 131

References 132

Other Sources Used 132

Further Reading 132

6 The Visual System 134

Key Themes 134

Familiar Principles and Processes, Applied to Higher-Level Representations 136

Two Parallel Pathways 136

A diversity of projections from V1 136

A functional dissociation of visual perception of what an object is vs. where it is located 137

Interconnectedness within and between the two pathways 142

The Organization and Functions of the Ventral Visual Processing Stream 144

Hand cells, face cells, and grandmother cells 144

Broader implications of visual properties of temporal cortex neurons 149

A hierarchy of stimulus representation 150

Object-based (viewpoint-independent) vs. image-based (viewpoint-dependent) representation in IT 153

A critical role for feedback in the ventral visual processing stream 153

Taking Stock 158

End-of-Chapter Questions 158

References 159

Other Sources Used 159

Further Reading 160

7 Spatial Cognition and Attention 161

Key Themes 161

Unilateral Neglect: A Fertile Source of Models of Spatial Cognition and Attention 163

Unilateral neglect: a clinicoanatomical primer 163

Hypotheses arising from clinical observations of neglect 164

The Functional Anatomy of the Dorsal Stream 166

Coordinate transformations to guide action with perception 169

From Parietal Space to Medial-Temporal Place 172

Place cells in the hippocampus 173

How does place come to be represented in the hippocampus? 175

The Neurophysiology of Sensory Attention 175

A day at the circus 176

Attending to locations vs. attending to objects 176

Mechanisms of spatial attention 180

Effects of attention on neuronal activity 181

Turning Our Attention to the Future 185

End-of-Chapter Questions 185

References 186

Other Sources Used 186

Further Reading 187

8 Skeletomotor Control 188

Key Themes 188

The Organization of the Motor System 190

The anatomy of the motor system 190

The corticospinal tract 190

The cortico-cerebellar circuit 190

The cortico-basal ganglia-thalamic circuits 192

Functional Principles of Motor Control 193

The biomechanics of motor control 193

Motor cortex 196

The neurophysiology of movement 196

Motor Control Outside of Motor Cortex 202

Parietal cortex: guiding how we move 202

A neurological dissociation between perceiving objects and acting on them 203

Cerebellum: motor learning, balance, . . . and mental representation? 204

Synaptic plasticity 205

Basal ganglia 206

Cognitive Functions of the Motor System 211

Mirror neurons 212

Holding a mirror up to nature? 213

It’s All About Action 214

End-of-Chapter Questions 214

References 215

Other Sources Used 215

Further Reading 216

9 Oculomotor Control and the Control of Attention 218

Key Themes 218

Attention and Action 220

Whys and Hows of Eye Movements 220

Three categories of eye movements 220

The Organization of the Oculomotor System 221

An overview of the circuitry 221

The superior colliculus 222

The posterior system 222

The frontal eye field 223

The supplementary eye field 223

The Control of Eye Movements, and of Attention, In Humans 224

Human oculomotor control 224

Human attentional control 226

The Control of Attention via the Oculomotor System 227

Covert attention 227

Where’s the attentional controller? 230

Are Oculomotor Control and Attentional Control Really the “Same Thing”? 233

The “method of visual inspection” 234

“Prioritized maps of space in human frontoparietal cortex” 235

Of Labels and Mechanisms 238

End-of-Chapter Questions 238

References 238

Other Sources Used 239

Further Reading 240

Section III: Mental Representation 241

10 Visual Object Recognition and Knowledge 243

Key Themes 243

Visual Agnosia 245

Apperceptive agnosia 245

Associative agnosia 245

Computational Models of Visual Object Recognition 247

Two neuropsychological traditions 247

The cognitive neuroscience revolution in visual cognition 249

Category Specificity in the Ventral Stream? 249

Are faces special? 249

Perceptual expertise 251

Evidence for a high degree of specificity for many categories in ventral occipitotemporal cortex 252

Evidence for highly distributed category representation in ventral occipitotemporal cortex 253

Demonstrating necessity 256

The code for facial identity in the primate brain (!?!) 258

Visual Perception as Predictive Coding 261

Playing 20 Questions With the Brain 262

End-of-Chapter Questions 264

References 264

Other Sources Used 265

Further Reading 265

11 Neural Bases of Memory 267

Key Themes 267

Plasticity, Learning, and Memory 269

The Case of H.M. 269

Bilateral medial temporal lobectomy 269

Hippocampus vs. MTL? 272

Association Through Synaptic Modification 273

Long-term potentiation 273

The necessity of NMDA channels for LTM formation 277

How Might the Hippocampus Work? 277

Fast-encoding hippocampus vs. slow-encoding cortex 278

Episodic memory for sequences 279

Episodic memory as an evolutionary elaboration of navigational processing 282

What Are the Cognitive Functions of the Hippocampus? 283

Standard anatomical model 283

Challenges to the standard anatomical model 283

Consolidation 285

Reconsolidation 286

To Consolidate 286

End-of-Chapter Questions 288

References 288

Other Sources Used 289

Further Reading 290

12 Declarative Long-Term Memory 291

Key Themes 291

The Cognitive Neuroscience of LTM 293

Encoding 293

Neuroimaging the hippocampus 293

Incidental encoding into LTM during a short-term memory task 296

The Hippocampus in Spatial Memory Experts 299

Retrieval 299

Retrieval without awareness 300

Documenting contextual reinstatement in the brain 301

Familiarity vs. recollection 303

Knowledge 306

End-of-Chapter Questions 306

References 307

Other Sources Used 308

Further Reading 308

13 Semantic Long-Term Memory 310

Key Themes 310

Knowledge in the Brain 312

Definitions and Basic Facts 312

Category-Specific Deficits Following Brain Damage 313

Animacy, or function? 313

A PDP model of modality specificity 314

The domain-specific knowledge hypothesis 314

How definitive is a single case study? A double dissociation? 315

The Neuroimaging of Knowledge 316

The meaning, and processing, of words 316

An aside about the role of language in semantics and the study of semantics 316

PET scanning of object knowledge 317

Knowledge retrieval or lexical access? 318

Repetition effects and fMRI adaptation 319

The Progressive Loss of Knowledge 321

Primary Progressive Aphasia or Semantic Dementia, Nonverbal deficits in fluent primary progressive aphasia? 322

