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Biopsychology
1. What categories of research methods are used to study human brain function? What are some examples, advantages, and disadvantages of each?
2. What are three key functions that the brain has evolved to perform as a biological organ?
3. Describe three common forms of cognitive bias in human thinking. Can you think of an example of each type in everyday life?
4. What are the key elements of the scientific method? How might each of these elements help to overcome some of the brain’s built-in biases?
Brain and Behavior A Cognitive Neuroscience Perspective.pdf
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A Cognitive Neuroscience Perspective
Brain and Behavior
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DAVID EAGLEMAN Baylor College of Medicine, Department of Neuroscience Director, Initiative on Neuroscience and Law
JONATHAN DOWNAR Department of Psychiatry and Institute of Medical Science, University of Toronto Toronto Western Hospital, University Health Network
New York Oxford Oxford University Press
A Cognitive Neuroscience Perspective
Brain and Behavior
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Library of Congress Cataloging-in-Publication Data Eagleman, David. Brain and behavior : a cognitive neuroscience perspective / David Eagleman, Baylor College of Medicine, Department of Neuroscience, Director, Initiative on Neuroscience and Law, Jonathan Downar, Department of Psychiatry and Institute of Medical Science, University of Toronto, Toronto Western Hospital, University Health Network. pages cm Includes bibliographical references and index. ISBN 978-0-19-537768-2 1. Cognitive neuroscience. 2. Neuropsychiatry. I. Downar, Jonathan. II. Title. QP360.5.E24 2016 612.8'233—dc23
2015013925
Printing number: 9 8 7 6 5 4 3 2 1
Printed in the United States of America on acid-free paper
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DEDICATION
From David: To Cirel, Arthur, Read, Francis, and Sarah, in the order
that I met you.
From Jonathan: In loving memory of Ann Downar—devoted mother of
two sons, and keen-eyed editor of many a manuscript—a gifted wielder of the red pen, from
whom no detail was too small to escape notice. With gratitude that you were able to see this book take shape
in its early days and with the hope that its final form would have made you proud.
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BRIEF CONTENTS PREFACE xv ABOUT THE AUTHORS xxi
PART I THE BASICS CHAPTER 1 Introduction 2 CHAPTER 2 The Brain and Nervous System 36 CHAPTER 3 Neurons and Synapses 74 CHAPTER 4 Neuroplasticity 102
PART II HOW THE BRAIN INTERACTS WITH THE WORLD CHAPTER 5 Vision 130 CHAPTER 6 Other Senses 162 CHAPTER 7 The Motor System 196
PART III HIGHER LEVELS OF INTERACTION CHAPTER 8 Attention and Consciousness 232 CHAPTER 9 Memory 270 CHAPTER 10 Sleep 308 CHAPTER 11 Language and Lateralization 336
PART IV MOTIVATED BEHAVIORS CHAPTER 12 Decision Making 362 CHAPTER 13 Emotions 398 CHAPTER 14 Motivation and Reward 438 CHAPTER 15 Social Cognition 472
PART V DISORDERS OF BRAIN AND BEHAVIOR CHAPTER 16 Neurological and Psychiatric Disorders 514
GLOSSARY 556 REFERENCES 585 CREDITS 637 NAME INDEX 643 SUBJECT INDEX 657
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The Payoffs of Cognitive Neuroscience 28 Healing the Disordered Brain 29 Enhancing Human Abilities 30 Blueprints for Artificial Cognition 31 Brain-Compatible Social Policies 31
Conclusion 33 Key Principles 33 Key Terms 34 Review Questions 34 Critical-Thinking Questions 35
CHAPTER 2 The Brain and Nervous System 36
LEARNING OBJECTIVES 36 STARTING OUT: The Brains of Creatures Great and
Small 38 An Overview of the Nervous System 39
Why Put Your Neurons in a Brain at All? 39 The Common Features of Every Central Nervous System 40 Getting Oriented in the Brain 42
The Peripheral Nervous System 43 Separate Systems for the Inner and Outer Environments 43 A Nervous System with Segmental Organization 44
The Spinal Cord 47 Circuits within a Segment: Spinal Reflexes 47
Case Study: Christopher Reeve, 1952–2004 47
Complex Circuits across Segments: Central Pattern Generators 49
The Bigger Picture: In Search of a Cure for Spinal Cord Injury 50
The Brainstem 51 Medulla Oblongata and Pons 51
Neuroscience of Everyday Life: Why Do We Get the Hiccups? 53
Midbrain 53 Most Cranial Nerves Emerge from the Brainstem 54
The Cerebellum 54 Circuitry of the “Little Brain” 56 Functions of the Little Brain 56
The Diencephalon: Hypothalamus and Thalamus 57 Hypothalamus: A Keystone Structure in Homeostasis 57 Thalamus 59
Case Study: Waking the Brain 61
PREFACE xv ABOUT THE AUTHORS xxi
PART I THE BASICS CHAPTER 1 Introduction 2
LEARNING OBJECTIVES 2 STARTING OUT: A Spark of Awe in the Darkness 4
Who Are We? 5 The Mission of Cognitive Neuroscience 5 Neuroscience Is a Relatively New Field 6
In Pursuit of Principles 6 The Functions behind the Form 6 Which Parts Matter? 7 What Is the Brain For? 8
How We Know What We Know 9 Connectional Methods 10 Correlational Methods 11
Research Methods: Magnetic Resonance Imaging 12
Lesion Methods 13 Stimulation Methods 14 A Toolbox of Complementary Methods 16
Thinking Critically about the Brain 16 Is the Brain Equipped to Understand Itself? 16 Biases and Pitfalls in Human Cognition 17 A Toolbox of Critical-Thinking Techniques 18
The Big Questions in Cognitive Neuroscience 19 Why Have a Brain at All? (Chapter 2) 19 How Is Information Coded in Neural Activity? (Chapter 3) 20 How Does the Brain Balance Stability against Change? (Chapter 4) 20 Why Does Vision Have So Little to Do with the Eyes? (Chapter 5) 21 How Does the Brain Stitch Together a Picture of the World from Different Senses? (Chapter 6) 21 How Does the Brain Control Our Actions? (Chapter 7) 22 What Is Consciousness? (Chapter 8) 22 How Are Memories Stored and Retrieved? (Chapter 9) 23 Why Do Brains Sleep and Dream? (Chapter 10) 24 How Does the Human Brain Acquire Its Unique Ability for Language? (Chapter 11) 24 How Do We Make Decisions? (Chapter 12) 25 What Are Emotions? (Chapter 13) 25 How Do We Set Our Priorities? (Chapter 14) 27 How Do I Know What You’re Thinking? (Chapter 15) 27 What Causes Disorders of the Mind and the Brain? (Chapter 16) 28
CONTENTS
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Key Principles 100 Key Terms 100 Review Questions 101 Critical-Thinking Questions 101
CHAPTER 4 Neuroplasticity 102 LEARNING OBJECTIVES 102 STARTING OUT: The Child with Half a Brain 104
The Brain Dynamically Reorganizes to Match Its Inputs 105 Changes to the Body Plan 105
Case Study: Phantom Sensation 106 Research Methods: Mapping Out the Brain 107
Changes to Sensory Input 107
The Brain Distributes Resources Based on Relevance 109 The Role of Behavior 109 The Role of Relevance: Gating Plasticity with Neuromodulation 109
Neuroscience of Everyday Life: Pianists and Violinists Have Different Brains 110
Case Study: The Government Worker with the Missing Brain 112
The Brain Uses the Available Tissue 113 Maps Adjust Themselves to the Available Brain Tissue 113 Cortical Reorganization after Brain Damage 114
A Sensitive Period for Plastic Changes 114 A Window of Time to Make Changes 114
Case Study: Danielle, the Feral Child in the Window 115
The Sensitive Period in Language 116 Neuromodulation in Young Brains 116
Hardwiring versus World Experience 117 Aspects of the Brain Are Preprogrammed 117 Experience Changes the Brain 117 Brains Rely on Experience to Unpack Their Programs Correctly 119
The Mechanisms of Reorganization 120 Neurons Compete for Limited Space 120 Competition for Neurotrophins 121 Rapid Changes: Unmasking Existing Connections 122 Slow Changes: Growth of New Connections 123
Changing the Input Channels 123 Case Study: The Man Who Climbs with His Tongue 125 The Bigger Picture: Adding New Peripherals 126
Conclusion 126 Key Principles 127 Key Terms 127 Review Questions 128 Critical-Thinking Questions 128
The Telencephalon: Cerebral Cortex and Basal Ganglia 61 Cerebral Cortex 61 Basal Ganglia 64
Research Methods: Cytoarchitecture of the Cortex 64
Uniting the Inside and Outside Worlds 66 The Limbic System 66 The Ventricular System and Brain Function 68
Conclusion 69 Key Principles 71 Key Terms 71 Review Questions 72 Critical-Thinking Questions 73
CHAPTER 3 Neurons and Synapses 74 LEARNING OBJECTIVES 74 STARTING OUT: The Kabuki Actor and the Pufferfish 76
The Cells of the Brain 77 Neurons: A Close-Up View 77 Many Different Types of Neurons 79 Glial Cells 80
Research Methods: Visualizing Neurons and Their Products 81
Synaptic Transmission: Chemical Signaling in the Brain 83 Release of Neurotransmitter at the Synapse 83 Types of Neurotransmitters 84 Receptors 85 Postsynaptic Potentials 86
The Bigger Picture: Psychoactive Drugs 87
Spikes: Electrical Signaling in the Brain 88 Adding Up the Signals 88 How an Action Potential Travels 89 Myelinating Axons to Make the Action Potential Travel Faster 90 Action Potentials Reach the Terminals and Cause Neurotransmitter Release 91
Case Study: Multiple Sclerosis 91 Neuroscience of Everyday Life: The Magic of a Local
Anesthetic 92
What Do Spikes Mean? The Neural Code 93 Encoding Stimuli in Spikes 93 Decoding Spikes 95
Research Methods: Recording Action Potentials with Electrodes 96
