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AN INTRODUCTION TOBRAIN AND BEHAVIOR
The image on the cover illustrates a cell-staining technique called Brainbowdeveloped by Jean Livet and his colleagues at ...
To the first neuron, our ancestors, our families, and                             students who read this bookSenior Publish...
AN INTRODUCTION TOBRAIN AND BEHAVIOR                                 Third Edition             BRYAN KOLB             Univ...
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About the Authors                                  Bryan Kolb received his Ph.D. from the Penn-                           ...
Index of Disorders     Name of disorder is followed by chapter     Deafness 10                                 MPTP poison...
Contents in BriefPreface xviiCHAPTER 1      What Are the Origins of Brain and Behavior? 1CHAPTER 2      How Does the Nervo...
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Contents              Preface xvii                                                Plastic Patterns of Neural Organization ...
x        ContentsCells of the Nervous System 71                                                     Voltage-Sensitive Chan...
Contents     xi                                  RESEARCH FOCUS 6-1 Stimulating the Brain                   Origins of Neu...
xii        ContentsFactors Influencing Individual Responses                                                The Visual Brai...
Contents        xiii                     RESEARCH FOCUS 11-1 The Brain–Computer                     Biology, Evolution, an...
xiv      ContentsPacemaking Circannual Rhythms 454                                                             Neural Circ...
Contents      xv                                     CLINICAL FOCUS 15-4 The Sodium Amobarbital                   RESEARCH...
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Preface     he Third Edition of An Introduction to Brain and Behavior continues to reflect theT    evolution of the field ...
xviii    PREFAC E Chapter-by-Chapter Updates in the Third Edition Chapter 1 now opens with a Clinical Focus on traumatic b...
Preface   xixplay of genes and environment. This ongoing interplay influences a person’s brain andbehavior from earliest i...
xx   PREFAC E                introduction to brain science. We provide the philosophical and anatomical back-             ...
Preface   xxiexplains what we do, whether it is talking, sleeping, seeing, or learning. Readers willtherefore find nearly ...
xxii   PREFAC E                    Big-Picture Emphasis                  One challenge in writing an introductory book on ...
Preface   xxiiielectronic versions of illustrations in the book (through Worth Publishers’ Image andLecture Gallery), and ...
xxiv   PREFAC E                  wide variety of applied, conceptual, and factual questions have been thoroughly re-      ...
Preface   xxvThe Brain and Behavior Video Segments Available on VHS, this special collection in-cludes targeted selections...
xxvi   PREFAC E                  specialists who found photographs and other illustrative materials that we would not     ...
CHAPTERWilliam West/AFP/Getty Images                                                                                      ...
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  1. 1. AN INTRODUCTION TOBRAIN AND BEHAVIOR
  2. 2. The image on the cover illustrates a cell-staining technique called Brainbowdeveloped by Jean Livet and his colleagues at Harvard University. (The name is a playon the word “rainbow.”) In this Brainbow image of a mouse hippocampus, the braincells are expressing fluorescent proteins. The Brainbow technique, which labels cellsby highlighting them with distinct colors, involves introducing into mice genes thatproduce cyan (blue), green, yellow, and red fluorescent proteins. The “red” gene isobtained from coral and the “blue” and “green” genes are obtained from jellyfish. Theextent to which each gene is activated varies somewhat, owing to chance factors, andby mixes of the primary colors produces fluorescence in one hundred or moredifferent hues. Brainbow offers a way to image the shape and location of cells and todescribe neuronal circuits—that is, where each neuron sends its processes and how itinterconnects with other neurons. Lichtman Laboratory/Harvard University
  3. 3. To the first neuron, our ancestors, our families, and students who read this bookSenior Publisher: Catherine WoodsSenior Acquisitions Editor: Charles LinsmeierDevelopment Editor: Barbara BrooksMarketing Coordinator: Jennifer BilelloSenior Project Editor: Georgia Lee HadlerManuscript Editors: Patricia Zimmerman and Penelope HullProduction Manager: Sarah SegalMedia and Supplements Editor: Peter TwicklerText and Cover Design: Vicki TomaselliPhoto Editor: Ted SzczepanskiSenior Illustration Coordinator: Bill PageIllustrations: Dragonfly Media Group and Northeastern Graphic, Inc.Composition: Northeastern Graphic, Inc.Printing and Binding: Worldcolor VersaillesLibrary of Congress Control Number: 2009936819ISBN-13: 9780716776918ISBN-10: 071677691XCopyright © 2011, 2006, 2001 by Worth PublishersAll rights reservedPrinted in the United States of AmericaFirst printingWorth Publishers41 Madison AvenueNew York, NY 10010www.worthpublishers.com