The locus of damage in fluent primary progressive aphasia? 322

Distal effects of neurodegeneration 324

Entente cordiale 324

Nuance and Challenges 326

End-of-Chapter Questions 326

References 327

Other Sources Used 328

Further Reading 329

14 Working Memory 330

Key Themes 330

“Prolonged Perception” Or “Activated LTM?” 332

Definitions 332

Working Memory and the PFC? The Roots of a Long and Fraught Association 333

Early focus on role of PFC in the control of STM 334

Single-unit delay-period activity in PFC and thalamus 335

Working Memory Capacity and Contralateral Delay Activity 342

The electrophysiology of visual working memory capacity 343

Novel Insights From Multivariate Data Analysis 349

The tradition of univariate analyses 349

MVPA of fMRI 349

Retrospective MVPA of single-unit extracellular recordings 356

Activity? Who Needs Activity? 357

Four-Score and a Handful of Years (and Counting) 360

End-of-Chapter Questions 360

References 360

Other Sources Used 362

Further Reading 362

Section IV: High-Level Cognition 363

15 Cognitive Control 365

Key Themes 365

The Lateral Frontal-Lobe Syndrome 367

Environmental-dependency syndrome 367

Perseveration 368

Electrophysiology of the frontal-lobe syndrome 370

Integration? 371

Models of Cognitive Control 371

Developmental cognitive neuroscience 371

Generalizing beyond development 374

What makes the PFC special? 375

Influence of the DA reward signal on the functions of PFC 376

Neural Activity Relating to Cognitive Control 378

Error monitoring 378

Going Meta 386

Where is the controller? 388

End-of-Chapter Questions 389

References 389

Other Sources Used 390

Further Reading 391

16 Decision Making 392

Key Themes 392

Between Perception and Action 394

Perceptual Decision Making 394

Judging the direction of motion 394

LIP 396

Modeling perceptual decision making 396

Controversy and complications 399

Perceptual decision making in humans 401

Value-Based Decision Making 402

The influence of expected value on activity in LIP 403

Common currency in the omPFC 404

Has neuroeconomics taught us anything about the economics of decision making? 409

Foraging 410

Boys being boys 411

Peer pressure 411

Next Stop 412

End-of-Chapter Questions 412

References 412

Other Sources Used 413

Further Reading 414

17 Social Behavior 415

Key Themes 415

Trustworthiness: A Preamble 417

Delaying gratification: a social influence on a “frontal” class of behaviors 417

The Role of vmPFC in the Control of Social Cognition 418

Phineas Gage 418

Contemporary behavioral neurology 420

Theory of Mind 422

The ToM network 422

The temporoparietal junction (TPJ) 423

False beliefs (?) about Rebecca Saxe’s mind 425

A final assessment of the role of RTPJ in ToM mentalization 429

Observational Learning 430

Predicting the outcome of someone else’s actions 430

Trustworthiness, Revisited 435

End-of-Chapter Questions 435

References 436

Other Sources Used 437

Further Reading 437

18 Emotion 438

Key Themes 438

What is an Emotion? 440

Approach/withdrawal 440

From “feeling words” to neural systems 440

At the nexus of perception and social cognition 440

Trustworthiness Revisited – Again 440

A role for the amygdala in the processing of trustworthiness 441

Implicit information processing by the amygdala 443

The Amygdala 444

Klüver–Bucy syndrome 444

Pavlovian fear conditioning 444

Emotional content in declarative memories 446

The amygdala’s influence on other brain systems 449

The Control of Emotions 450

Extinction 450

How Does That Make You Feel? 455

End-of-Chapter Questions 457

References 458

Other Sources Used 458

Further Reading 459

19 Language 460

Key Themes 460

A System of Remarkable Complexity 462

Wernicke–Lichtheim: The Classical Core Language Network 462

The aphasias 462

The functional relevance of the connectivity of the network 463

Speech Perception 464

Segregation of the speech signal 464

Dual routes for speech processing 468

Grammar 469

Genetics 469

Rules in the brain? 471

Broca’s area 472

The electrophysiology of grammar 475

Speech Production 477

A psycholinguistic model of production 477

Forward models for the control of production 477

Prediction 479

Integration 480

End-of-Chapter Questions 481

References 481

Other Sources Used 483

Further Reading 483

20 Consciousness 485

Key Themes 485

The Most Complex Object in the Universe 487

Different Approaches to the Problem 487

The Physiology of Consciousness 488

Neurological syndromes 488

Sleep 492

Anesthesia 494

Summary across physiological studies 495

Brain Functions Supporting Conscious Perception 495

Are we conscious of activity in early sensory cortex? 497

Manipulating extrinsic factors to study conscious vs. unconscious vision 500

Are Attention and Awareness the Same Thing? 501

Theories of Consciousness 503

Global Workspace Theory 503

Recurrent Processing Theory 505

Integrated Information Theory 506

Updating the Consciousness Graph 508

End-of-Chapter Questions 509

References 509

Other Sources Used 511

Further Reading 511

Glossary G-1

Index I-1