Individuals and Populations 97 Populations of Neurons 97 Forming a Coalition: What Constitutes a Group? 98 Open Questions for Future Investigation 99
Conclusion 100
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CHAPTER 6 Other Senses 162 LEARNING OBJECTIVES 162 STARTING OUT: The Man with the Bionic Ear 164
Detecting Data from the World 165 Hearing 165
Research Methods: Psychophysics 166
The Outer and Middle Ear 167 Converting Mechanical Information into Electrical Signals: The Inner Ear 168
Neuroscience of Everyday Life: The Undetectable Cell Phone 169
The Auditory Nerve and Primary Auditory Cortex 170 The Hierarchy of Sound Processing 171 Sound Localization 172 Balance 173
The Somatosensory System 174 Touch 174 Temperature 175 Pain 175
Case Study: The Pain of a Painless Existence 176
Proprioception 177 Interoception 177 The Somatosensory Pathway 179
Chemical Senses 180 Taste 180 Smell 183 The Sense of Flavor 184 Pheromones 185
The Brain Is Multisensory 185 Synesthesia 186 Combining Sensory Information 186 The Binding Problem 188 The Internal Model of the World 189
Case Study: The Paralyzed Supreme Court Justice Who Claimed He Could Play Football 189
Time Perception 190 Conclusion 193 Key Principles 193 Key Terms 194 Review Questions 194 Critical-Thinking Questions 195
CHAPTER 7 The Motor System 196 LEARNING OBJECTIVES 196 STARTING OUT: “‘Locked-In Syndrome”’ 198
Muscles 199 Skeletal Muscle: Structure and Function 199 The Neuromuscular Junction 200
PART II HOW THE BRAIN INTERACTS WITH THE WORLD
CHAPTER 5 Vision 130 LEARNING OBJECTIVES 130 STARTING OUT: Vision Is More Than the Eyes 132
Visual Perception 132 What Is It Like to See? 132 Signal Transduction 133
Anatomy of the Visual System 134 Sensory Transduction: The Eye and Its Retina 134
Case Study: The Bionic Retina 137
Path to the Visual Cortex: The Lateral Geniculate Nucleus 139 The Visual Cortex 139 Two Eyes Are Better Than One: Stereo Vision 141
Neuroscience of Everyday Life: Random-Dot Stereograms 142
Higher Visual Areas 142 Secondary and Tertiary Visual Cortex: Processing Becomes More Complex 142 Ventral Stream: What an Object Is 143
The Bigger Picture: Reading the Movies in Our Minds 145
Dorsal Stream: How to Interact with the World 146 Case Study: The World in Snapshots 147
Attention and the Dorsal Stream 148 Comparing the Ventral and Dorsal Processing Streams 149 The Bigger Picture of the Visual Brain 150
Case Study: The Blind Woman Who Could See, Sort Of 150
Perception Is Active, Not Passive 151 Interrogating the Scene with Our Eyes 151 The Blind Spot 152 Seeing the Same Object Different Ways: Multistability 152 Binocular Rivalry: Different Images in the Two Eyes 152 We Don’t See Most of What Hits Our Eyes: Fetching Information on a Need-to-Know Basis 153
Vision Relies on Expectations 154 Change Blindness 154 Saving Resources by Embedding Prior Experience 155 Unconscious Inference 156 Activity from Within 157 Feedback Allows an Internal Model 157
Conclusion 158 Key Principles 159 Key Terms 160 Review Questions 160 Critical-Thinking Questions 161
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Inattentional Blindness 236 Neuroscience of Everyday Life: Stage Magic 237
Approaches to Studying Attention and Awareness 238 Attentional Orienting Paradigms: Aiming the “Spotlight” of Attention 238 The Oddball Paradigm: Monitoring a Physiological Measure of Attention 239 Uncoupling Sensory Input from Perception: Sensory Rivalry 240
Neural Mechanisms of Attention and Awareness 241 Seeking the Correlates of Consciousness 241 Hemineglect: A Disorder of Attention and Awareness 242
Case Study: Unaware of Half of the World 243
Neural Correlates of Attention: A Single Network or Many? 245
Case Study: Whose Arm Is This, Anyway? 247
Sites of Attentional Modulation: Neurons and Neural Populations 247 The Biased-Competition Model of Attention 247 Attention and Single Neurons: Enhancing the Signal 248 Attention and Local Groups of Neurons 250
Synchronization, Attention, and Awareness 250 Coma and Vegetative State: Anatomy of the Conscious State 253 Why Should Synchronization Matter? 253 Unconsciousness: Coma and Vegetative State 254
Case Study: Waking the Brain 255
Midbrain and Thalamus: Key Players in the Conscious State 256
Anesthesia and Sleep: Rhythms of Consciousness 258 Sleep: Unraveling the Rhythm of Consciousness 258 Anesthesia: Reversible, Artificial Unconsciousness 260
Theories of Consciousness 262 Dualism: The Mind–Body Problem 262 Functionalist Theories of Consciousness 263 Consciousness and the Integration of Information 265
Conclusion 266 Key Principles 267 Key Terms 267 Review Questions 268 Critical-Thinking Questions 269
CHAPTER 9 Memory 270 LEARNING OBJECTIVES 270 STARTING OUT: The Woman Who Cannot Forget 272
The Many Kinds of Memory 273 Working and Long-Term Memory 273
The Spinal Cord 201 Lower Motor Neurons 201 Spinal Motor Circuits: Reflexes 202 Spinal Motor Circuits: Central Pattern Generators 203 Descending Pathways of Motor Control 204
The Cerebellum 205 The Circuitry of the Cerebellum 206 Motor Functions of the Cerebellum 208 Nonmotor Functions of the Cerebellum 208
The Motor Cortex 209 Motor Cortex: Neural Coding of Movements 211 Motor Cortex: Recent Controversies 212
The Bigger Picture: Neural Implants for Motor Control 214
The Prefrontal Cortex: Goals to Strategies to Tactics to Actions 215 The Functional Organization of the Prefrontal Cortex in Motor Control 215 Sensory Feedback 216 Mirror Neurons in Premotor Cortex 217 Control Stages of the Motor Hierarchy 218
Basal Ganglia 219 Components of the Basal Ganglia 219 Circuitry of the Basal Ganglia 220 Diseases of the Basal Ganglia 220
Medial and Lateral Motor Systems: Internally and Externally Guided Movement Control 222 Organization of Medial Motor Areas 222 Functions of Medial and Lateral Motor Systems 222
Neuroscience of Everyday Life: Why Can’t I Multitask? 225
Did I Really Do That? The Neuroscience of Free Will 226
Research Methods: Neurosurgical Stimulation 228 Case Study: Alien Hand Syndrome 229
Conclusion 229 Key Principles 230 Key Terms 230 Review Questions 231 Critical-Thinking Questions 231
PART III HIGHER LEVELS OF INTERACTION
CHAPTER 8 Attention and Consciousness 232
LEARNING OBJECTIVES 232 STARTING OUT: The Stream of Consciousness 234
Awareness Requires Attention 235 Change Blindness 235
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The Limitations of Neural Networks 302 Neural Networks: Solving the Wrong Problem? 303 Remembering Relationships, Not Features 303 The Future of Memory Research 303
Conclusion 304 Key Principles 305 Key Terms 305 Review Questions 306 Critical-Thinking Questions 307
CHAPTER 10 Sleep 308 LEARNING OBJECTIVES 308 STARTING OUT: Caught between Sleeping and
Waking 310 Sleep and the Brain 311
The Brain Is Active during Sleep 311 Research Methods: Electroencephalography 312
The Neural Networks of Sleep 314 The Brain during REM Sleep 314
The Circadian Rhythm 316 Entrainment of the Circadian Rhythm by Light Cues 316 The Circadian Rhythm Is Not Fixed 317
Case Study: The Shifted Circadian Rhythm 318
The Circadian Rhythm and Napping 319 The Bigger Picture: Schools and Circadian Rhythms 320
Why Do Brains Sleep? 320 Four Theories of Sleeping: Restoration, Survival, Simulation, Learning 320 Rehearsal 321 Forgetting 323 Insight and the Restructuring of Information 324
Dreaming 324 Dream Content 325
Neuroscience of Everyday Life: Lucid Dreaming 326
Can Dreams Shed Light on Consciousness? 327 Dreams of the Future and How to Study Them 327
Sleep Deprivation and Sleep Disorders 328 Sleep Deprivation 328
Case Study: Staying Awake 329
Insomnia 330 Hypersomnia 331
Case Study: The Family Who Couldn’t Sleep 332
Parasomnias 333 Conclusion 333 Key Principles 334 Key Terms 335 Review Questions 335 Critical-Thinking Questions 335
Implicit Memory 274 Explicit Memory 275
Travels in Space and Time: The Hippocampus and Temporal Lobe 277
Case Study: Gone but Not Forgotten: Henry Molaison, 1926–2008 277
A Map of the Medial Temporal Lobe 278 Episodic Memory 279 Spatial Memory 280 Theories of Hippocampal Function 281 Unifying the Functions of the Hippocampus 282
Remembering the Future: Prospection and Imagination 282 How We Imagine Future Experiences 282
Research Methods: Localizing Human Brain Function 283
The Circuitry of Prospection and Recollection 284 Neuroscience of Everyday Life: Simonides and the
Champions of Memory 285
Prospection in Other Species 286 Models of Prospection 287
The Confabulation of Reality 288 Confabulation in the Injured Brain 288
Case Study: The Woman with a Thirty-Year-Old Baby 288
The Anatomy of Spontaneous Confabulation 289 Confabulation in the Normal Brain 290 The Anatomy of a False Memory 291
The Bigger Picture: Scanning for the Truth 292
The Mechanisms of Memory 292 General Mechanisms of Learning and Memory 292 Memory as Synaptic Change 293 Long-Term Potentiation and Depression of Synaptic Connections 293 The NMDA Receptor 293 Consolidation and Reconsolidation 294 Associative Neural Networks 295
Beyond Synaptic Plasticity: The Frontiers of Memory Mechanisms 296 Whole Neurons as a Substrate for Memory? 296 New Neurons for New Memories 297 Spines: Another Structural Basis for Memory? 298 Looking inside the Cell: Memory in Chemical Reactions 298
Case Study: The Flies with Photographic Memory 299
Epigenetics: Making a Single Genome Play Different Tunes 300
The Mysteries of Memory 300 Are the Roles of LTP and LTD Overstated? 301 The Timing of Spikes 301
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Where Do Our Irrational Decisions Come From? 371 Decision Making in Other Species 372 Do Irrational Decisions Come from Irrational People? 373 One Brain, Two Systems 373
How the Brain Decides 374 The Neural Mechanisms of Delay Discounting 374 Neural Mechanisms of Decisions under Risk 375 The Neural Basis of the Endowment Effect 377 The Neural Basis of the Framing Effect 378
The Common Currency of Subjective Value 379 Comparing Apples to Oranges 379
Research Methods: Charting the Landscape of Subjective Value 380
A Consistent Neural Basis for Subjective Value 380 Evaluation and the Orbitofrontal Cortex 381 One Currency, but Many Markets 382
Neuroscience of Everyday Life: Snack Food or Brussels Sprouts? 383
A Hierarchy of Internally Guided Decision Making 384 Internally and Externally Guided Decision Making 384 Values into Goals 385 Goals into Plans 387 Plans into Behavior and Action 388
Modulators of Decision Making 389 Strategic Use of Decision-Making Systems 389 Neurotransmitter Effects on Decision Making 391
The Bigger Picture: How to Avoid the Scorpion’s Sting 394
Conclusion 395 Key Principles 396 Key Terms 396 Review Questions 397 Critical-Thinking Questions 397
CHAPTER 13 Emotions 398 LEARNING OBJECTIVES 398 STARTING OUT: Sadness, at the Flip of a Switch 400
Early Theories of Emotion 401 Emotional Expressions: Signposts on a Landscape of Inner States 401 The James–Lange Theory of Emotion: A Bottom-Up Theory 402 The Cannon–Bard Theory: A Top-Down Theory 404
Case Study: Pathological Laughter and Crying 405
Two-Factor Theories: Reconciling Central and Peripheral Influences on Emotion 406
CHAPTER 11 Language and Lateralization 336
LEARNING OBJECTIVES 336 STARTING OUT: The Stuttering King 338
Speech, Language, and Communication 338 Aphasia: The Loss of Language 339
Case Study: The Woman Who Couldn’t Find Her Words 340
Broca’s Aphasia 341 Wernicke’s Aphasia 341
Case Study: The Woman Who Makes Up Words 342
A Language Network 343 The Larger Picture of Language-Specific Regions 344 Dyslexia 347 Stuttering 348
Lateralization: The Two Hemispheres Are Not Identical 348 Tests for Dominance 349 Apraxia 349 Hemispheric Differences 349 Two Brains in One? The Case of the Split-Brain Patients 350 Thinking about Cerebral Asymmetry 352
Development of Language 352 Learning Language from Experience 353 Innate Language Tendencies 354 Socially and Emotionally Directed Learning 356
Research Methods: The Baby with No Privacy 357
Conclusion 359 Key Principles 359 Key Terms 360 Review Questions 360 Critical-Thinking Questions 361
PART IV MOTIVATED BEHAVIORS CHAPTER 12 Decision Making 362
LEARNING OBJECTIVES 362 STARTING OUT: A Fatal Mistake, at the Highest
Place on Earth 364 How Do We Decide What to Do? 365
The Scorpion and the Frog 365 The Search for a “Physics” of Human Decisions 366 Homo economicus and Rational Choice Theory 366
The Predictably Irrational Homo sapiens 367 Homo sapiens versus Homo economicus 367 Confused by Uncertainty 368 The Framing Effect and the Endowment Effect 369 The Illusory Value of Procrastination 370
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Midbrain Dopamine Neurons and the Common Currency of Motivation 447
Reward, Learning, and the Brain 447 Defining Reward 448 Learning from Reward Using Prediction Error 448 “Liking” Is Different from “Wanting” 450
Opioids and the Sensation of Pleasure 451 Opioids, Opioid Receptors, and Opioid Functions 451 Opioids and Reward 453
Dopamine, Learning, Motivation, and Reward 453 Dopamine Functions in Motivation and Reward 454 Unifying the Functions of Dopamine 455
Research Methods: Measuring Neurotransmitter Levels in the Brain 456
Neurotransmitters Are Messengers, Not Functions 457
Addiction: Pathological Learning and Motivation 457 Addictive Substances Have Distorted Reward Value 457
Neuroscience of Everyday Life: The Pursuit of Happiness 458
Addiction Is a Result of Pathological Learning 460 The Circuitry and Chemistry of Addiction 461
Unlearning Addiction 462 The Challenge of Treatment 462
Case Study: Pathological Gambling in a Patient with Parkinson’s Disease 463
Existing Approaches to Treatment 464 Future Approaches to Treatment 465
The Bigger Picture: Finding the Motivation to Change 468
Conclusion 469 Key Principles 469 Key Terms 470 Review Questions 471 Critical-Thinking Questions 471
CHAPTER 15 Social Cognition 472 LEARNING OBJECTIVES 472 STARTING OUT: Why Risk Your Life for a Yellow
T-shirt? 474 Social Perception 475
What’s in a Face? 475 Do I Look Like a Liar to You? 476
Neuroscience of Everyday Life: A Poker Face 476
Social Knowledge and the Temporal Pole 478 Social Signals and the Superior Temporal Sulcus 479
Social Thinking: Theory of Mind 480 What Is Theory of Mind? 480 Neural Mechanisms of Theory of Mind 481
Core Limbic Structures: Amygdala and Hypothalamus 408 Hypothalamus: Internal States, Homeostatic Drives 409
Case Study: An Internal Growth of Rage 411
Do Hypothalamic Circuits Generate Inner Emotional Experiences? 412 Amygdala: Externally Generated States and Drives 412 The Amygdala and Emotional Experience 413
Case Study: The Woman Who Knows No Fear 414
Hippocampus: Emotional Memories 416 Ventral Striatum: Pleasure and Reward 417 Bringing It All Together: The Circuit of Papez and the Ring of Limbic Cortex 418
The Bigger Picture: The Ethics of Brain Stimulation in Human Beings 419
The Limbic Cortex and Emotions 420 The Interoceptive Insula: The “Feeling” Side of Emotions 420 Cingulate Cortex: A Motor Cortex for the Limbic System 421
Neuroscience of Everyday Life: Mental Effort 422
Ventromedial Prefrontal Cortex: A Generator of Gut Feelings 423
Limbic Association Cortex: Modulation of Emotion 425 The Mechanisms of Emotional Reappraisal 425 Brain Injury, Brain Stimulation, and Emotion Regulation 426
Neurochemical Influences on Emotion 428 Case Study: A Cure Born of Desperation 429
Serotonin and Mood 430 Norepinephrine and Mood 431 GABA and Anxiety 432
Conclusion 433 Key Principles 435 Key Terms 435 Review Questions 436 Critical-Thinking Questions 437
CHAPTER 14 Motivation and Reward 438 LEARNING OBJECTIVES 438 STARTING OUT: “More Important Than Survival
Itself ” 440 Motivation and Survival 440
Addiction: An Illness of Motivation 440 Why Motivation Matters 441 Feelings: The Sensory Side of Motivation 442
The Circuitry of Motivation: Basic Drives 443 Hypothalamus and Homeostatic Drives 443 Amygdala and External-World Drives 445
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Huntington’s Disease: A Genetic Rarity, in Two Senses 524
Tourette Syndrome: A Case of Involuntary Volition? 527
Obsessive–Compulsive Disorder: Neurological or Psychiatric? 529
Research Methods: Voxel-Based Morphometry 532
Schizophrenia: A Dementia of the Young 534 Bipolar Disorder 540 Depression: A Global Burden 544
Impact of Depression 544 Case Study: A Lifetime Studying, and Living with, Bipolar
Disorder 545
Causes of Depression 546 Neurochemical Effects of Depression on the Brain 548 Functional Effects of Depression on the Brain 549 Treatment of Depression 550
Conclusion 552 Key Principles 553 Key Terms 554 Review Questions 554 Critical-Thinking Questions 555
GLOSSARY 556 REFERENCES 585 CREDITS 637 NAME INDEX 643 SUBJECT INDEX 657
Mirror Neurons and Theory of Mind 483 Disorders of Theory of Mind 484
Social Feelings: Empathy and Its Many Components 487 An Emotional Theory of Mind 487 Empathy, Sympathy, and Compassion 488 Neural Mechanisms of Emotional Mimicry and Contagion 489 Neural Mechanisms of Empathy, Sympathy, and Antipathy 490 Disorders of Empathy 491
Social Emotions, Motivations, and Behavior 493 Social Emotions from Theory of Mind 493
Case Study: Acquired Sociopathy 493
Social Emotions from Social Values 495 Social Reward and Social Aversion 496 The Anatomy of a Lie 498
Neurotransmitters and Social Behavior 499 Research Methods: Transcranial Direct Current
Stimulation 500
An Ancient and Fundamental System 501 Oxytocin 501 Vasopressin 503
The Bigger Picture: The Brave New World of the “Cuddle Hormone”? 504
The Social Self 505 The Wondrous Self-Awareness of the Human Brain 505 Forms of Self-Awareness 505 Why Bother with Self-Awareness? 506 Neural Correlates of Self-Awareness 507 Disorders of Self-Awareness 508 Self-Awareness and Social Cognition 510
Case Study: The Man in the Mirror 510
Conclusion 511 Key Principles 511 Key Terms 512 Review Questions 512 Critical-Thinking Questions 513
PART V DISORDERS OF BRAIN AND BEHAVIOR
CHAPTER 16 Neurological and Psychiatric Disorders 514
LEARNING OBJECTIVES 514 STARTING OUT: Epilepsy: “The Sacred Disease” 516
Alzheimer’s Disease: Burning Out with Age? 517 Frontotemporal Dementia: Like a Cancer of the Soul 521
Case Study: Ravel and “Bolero” 523
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Approach Brain and Behavior is a new kind of textbook for the emerg- ing field of cognitive and behavioral neuroscience. No other textbook tells the story of the brain in such a logical and meaningful way. Through the use of overarching principles rather than lists or facts, Brain and Behavior highlights what we understand about the function of the brain, as well as what we have left to learn and future directions.
Brain and Behavior illustrates current thinking in the field and builds scaffolding for you to learn new concepts. Without compromising important ideas, it covers a wide swath of territory critical for understanding the brain, from the basics of the nervous system to sensory and motor sys- tems, the frontal lobes, sleep, language, memory, drug addic- tion, and brain disorders. Throughout the book, the narrative emphasizes the dynamically changing nature of the brain (neuroplasticity) using clear and vibrant writing and fasci- nating real-life examples and applications.
Brain and Behavior presents the concepts of cognitive neuroscience as thoroughly as possible, using an easy and ac- cessible style that does not presuppose advanced knowledge. It features the following: • A principles-based approach. Students of all ages, and
especially undergraduates, find themselves frustrated with lists of unrelated facts to memorize. Overarching principles enable you to wrap your head around the big picture and learn how to mine for further details.
• A progressive structure. This book unfolds logically, be- ginning with the basics of the nervous system before moving to the brain’s interaction with the world (sensory and motor systems) and then to more complex interac- tions (attention, learning, sleeping, and dreaming). Building on this foundation, the book introduces still more complex interactions (language, decisions, emo- tions, motivation, and reward), before exploring the ways that the system can go awry (drug addiction, mental dis- orders, and neurological disorders).
• Engaging features. Several tools and features throughout each chapter help in the preparation for exams and high- light real-life examples and applications of the material.
Chapter Opening Each chapter opens with a rendering of the human body, emphasizing a key aspect related to the chapter topic, and includes a list of the major sections, features, and learning objectives to be covered.
The human brain is the most complex object we have found in the universe. There are more connections in a cubic milli- meter of neural tissue than there are stars in the Milky Way galaxy. So it is no surprise that even in the glow of remarkable advancement in recent decades, we find ourselves squinting to find the lay of the land. Even for experts in the field, the brain’s complexity can feel daunting at the best of times.
With this point in mind, we set out to write a cognitive neuroscience textbook that would help readers make sense of this complexity by focusing on fundamental scientific principles, patterns, and ways of thinking. Throughout the text, we prize understanding integration of principles over simple memorization of brain structures and scattered find- ings. As students from all backgrounds become increasingly interested in the brain, we wanted to capture the state of the science while distilling the expansive territory into under- standable parts.
Brain and Behavior covers a wide swath of territory criti- cal for understanding the brain, from the basics of the ner- vous system to sensory and motor systems, sleep, language, memory, emotions and motivation, social cognition, and brain disorders. Throughout the narrative we have sought to emphasize the dynamically changing nature of the brain through the mechanisms of neuroplasticity. In addition, wherever possible, we make reference to elements of neuro- science that are encountered in everyday life. We illustrate key points and concepts using case studies of rare but illumi- nating brain disorders. Brain and Behavior pulls together the best of our current knowledge about the brain while ac- knowledging our current areas of ignorance and pointing the reader toward our most promising directions for future research.
Brain and Behavior aims to present key concepts as thor- oughly as possible, in a reader-friendly style that does not presuppose advanced knowledge of the field. Our intention was to make the topic as accessible as possible to a wide un- dergraduate audience. However, it is our hope that students at all levels, and in other fields, will find this text to be a help- ful introductory guide to the complexities of the human brain.
Whether you are reading this book as an aspiring neuro- scientist or whether you are reading it simply as a fellow human being who wishes to better understand the miniature universe we carry inside our heads, we hope that you will come away from Brain and Behavior having gained a better understanding both of the human brain and of the human experience.
With best wishes, David Eagleman and Jonathan Downar
PREFACE
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xvi Preface
Inside the Chapter CASE STUDIES interwoven with the text present fascinat- ing human-interest stories that illustrate key content. These clinical cases include a woman incapable of feeling fear and a blind mountain climber who “sees” via electrical signals on his tongue.
Changing the Input Channels 125
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we think of as a visual cortex; plug an auditory stream into it, and it will become an auditory cortex.
As we have seen so far, the brain has a remarkable capac- ity to reconfigure itself in the face of new inputs, outputs, neural gains, or neural losses. That versatility opens the door to technologies that can deliver information to the brain through unusual sensory channels. For example, what if a blind person used the data stream from a video camera and converted it into sounds in her headphones? Would she eventually be able to see the world by listening? Welcome to the world of sensory substitution experiments, in which a deficient sensory channel is circumvented in favor of other routes to the brain (Lenay, Hanneton, Marque, & Genouel, 2003; Maidenbaum, Abboud, & Amedi, 2013; Poirier, De Volder, & Scheiber, 2007).
Why does the BrainPort use the tongue, of all places? Although we normally think of the tongue as a taste organ, it also has the finest sense of touch on the entire body. It can distinguish stimuli only 1.6 mm apart—roughly twice as fine as a typical fingertip (Wilson, Walton, Tyler, & Wil- liams, 2012). This makes the tongue a great site for passing on new kinds of information, even learning to “see.” A grid of electrodes, the size of a postage stamp, zaps the tongue, converting the lattice of video pixels into “pixels” in the mouth (Bach-y-Rita, 2004; Bach-y-Rita, Collins, Saunders,
White, & Scadden, 1969). With practice, the tongue learns to interpret the signals that correspond to the visual prop- erties, such as how large an object is, how far away it is, and whether it’s moving in a particular direction. With the BrainPort, blind users can learn to navigate complex ob- stacle courses and throw balls into buckets. For sighted people, the BrainPort can be used to see in the dark. How is any of this possible? Vision isn’t about the eyes. It’s about the brain (Khoo, Seidel, & Zhu, 2012). Other sen- sory substitution devices for the blind convert video streams into patterns of touch on the lower back, sound for the ears, or small electric shocks to the skin of the fore- head. Similarly, a sensory substitution device for the deaf uses a vest covered in vibratory motors to translate sound into patterns on the skin (Novich & Eagleman, under review). These amazing substitutions are possible only be- cause the brain can dynamically shape itself around what- ever input is presented. It even seems possible that in the near future people will feed information streams directly into their cortex.
In conclusion, the rerouting of information and the suc- cess of sensory substitution underscores the dynamic plas- ticity of brains. The principles of competition constantly reorganize the circuitry to optimize their representation of the input.
CASE STUDY: The Man Who Climbs with His Tongue Eric Weihenmayer is a mountaineer who has scaled Mount Everest—a feat made even more impressive by the fact that he is blind. As a child, Eric progressively lost his vision to a rare eye disease called retinoschi- sis, and he was rendered entirely blind by the age of 13. But that didn’t slow his ambition to become a climber. Given his condition, it’s cap- tivating to watch Eric scale shear rock faces, holding on to small crev- ices and protrusions. How does he know where to reach next? How does he do it?
Eric climbs with an electrode grid in his mouth called the BrainPort (FIGURE 4.26). The grid delivers little impulses to his tongue that mirror
the visual signals from a camera at- tached to his forehead. Eric reports that he first had to think hard about how the tongue stimulation might translate into edges and shapes. But
he learned, eventually, to recognize the stimulation as direct perception (Levy, 2008). He is now able to use the device for a low-resolution but effec- tive sense of his visual surroundings.
FIGURE 4.26 BrainPort. The BrainPort converts a video feed to corresponding electrical activity on the tongue. With this technology, blind users can come to understand their visual surroundings with high accuracy.
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Three special features appear throughout each chapter to highlight key themes and concepts:
NEUROSCIENCE OF EVERYDAY LIFE explains how neuroscience directly relates to our daily lives, such as why people have difficulty multitasking.
Medial and Lateral Motor Systems: Internally and Externally Guided Movement Control 225
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NEUROSCIENCE OF EVERYDAY LIFE: Why Can’t I Multitask? Why is it so hard to do more than one thing at once? Why should doing one task interfere with doing another? Our mouths and lips are not neces- sary to drive a car, so why is it dan- gerous to talk on a hands-free cell phone while driving? When we look at a face, we don’t need to examine each part of the face one at a time to tell who we are looking at. We simply do it all at once. Why can’t we per- form many behaviors at once, in the same way that we perform many sensory tasks at once?
Neuroimaging studies have begun to search for the apparent “bottleneck” in information pro- cessing that limits our ability to mul- titask (FIGURE 7.28). One study (Dux, Ivanoff, Asplund, & Marois, 2006) had subjects perform two tasks almost simultaneously: pressing buttons according to an auditory cue and pronouncing syllables accord- ing to a visual cue. When these tasks occurred close together in time, the subjects’ reaction times began to slow: the limits of their multitasking abilities had been reached.
Functional MRI revealed a possi- ble basis for the slowing. Some brain regions were active for only one of these two tasks. However, other areas responded for both tasks; these areas could be poten- tial sites of the multitasking bottle- neck. Two of these regions showed responses that reflected the degree of reaction-time slowing (Dux et al., 2006). The posterior lateral prefron- tal cortex and, to some extent, the SMA both showed this apparent slowing of response activity. For competition between these two simple cue–response tasks, the bottleneck appeared to be in areas low in the motor hierarchy. However,
multitasking at higher levels of cog- nitive control may have a different basis. For example, compared with healthy individuals, patients with damage to the frontopolar cortex have trouble multitasking among competing goals rather than com- peting responses (Dreher et al., 2008; Shallice & Burgess, 1991).
So why do we need to perform many motor tasks slowly, step by step (for example, performing long divi- sion to find the answer to 247 divided
by 13), although we can perform many kinds of much more complex sensory tasks simultaneously, quickly, and ef- fortlessly (for example, recognizing a face as male or female)? The answer may lie in the differing architecture of motor control versus sensory per- ception systems. Sensory perception involves integrating multiple features of a stimulus into a final result—a set of computations that can be done si- multaneously (in parallel), to arrive at an answer quickly (FIGURE 7.29a). Most
FIGURE 7.28 An fMRI study of multitasking. (a) Subjects had to press buttons according to an auditory cue and then, after a short interval (SOA), pronounce a syllable according to a visual cue (or vice versa). When the intervals were short, reaction times were delayed, suggesting a “bottleneck” in information processing. (b) fMRI localized this bottleneck to the medial and lateral prefrontal cortex, which were slower to activate as reaction times increased during multitasking (Dux et al., 2006).
(b)
(a)
Auditory stimulus
Manual response
Visual stimulus
Vocal response
Visual stimulus
SOA
SOA
Vocal response
Auditory stimulus
Manual response
Supplementary motor area Posterior lateral prefrontal cortex
T1 VVAM T2 VVAM T1 AMVV T2 AMVV
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ec )
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LEARNING OBJECTIVES provide a guide to what you will read and learn, helping you focus on the most im- portant points. Each learning objective corresponds to a major section of text.
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LEARNING OBJECTIVES By the end of this chapter, you should be able to: • Describe the basic underlying organization of all
vertebrate central nervous systems.
• Summarize the basic organization and structure of the peripheral nervous system.
• Explain the circuitry and function of spinal reflexes and central pattern generators.
• Distinguish the major components of the brainstem and their functions.
• Characterize the anatomy of the cerebellum and its role in motor function.
• Illustrate the role of the hypothalamus in homeostasis and the role of the thalamus as a relay and synchronization center, using examples.
• Identify the locations of the four lobes of the cerebral cortex, the locations of the major gyri and sulci, and their functions.
• Characterize the components of the basal ganglia and their functions.
• Distinguish the major components of the limbic system and their functions.
36
The Brain and Nervous System
STARTING OUT: The Brains of Creatures Great and Small
An Overview of the Nervous System The Peripheral Nervous System The Spinal Cord
CASE STUDY: Christopher Reeve, 1952–2004
THE BIGGER PICTURE: In Search of a Cure for Spinal Cord Injury
The Brainstem
NEUROSCIENCE OF EVERYDAY LIFE: Why Do We Get the Hiccups?
The Cerebellum The Diencephalon: Hypothalamus and Thalamus
CASE STUDY: Waking the Brain
The Telencephalon: Cerebral Cortex and Basal Ganglia
RESEARCH METHODS: Cytoarchitecture of the Cortex
Uniting the Inside and Outside Worlds
CHAPTER 2
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STARTING OUT scenarios begin each chapter with a gripping real-world example of chapter concepts, from hikers on Mount Everest who make a deadly mistake to a boy who functions normally following the removal of half of his brain.
104 PART 1 • ChAPTeR 4 Neuroplasticity
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STARTING OUT: The Child with half a Brain By the time Matthew S. was 6 years old, he began to have several epi- leptic seizures each day—some- times every few minutes. Medications were of no use. He was diagnosed with Rasmussen’s en- cephalitis, a rare, chronic inflam- matory disease that typically affects only a single brain hemisphere. His parents explored their options and were shocked to learn that there was only one known treatment for Rasmussen’s: removal of an entire hemisphere of the brain (Borgstein & Gottendorst, 2002).
But how could it be possible to live with half of the brain missing? Aren’t the functions of the brain dis- tributed widely across its territo- ries? Wouldn’t removal of one half be fatal—or at least devastating to Matthew’s quality of life?
With no remaining options, Mat- thew’s parents took him to Johns Hopkins Hospital in Baltimore, Maryland, where he underwent a hemispherectomy: the complete removal of half the cerebrum (FIGURE 4.1). The empty half of the skull filled up with cerebrospinal fluid, which shows up as a black void in neuroimaging.
Matthew walks with a slight limp on the opposite side of his body. Otherwise, he lives a normal life with almost no measurable deficit in cognition or behavior. How
can this be possible? Because the remainder of his brain has dynami- cally rewired to take over the miss- ing functions. The normal maps of the brain have redrawn themselves
on a smaller piece of neural real estate. How the brain accomplishes this remarkable feat—something no manmade machine can yet do— is the subject of this chapter.
FIGURE 4.1 Hemispherectomy. In a hemispherectomy, half the brain is surgically removed. This surgery has become standard operating procedure for Rasmussen’s encephalitis, a rare inflammatory disease that often affects only one hemisphere. Amazingly, as long as the surgery is performed before the age of 8, the child does remarkably well: the remainder of the brain dynamically rewires to take over the missing functions.
The brain is often thought of as a fixed organ with different regions dedicated to specific tasks. But the brain is better un- derstood as a dynamic system, constantly modifying its own circuitry to match the demands of the environment and the goals of the animal. This ongoing rewiring is the brain’s most fundamental principle and the source of its utility. Whereas your computer is built with hardwiring that remains fixed
from the assembly line onward, the brain dynamically recon- figures, ever so subtly, with each new experience. It reorga- nizes itself from the level of molecules in the synapses to the level of the gross anatomy visible to the naked eye. When you learn something new (such as your professor’s name), your brain physically changes. This ability to physically change, and to hold that change, is known as plasticity—just like the
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Preface xvii
A CLEAR, MODERN ART PROGRAM provides at- tractive biological drawings to help convey important con- cepts and information. Photographs and historical images also connect chapter content to the world around us.
316 PART 3 • CHAPTER 10 Sleep
rhythms in the absence of light cues like the sun (Rado, Gev, Goldman, & Terkel, 1991). Incredibly, this complex network of biochemical cascades results in a clock whose natural period is 24 hours and 11 minutes. Across subjects, the vari- ation is remarkably small: plus or minus only 16 minutes (Czeisler et al., 1999).
� e primary clock in mammals lies in the suprachias- matic nucleus (SCN) of the hypothalamus (from supra meaning “above” and chiasmatic referring to the optic chiasm, or crossing of the optic nerves; the two nuclei (le� and right hemisphere) are located just above this crossing, see FIGURE 10 . 9 ) (Klein, Moore, & Reppert, 1991). Cells of the SCN maintain their own rhythm when cultured in a dish, and damage of the SCN obliterates a regular sleep–wake rhythm in animals (Ibuka & Kawamura, 1975; Welsh, Logo- thetis, Meister, & Reppert, 1995).
Entrainment of the Circadian Rhythm by Light Cues Although the circadian rhythm is endogenously generated, it becomes entrained to various environmental stimuli, known as zeitgebers (Ascho� , Daan, & Honma, 1982). � e most important of these is the light–dark cycle—that is, the phase of the circadian rhythm is set by the planet’s rota- tion into and out of light from the sun. � e SCN receives information about external light levels from the eyes, but not via the rods or cones that we learned about in Chapter 5. � e retina contains a third type of light-sensitive cell: retinal
The Circadian Rhythm � ere are two important aspects to the sleep–wake cycle: how much you sleep and when you sleep. As for how much , not everyone needs to sleep the same amount. In fact, most people sleep about 6.5–7 hours a night, with a range between 4 and 11 hours (National Sleep Foundation, 2005, 2011, 2013). How many hours you sleep correlates with what you have done during the day, and periods of intense stimulation (such as going to a museum, an amusement park, and so on) during the day tend to make people sleep longer that night (Shapiro, Bortz, Mitchell, Bartel, & Jooste, 1981). How much sleep you require in general appears to have at least some ge- netic basis, although this is poorly understood (Franken, Malafosse, & Ta� i, 1999).
� e rest of this section discusses the second half of the story: when you sleep. Why is sleep on such a regular cycle, and with what environmental factors is the sleep cycle syn- chronized? � e answer to that question involves the circa- dian rhythm , a natural internal rhythm that runs on an approximately 24-hour cycle ( circa meaning “about” and dian referring to a “day”) and controls our sleep–wake cycles (Richardson, 2005). All animals appear to have some form of circadian cycle. � is circadian rhythm in� uences not just sleep and wakefulness, but also coordination, blood pres- sure, alertness, and body temperature ( FIGURE 10 . 8 ).
� e circadian rhythm is endogenously generated , meaning that it comes from programmed mechanisms in our brain and persists even in the absence of external cues. Some animals, like the blind mole rat, maintain their endogenous
FIGURE 10 . 8 Physiological changes tied to the circadian rhythm. The circadian rhythm runs on an approximately 24-hour cycle and controls sleep-wake cycles. It also influences other physiological and cognitive processes, including temperature, alertness, blood pressure, and hormone levels.
14:30 Best coordination
15:30 Best reaction time
17:00 Greatest heart efficiency and muscle strength
18:30 Highest blood pressure
19:00 Highest body temperature
21:00 Melatonin secretion begins2:00
Deepest sleep
4:30 Lowest body temperature
6:45 Rise in blood pressure
7:30 Melatonin secretion stops
9:00 High testosterone secretion
10:00 High alertness
00:00 Midnight
18:006:00
12:00 Noon
Chapter Ending KEY PRINCIPLES summarize the main points covered in the chapter—with one principle corresponding to each main heading—to remind you of what you have learned.
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Key Principles 469
Conclusion The architecture of motivation and reward is ancient and es- sential to survival. Motivations arise out of basic survival needs, and resources become rewards when they help us meet those needs. The brain has an elaborate and ancient mechanism for learning to predict the value of rewards in meeting future needs. This mechanism relies on accurate measurements of the internal state and accurate measure- ments of prediction error: the difference between the ex- pected and the actual outcomes. When these mechanisms operate normally, they are effective at keeping us alive.
Unfortunately, these mechanisms can be hijacked, with devastating consequences. By altering neurotransmitter sig- nals, some substances can create powerful illusions of well- being and illusions of always being better than expected. When this happens, a vicious circle of pathological learning begins. This pathological learning creates an unnaturally strong motivation to obtain and consume the addictive sub- stance. The strength of this motivation increases over subse- quent exposures, in time eclipsing other survival needs.
Undoing this process is as difficult as unlearning a skill. Current treatments for addiction rely on a combination of counseling and, in some cases, medications to alleviate with- drawal and reduce the high risk of relapse. Unfortunately, effective treatments for addiction have been hard to find, and substance abuse ranks among the most devastating causes of disability and death worldwide. Finding a safe and effective treatment for substance dependence is one of the major goals in the neuroscience of the 21st century.
Paradoxically, this often requires that the doctor take the part of discussing the reasons not to change. The pa- tient then has no option but to take up the other side of the argument. For example, a physician might begin by asking the patient, “On a scale of 1 to 10, how much do you want to quit smoking?” The patient might reply, “Not much, maybe a 2.” Although the physician’s natural reaction might be to splutter, “ Don’t you realize the damage that smoking does to your health?”, such an approach only makes the patient defensive, encouraging him to generate self- convincing reasons why it’s too hard or too stressful to quit right now. A doctor trained in MI might instead say, “A 2 out of 10 . . . OK, so why not a 1? Why not a zero?” This gets the person thinking and talking about the reasons they actually do want to change: setting a good example for their children, not wanting to leave their spouse alone and bereft in old age. By eliciting the other side of the argu- ment, the physician gets the patient to think a little harder about his or her own motivations for overcoming the habit of smoking. With time and discussion, these motivations move to the forefront, increasing the chances that the person will make a break from prior habits (Rollnick, Miller, & Butler, 2007).
MI is not a cure- all, but it seems to work better than be- rating people for their bad habits in the hopes this will per- suade them to change. Its take- home point: the motivation to change is hard to implant in someone from the outside. Instead, it must be found within the person’s own set of values, goals, hopes, and ideals and then carefully coaxed out into the forefront. So next time you find yourself “wanting someone to want” something, give this approach a try.
KEY PRINCIPLES
• Natural motivations, such as eating, drinking, and reproductive behaviors, help inform the brain what is needed and how to value those needs at the cur- rent time.
• The hypothalamus is important for homeostasis and for evaluating internally driven motivation, whereas the amygdala receives input from the ex- ternal world and evaluates the importance of these outside factors. Dopamine is the neurotransmitter that most commonly conveys these reward and motivation signals.
• The brain learns to predict rewards by comparing the expected outcome of an action to the actual outcome. “ Better- than-expected” outcomes in- crease the motivation toward that action in the future.
• “Liking” and “wanting” are two different things in the brain. Liking refers to the sensation of well- being that is being experienced at that time, whereas wanting refers to a future expectation of well- being.
• Opioids are naturally occurring chemicals in the brain that reduce pain and increase pleasure. These effects can be mimicked by synthetic opi- oids, such as morphine, heroin, and codeine. Recent research has identified several types of opioid receptors. Stimulation of some of these re- duces pain, but stimulation of others produces un- pleasant sensations.
• The neurotransmitter dopamine is important for motivation and in learning to predict rewards. It is especially important for assigning value to those
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terms listed by subsection, with page numbers, for easy exam review.
THE BIGGER PICTURE connects neuroscience to larger concepts and questions—social, ethical, legal, and historical. Examples include how neuroscience can help us make better decisions and whether we will one day be able to equip our brains with new senses.
214 PART 2 • ChAPTeR 7 The Motor System
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THE BIGGER PICTURE Neural Implants for Motor Control Remember the tragic case of Jean- Dominique Bauby from the begin- ning of the chapter? He had suffered a stroke affecting the medulla and the nearby pyramidal decussations: the outgoing pathway for virtually all of the motor cortex. Locked-in syndrome is the usual result of this kind of stroke. Without any connec- tions to the spinal cord or lower cra- nial nerves, the cortex is unable to send output to nearly any parts of the body. A few cranial nerves above the injury may be spared, allowing some movement of the eyes and eyelids only. Since these pathways do not easily regenerate, the hopes for recovery are slim.
Yet the motor cortex itself is still intact. What if there were some way to read the activity of the upper motor neurons directly? Could we use them to drive an ar- tificial arm, or a wheelchair, or a computer cursor? Perhaps even a simple speech synthesizer? In fact, for more than a decade, neu- roscientists have been using brain– computer interface tech- nologies to help locked-in patients communicate with the outside world (Kennedy, Bakay, Moore, Adams, & Gold waithe, 2000). One strategy involves implanting a set of electrodes directly in the motor cortex, in the part of the homuncu- lus that controls the hand or the mouth. The electrodes may be coated with neural growth factors to encourage the neurons to grow connections to the implant itself.
As one example, in 2006, a re- search team from Massachusetts General Hospital and Brown Univer- sity inserted a small electrode array (FIGURE 7.17) into the motor cortex of Matthew N., a 25-year-old who had
been quadriplegic since a knife attack five years earlier. A success- ful example of “neuromotor pros- thetics” in humans, the sensor was able to read the signals that Mat- thew’s brain was trying to send out to his body, and convert those sig- nals into commands that could direct the movements of a pros- thetic arm or a pointer on a com- puter screen (Hochberg et al., 2006).
Decoding the neural activity into meaningful signals can be difficult. Computer algorithms can learn to in- terpret the collective activity of a pop- ulation of neurons as signaling a particular movement, or syllable. The patients themselves can learn to use the prostheses, rewiring the local connections to improve their ability to communicate over time. More
recently, a young man with locked-in syndrome has begun to use this system to produce simple syllables with reasonable accuracy after sev- eral years of training for both patient and computer (Guenther et al., 2009).
So far, the technology is still in its infancy. Interpreting the neural signals remains difficult. The sig- nals themselves fade over time as glial cells gradually build up around the implanted electrodes, render- ing the electrodes useless. At pres- ent, no implant has been able to match the efficacy of the simple blink-coding card used by M. Bauby to write his memoirs. Developing effective brain–computer inter- faces will be one of the most impor- tant technological challenges of the 21st century.
FIGURE 7.17 A neuromotor prosthetic. A tiny electrode array implanted in the motor cortex of Matthew N. enabled him to control his wheelchair and actions on a screen.
07-Eagleman_Chap07.indd 214 28/10/15 3:06 pmRESEARCH METHODS shows how we know what we know about the brain, presenting important research tech- niques and indicating the types of research questions that these techniques have been used to investigate.
The Cells of the Brain 81
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RESEARCH METHODS: Visualizing Neurons and Their Products The discovery that neurons are dis- crete, fundamental units of the ner- vous system was made possible by the ability to stain them. Several techniques allow the visualization of neurons. Golgi staining is a tech- nique that impregnates some frac- tion of neurons with a dark material, allowing the entirety of individual cells to be seen under a microscope (FIGURE 3.10a). This is the method that birthed Ramón y Cajal’s neuron doctrine. Another technique, Nissl staining, uses a chemical that binds to the RNA in cell bodies, thereby allowing the visualization of somas (FIGURE 3.10b). Nissl staining is most commonly used for judging sizes of cells and their densities.
Several other methods are utilized to obtain detailed pictures of nervous tissue. In the technique of autoradi- ography, a radioactive substance is designed to be taken up by specific cells but not by others (FIGURE 3.10c). Then, when a photographic emulsion is placed over thin slices of the brain tissue, the emulsion is exposed by the radioactivity in the same way that film is exposed by light. In this way, it can be seen which cell types absorbed the substance in question (for example, a pharmaceutical drug).
In the technique of immunocyto- chemistry, antibodies are developed that bind only to specific proteins (FIGURE 3.10d). These antibodies are washed onto a slice of brain tissue, and they attach wherever the protein of interest is being expressed. With some chemical steps, these antibod- ies can be visualized, revealing the
exact locations of the protein within the cell. A related technique is to use radioactively labeled stretches of RNA or DNA that will bind to specific
stretches of messenger RNA (mRNA); this is called in situ hybridization, and it reveals which cells have expressed a gene of interest (FIGURE 3.10e).
A
C
FIGURE 3.10 Different techniques to bring the invisibly small world of neurons to light. (a) Golgi staining, (b) Nissl staining, (c) autoradiography, (d) immunocytochemistry, and (e) in situ hybridization.
(a) (b)
(d)
(e)(c)
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# 158305 Cust: OUP Au: Eagleman Pg. No. xviii Title: Brain and Behavior, 1e
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xviii Preface
• Chapter Summaries: Full summaries of each chapter provide a thorough review of the important facts and con- cepts covered.
• Flashcards: Interactive flashcard activities are an effec- tive way for students to learn and review all of the impor- tant terminology.
• Practice Quizzes: Each chapter includes a practice quiz, which students can use as a self-review exercise, to check their understanding.
For Instructors An extensive and thoughtful supplements program offers in- structors everything they need to prepare their course and lectures, and assess student progress.
Ancillary Resource Center (ARC) For more information, go to www.oup.com/us/eagleman
Available online exclusively to adopters, the Ancillary Resource Center (ARC) includes all of the instructor re- sources that accompany Brain and Behavior: A Cognitive Neuroscience Perspective.
Instructor’s Manual: For each chapter of the textbook, the Instructor’s Manual includes the following: • Chapter Overview • Chapter Outlines • Key Concepts • Suggested Online Activities • Journal Articles and Press Releases
Textbook Figures and Tables: All of the textbook’s il- lustrations and tables are provided in a variety of formats, including high and low resolution, with and without balloon captions, and unlabeled (all balloon captions, labels, and leaders removed).
PowerPoint Resources: • Figures and Tables: This presentation includes all of the
figures and tables (all formats) from the chapter, with titles.
• Lecture: A complete lecture outline, ready for use in class. Includes coverage of all important facts and concepts pre- sented in the chapter along with selected figures and tables. Animations: All of the animations from Dashboard are
available in the ARC for download, making it easy to include them in lecture presentation and online course materials. (Also available in Dashboard.)
Videos: A collection of videos selected to accompany each chapter helps bring some of the key concepts from the textbook to life. Ideal for use as lecture starters or paired with assignments.
Test Bank: A complete test bank provides instructors with a wide range of test items for each chapter, including multiple-choice, fill-in-the-blank, short-answer, true/false, and essay questions. Questions are noted for whether they
REVIEW QUESTIONS test your recall and under- standing of the key information presented in the chapter to aid with studying.
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Critical- Thinking Questions 555
REVIEW QUESTIONS
1. In what parts of the brain do we tend to see the plaques and tangles of Alzheimer’s disease? Why do you think that removing these plaques and tangles turned out to be a failure as a strat- egy for treating Alzheimer’s disease?
2. Is it a reasonable metaphor to call frontotempo- ral dementia a “cancer of the soul?” What does this name tell us about our own intuitions on which faculties are proper to the “spirit,” as op- posed to the neural circuits of the brain?
3. Why does increased creativity emerge as a symptom of some neurological and psychiatric disorders? What do these disorders have in common?
4. In Huntington’s disease, a disruption of a single gene gives rise to a diverse set of neurological and psychiatric symptoms. What are these symptoms, what common disease process cre- ates them, and how does this common process disrupt such a wide variety of brain functions?
5. What different kinds of symptoms can arise in Tourette syndrome? What common neuro- anatomical changes might give rise to these symptoms?
6. Should we describe symptoms of Tourette syn- drome as “voluntary” or “involuntary”? Why so? Should ordinary blinking be considered a kind of motor tic? Why or why not?
7. Obsessive–compulsive disorder is traditionally considered a psychiatric rather than a neurolog- ical disorder. How would you classify the illness? Why? Do you think it should still be considered a single category of disease? Why or why not?
8. What are the positive and the negative symp- toms of schizophrenia? Which symptoms can be relieved with currently available treatments and which cannot? Why don’t these treatments provide a permanent cure?
9. What are three kinds of similarities between bi- polar disorder and schizophrenia? What are some of the key differences between these dis- orders? Should we think of them as two forms of the same illness? Why or why not?
10. What kinds of factors can lead to depression? Is depression really a chemical imbalance in the brain, as it is sometimes described? What and where are the brain abnormalities of depression? How has this knowledge led to new treatments?
CRITICAL- THINKING QUESTIONS
1. Imagine that you are the head of a pharmaceuti- cal research group that is dedicated to develop- ing new, effective drug treatments for Alzheimer’s disease. Given what you have learned in this chapter about the physiological basis and progression of Alzheimer’s, what pro- cesses would you design the drugs to target? At what stage of the disease do you think that such drugs would be most effective? Explain.
2. How do patients with Alzheimer’s disease, fron- totemporal dementia, Tourette syndrome, obsessive–compulsive disorder, and bipolar disorder differ from one another in terms of their self- awareness of their symptoms? How
could such differences be used to guide the de- velopment of new, specialized treatments for each disease?
3. Scientists are continuing to learn more about the influence of genetics on the likelihood of de- veloping diseases such as schizophrenia. How- ever, we are unable to know for certain whether individuals are destined to develop such dis- eases. If an effective neuroprotective treatment were discovered for schizophrenia, do you think it would be ethical to administer it to individuals who are at risk for developing the disease? What factors would influence your viewpoint? Explain your reasoning.
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CRITICAL-THINKING QUESTIONS ask you to apply and extend information from the chapter to new sce- narios to help you master the material.
# 158305 Cust: OUP Au: Eagleman Pg. No. 555 Title: Brain and Behavior, 1e
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Critical- Thinking Questions 555
REVIEW QUESTIONS
1. In what parts of the brain do we tend to see the plaques and tangles of Alzheimer’s disease? Why do you think that removing these plaques and tangles turned out to be a failure as a strat- egy for treating Alzheimer’s disease?
2. Is it a reasonable metaphor to call frontotempo- ral dementia a “cancer of the soul?” What does this name tell us about our own intuitions on which faculties are proper to the “spirit,” as op- posed to the neural circuits of the brain?
3. Why does increased creativity emerge as a symptom of some neurological and psychiatric disorders? What do these disorders have in common?
4. In Huntington’s disease, a disruption of a single gene gives rise to a diverse set of neurological and psychiatric symptoms. What are these symptoms, what common disease process cre- ates them, and how does this common process disrupt such a wide variety of brain functions?
5. What different kinds of symptoms can arise in Tourette syndrome? What common neuro- anatomical changes might give rise to these symptoms?
6. Should we describe symptoms of Tourette syn- drome as “voluntary” or “involuntary”? Why so? Should ordinary blinking be considered a kind of motor tic? Why or why not?
7. Obsessive–compulsive disorder is traditionally considered a psychiatric rather than a neurolog- ical disorder. How would you classify the illness? Why? Do you think it should still be considered a single category of disease? Why or why not?
8. What are the positive and the negative symp- toms of schizophrenia? Which symptoms can be relieved with currently available treatments and which cannot? Why don’t these treatments provide a permanent cure?
9. What are three kinds of similarities between bi- polar disorder and schizophrenia? What are some of the key differences between these dis- orders? Should we think of them as two forms of the same illness? Why or why not?
10. What kinds of factors can lead to depression? Is depression really a chemical imbalance in the brain, as it is sometimes described? What and where are the brain abnormalities of depression? How has this knowledge led to new treatments?
CRITICAL- THINKING QUESTIONS
1. Imagine that you are the head of a pharmaceuti- cal research group that is dedicated to develop- ing new, effective drug treatments for Alzheimer’s disease. Given what you have learned in this chapter about the physiological basis and progression of Alzheimer’s, what pro- cesses would you design the drugs to target? At what stage of the disease do you think that such drugs would be most effective? Explain.
2. How do patients with Alzheimer’s disease, fron- totemporal dementia, Tourette syndrome, obsessive–compulsive disorder, and bipolar disorder differ from one another in terms of their self- awareness of their symptoms? How
could such differences be used to guide the de- velopment of new, specialized treatments for each disease?
3. Scientists are continuing to learn more about the influence of genetics on the likelihood of de- veloping diseases such as schizophrenia. How- ever, we are unable to know for certain whether individuals are destined to develop such dis- eases. If an effective neuroprotective treatment were discovered for schizophrenia, do you think it would be ethical to administer it to individuals who are at risk for developing the disease? What factors would influence your viewpoint? Explain your reasoning.
16-Eagleman-ch16.indd 555 08-Jul-15 8:19:12 PM
Media and Supplements to Accompany Brain and Behavior: A Cognitive Neuroscience Perspective
For Students
Companion Website Available at no additional cost, the Companion Website pro- vides students with the following review resources: • Chapter Outlines: Detailed outlines give an overview of
each chapter.
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Preface xix
progress instantly. Dashboard includes the following resources: • Quizzes: For each chapter of the textbook, there is a quiz
to test student comprehension of important facts and concepts introduced in the chapter.
• A postlecture summative quiz designed to be used as an assessment of student mastery of the important facts and concepts introduced in the chapter, after the student has read the chapter and attended the relevant lecture/class period/discussion section.
• Animations: A set of detailed animations helps students understand some of the book’s more complex topics and processes by presenting them in a clear, easy-to-follow narrative.
LMS Course Cartridges For those instructors who wish to use their campus learning management system, a course cartridge containing all of the Dashboard resources is available for a variety of e-learning environments. (For more information, please contact your local Oxford representative.)
are factual or conceptual, and for level of difficulty. All ques- tions from the Dashboard and Companion Website quizzes (see below) are also included.
Computerized Test Bank: The Test Bank is also pro- vided in Blackboard Diploma format (software included). Diploma makes it easy to create quizzes and exams using any combination of publisher-provided questions and an in- structor’s own questions and to export those assessments for print or online delivery in a wide range of learning manage- ment system formats.
Dashboard For more information, go to www.oup.com/us/dashboard
Oxford’s Dashboard learning management system fea- tures a streamlined interface that connects instructors and students with the functions they perform most often, simpli- fying the learning experience to save instructors time and put students’ progress first. Dashboard’s prebuilt assess- ments were created specifically to accompany Brain and Behavior: A Cognitive Neuroscience Perspective and are au- tomatically graded so that instructors can see student
Acknowledgments Over the past several years, many talented brains have devoted their cognitive powers to making Brain and Behavior a reality. We express our heartfelt gratitude for the hard work, patience, and dedication of the team at Oxford University Press, without whom this book would not exist. Special thanks go to editorial director Patrick Lynch, who first saw the project’s potential, as well as John Challice, vice president and publisher, who also supported the book at an early stage. Kind thanks to our editor, Jane Potter, who encouraged us steadily onward from deadline to deadline, with patience and gentle persuasion. We are also ever grateful to development editor Anne Kemper, senior devel- opment editor Lisa Sussman, and assistant editor Maura Mac- Donald for their careful attention to detail throughout the manuscript. We also thank the Oxford production team for transforming the unadorned drafts of the manuscript chapters into such an eye-catching and engaging final form: Lisa Grzan,
production manager, Jane Lee and Keith Faivre, senior produc- tion editors, and Susan Brown, copyeditor. Kudos to art direc- tor Michele Laseau and senior designer Caitlin Wagner for their truly beautiful work on the interior and the cover design. Thanks also for the essential efforts of Eden Gingold, marketing manager, Kateri Woody, marketing associate, and Frank Mor- timer, director of marketing, for helping to bring this book before a wide audience of curious minds. Grateful acknowledg- ment is made to the talents of the team at Dragonfly Media for the art program for the book—specifically, art development and art direction by Mike Demaray; art production by Mike Demaray, Craig Durant, Helen Wortham, and Rob Fedirko; and chapter openers and cover art by Craig Durant. Finally, thanks to all of our reviewers, anonymous and otherwise, for their suggestions and insightful comments on the early drafts of this work:
Lewis Barker, Auburn University
Diane E. Beals, University of Tulsa
Patricia Bellas, Irvine Valley College
Annemarie Bettica, Manhattanville College
Christopher Braun, Hunter College
Blaine Browne, Valdosta State University
David Bucci, Dartmouth College
Amanda N. Carey, Simmons College
Cynthia R. Cimino, University of South Florida
Barbara Clancy, University of Central Arkansas
Howard Casey Cromwell, Bowling Green State University
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xx Preface
Kelly L. Curtis, High Point University
Deana Davalos, Colorado State University
Scott Decker, University of South Carolina
Dean Dessem, University of Maryland
Vonetta Dotson, University of Florida
Jeffrey Eells, Mississippi State University–Main
Paul Engelhardt, Michigan State University
Joseph Farley, Indiana University–Bloomington
Robert Faux, Duquesne University
Robert P. Ferguson, Buena Vista University
Jane Flinn, George Mason University
Jay Friedenberg, Manhattan College
Jonathan Gewirtz, University of Minnesota
Edward Golob, Tulane University
Kim Gorgens, University of Denver
Jinger S. Gottschall, Penn State University
Jay E. Gould, University of West Florida
Sayamwong E. Hammack, University of Vermont
Valerie Gray Hardcastle, University of Cincinnati
Linda Hermer, University of Florida
Elaine M. Hull, Florida State University
Daniel Hummer, Morehouse College
Mark Hurd, University of Texas
Eric Jackson, University of New Mexico
Daniel Jacobson, Madonna University
Mark Jareb, Sacred Heart University
Penelope L. Kuhn, California State University–Chico
Matthew Kurtz, Wesleyan University
Eric Laws, Longwood University
Ben Lester, University of Iowa
Linda Lockwood, Metropolitan State College of Denver
Jeannie Loeb, University of North Carolina–Chapel Hill
Keith B. Lyle, University of Louisville
Cyrille Magne, Middle Tennessee State University
Kai McCormack, Spelman College
Ming Meng, Dartmouth College
Maura Mitrushina, California State Northridge
Daniel Montoya, Fayetteville State University
Andrea Morris, University of California–Los Angeles
Ezequiel Morsella, San Francisco State University
Andrea Nicholas, University of California–Irvine
J. Ian Norris, Berea College
Jamie Olavarria, University of Washington
Matthew Palmatier, East Tennessee State University
Jim H. Patton, Baylor University
Tadd B. Patton, Georgia Regents University
Richard Payne, University of Maryland–College Park
Michael Sakuma, Dowling College
Haline Schendan, Plymouth University
Lynda Sharrett-Field, University of Kentucky
Robert W. Sikes, Northeastern University
Scott Slotnick, Boston College
Kenith V. Sobel, University of Central Arkansas
Jessica Stephens, Texas A&M University at Kingsville
Jeffrey Taube, Dartmouth College
Sheralee Tershner, Western New England University
Jason Themanson, Illinois Wesleyan University
Lucien T. Thompson, University of Texas at Dallas
Lucy J. Troup, Colorado State University
Jonathan Vaughan, Hamilton College
Sandy Venneman, University of Houston–Victoria
Todd D. Watson, Lewis & Clark College
Douglas A. Weldon, Hamilton College
Robin Wellington, St. John's University
Mark West, Rutgers University
Adrienne Williamson, Kennesaw State University
John L. Woodard, Wayne State University
With special thanks to our Advisory Panel:
Alex Michael Babcock, Montana State University
Peter Brunjes, University of Virginia
Arne Ekstrom, University of California–Davis
Samuel McClure, Stanford University
Juan Salinas, University of Texas at Austin
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David Eagleman is a neuroscientist, New York Times best- selling author, and Guggenheim Fellow who holds joint ap- pointments in the departments of neuroscience and psychiatry at Baylor College of Medicine in Houston, Texas. Dr. Eagleman’s areas of research include time perception, vision, synesthesia, and the intersection of neuroscience with the legal system. He directs the Laboratory for Percep- tion and Action and is the founder and director of Baylor College of Medicine’s Initiative on Neuroscience and Law. Dr. Eagleman has written several neuroscience books, in- cluding Incognito: The Secret Lives of the Brain and Wednesday Is Indigo Blue: Discovering the Brain of Synesthesia. He has also written an internationally bestselling book of literary fiction, Sum, which has been translated into 28 languages and turned into two operas in Sydney and London. Dr. Eagleman is the author and presenter of “The Brain,” an international six- hour series on PBS that poses the question, “What does it mean to be human?” from a neuroscientist’s point of view. Dr. Eagleman has written for the Atlantic, New York Times, Discover Magazine, Slate, Wired, and New Scientist and ap- pears regularly on National Public Radio and BBC.
Jonathan Downar is the Director of the MRI-Guided rTMS Clinic at the University Health Network in Toronto, Canada, and a scientist at the Toronto Western Research Institute. He currently holds appointments with the Department of Psychiatry and the Institute of Medical Science at the Uni- versity of Toronto.
As a physician-scientist, his clinical work focuses on using noninvasive brain stimulation to treat patients with severe and medication-resistant forms of psychiatric illness, including depression, bipolar disorder, obsessive– compulsive disorder, post–traumatic stress disorder, and eating disor- ders. His research work focuses on developing a new genera- tion of more effective, more accessible, and less costly techniques for brain stimulation in these disorders. His re- search laboratory also focuses on developing tests that use functional MRI and EEG to predict the most effective treat- ment parameters for individual patients.
In addition to his research and clinical work, he teaches undergraduate courses in the neuroscience of social cogni- tion, emotion regulation, decision making, and other forms of complex human behavior. He also teaches medical students and psychiatry resident physicians on the subjects of neuro- anatomy, neuroimaging, and therapeutic brain stimulation.
ABOUT THE AUTHORS
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A Cognitive Neuroscience Perspective
Brain and Behavior
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2
LEARNING OBJECTIVES By the end of this chapter, you should be able to: • Describe the central aims of cognitive
neuroscience.
• Distinguish between understanding the brain’s components and understanding its functioning.
• Compare and contrast the advantages and disadvantages of each of four categories of research methods for studying brain function.
• Summarize three common forms of cognitive bias in human thinking and how the key elements of the scientific method address cognitive biases.
• Describe five big questions in cognitive neuroscience.
• Show at least three ways in which advances in neuroscience may benefit human society.
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Introduction STARTING OUT: A Spark of Awe in the Darkness
Who Are We? In Pursuit of Principles How We Know What We Know
RESEARCH METHODS: Magnetic Resonance Imaging
Thinking Critically about the Brain The Big Questions in Cognitive Neuroscience The Payoffs of Cognitive Neuroscience
CHAPTER 1
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4 PART 1 • ChAPTeR 1 Introduction
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STARTING OUT: A Spark of Awe in the Darkness On October 9, 1604, a brilliant spark of light grew to life in the darkness of the night sky over Europe. A few days later, the astronomer Johannes Kepler began to gaze up at the new star that had appeared in the void, outshining all its peers, visible for a time even through the brightness of the day. Kepler wrote extensively on the astronomical properties of the new star, or stella nova, whose sudden appearance challenged the conventional wisdom that the heavens were fixed and unchang- ing (Kepler, [1604] 2004). Over the ensuing months, the new star faded gradually back into the ce- lestial background. Nothing simi- lar has appeared in our skies to surpass it since then, even four centuries later.
Today’s astronomers would have called Kepler’s star a super- nova and could have told him some astonishing details about the nature of the object that captured his attention on that clear night so long ago (FIGURE 1.1). They could have told him about a star several times more massive than the Sun, reaching the end of a lifespan mea- sured in eons, collapsing suddenly in upon itself to form a core blazing at a hundred billion degrees, then bursting outward again in a cata- clysmic explosion that, for a time, shone brighter than the entire sur- rounding galaxy. The light of that distant explosion was obliged to sear through space to arrive tens of thousands of years later in the night sky over Europe, drawing human eyes upward in wonder.
Yet a supernova, for all its mag- nificence and rarity, is still outshined
by marvels closer to home. As Kepler stood under the stars, be- holding the bright spark in the dark- ness before his eyes, even rarer and more wondrous events were taking place in the darkness behind his eyes. In that mysterious vault within his skull, a spark, not of light but of awe, was taking form in the warm, dark passageways of his brain. Where the supernova burned with common light and heat, his mind burned with a rare and incandes- cent emotion. Where the supernova shed its light blindly in all directions, his mind turned its attention to one tiny facet of the universe beyond, striving to see more clearly. Where the supernova was unaware of its own grandeur, Kepler’s mind was capable of reflect
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