  4. 4. AN INTRODUCTION TOBRAIN AND BEHAVIOR Third Edition BRYAN KOLB University of Lethbridge IAN Q. WHISHAW University of Lethbridge WORTH PUBLISHERS
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  6. 6. About the Authors Bryan Kolb received his Ph.D. from the Penn- sylvania State University in 1973. He conducted postdoctoral work at the University of Western Ontario and the Montreal Neurological Institute. He moved to the University of Lethbridge in 1976, where he is currently Professor of Neuro- science and holds a Board of Governors Chair in Neuroscience. His current research examines how neurons of the cerebral cortex change inresponse to various factors, including hormones, experience, psychoactive drugs,neurotrophins, and injury, and how these changes are related to behavior in thenormal and diseased brain. Kolb is a Fellow of the Royal Society of Canada, theCanadian Psychological Association (CPA), the American Psychological Associa-tion, and the Association of Psychological Science. He is a recipient of the HebbPrize from CPA and from the Canadian Society for Brain, Behaviour, and Cogni-tive Science (CSBBCS) and has received honorary doctorates from the Universityof British Columbia and Thompson Rivers University. He is currently a memberof the Experience-Based Brain and Behavioral Development program of the Cana-dian Institute for Advanced Research.Ian Q. Whishaw received his Ph.D. from theUniversity of Western Ontario in 1971. Hemoved to the University of Lethbridge in 1970,where he is currently Professor of Neuroscienceand holds a Board of Governors Chair in Neuro-science. He has held visiting appointments at theUniversity of Texas, University of Michigan,Cambridge University, and the University ofStrasbourg, France. He is a Fellow of Clair Hall, Cambridge. His current researchexamines how the precise details of movements are influenced by injury or diseaseto the motor systems of rodents and humans and how animals and humans movethrough real and mental space. Whishaw is a Fellow of the Canadian Psychologi-cal Association, the American Psychological Association, and the Royal Society ofCanada, and the Institute for Scientific Information includes him in its list of mostcited neuroscientists. He is a recipient of a Bronze medal from the CanadianHumane Society, and a recipient of the Ingrid Speaker medal for research. He hasreceived honorary doctorates from Thompson Rivers University and the Univer-sity of Lethbridge. v
  7. 7. Index of Disorders Name of disorder is followed by chapter Deafness 10 MPTP poisoning 5*, 16 number(s) where information appears; Deafferentation 11, 14 Multiple sclerosis (MS) 3*, 4*, 8, 16 * indicates a Focus box. Dementia 6, 14*, 16 Myasthenia gravis 4*, 8 For a summary of general treatment Depression 5, 6, 8*, 12*, 16 Myopia 9 categories, see the table on page 577. Developmental disorders 6*, 7, 11*, 14*, Narcolepsy 13 15*, 16 Neurotoxins 5, 8, 16 For a summary of research techniques, see Diaschisis 16 Night terrors 13 Chapter 6. Down syndrome 3, 7 Obesity 12 Drug-dependency insomnia 13 Obsessive compulsive disorder (OCD) Addiction 6, 8*, 12 Drug-induced behavioral sensitization 5, 8, 12, 16 Affective disorders 12*, 16 8, 14 Optic ataxia 9 Agenesis of the frontal lobe 7, 12* Drug-induced psychosis 6, 8* Pain 8, 11, 12*, 16 Agnosias 9 Dyskinesia 5*, 7, 8, 11, 16 Panic disorder 12* Akathesia 16 Dyslexia 14* Paraplegia 3, 11* Akinesia 6 Encephalitis 2* Parkinson’s disease 2, 5*, 6, 11, 16 Alcohol myopia 8 Environmental deprivation 6, 7*, 16 Persistent vegetative state 1* Alzheimer’s disease 2, 5, 6, 14*, 16 Epilepsy 4*, 6, 9*, 10*, 16 Petit mal seizure 16 Amblyopia 7 Extinction 15 Phenylketonuria (PKU) 7, 16 Amnesia 6, 14* Fetal alcohol syndrome (FAS) 7, 8* Phobias 12, 16 Amusia 10 Festination 16 Postictal depression 16 Amyotrophic lateral sclerosis (ALS) 4*, Focal seizure 16 Posttraumatic stress disorder (PTSD) 5, 11* Frontal leucotomy 12 8, 12, 16* Androgen insensitivity syndrome 12* Generalized anxiety 12, 16 Presbyopia 9 Androgenital syndrome 12* Genetic disorders 3, 6, 7, 16 Psychosis 8, 16 Anencephaly 7 Glioma 3* Quadrantanopia 9 Aneurysm 10* Grand mal seizure 16 Quadripelegia 11 Anorexia nervosa 12 Hemianopia 9 Restless legs syndrome (RLS) 13* Anterograde amnesia 14* Hemorrhagic stroke 2* Retrograde amnesia 14* Anxiety disorders 8, 12*, 16 Hormonal disorders 7, 8, 12, 16 Schizophrenia 5, 6, 7*, 8, 16 Aphasias 6, 10 Huntington’s chorea 3* Scotoma 9 Apraxia 11 Hydrocephalus 3 Seasonal affective disorder (SAD) 13* Arteriovenous malformations 10* Hyperopia 9 Seizure 4*, 10, 16 Ataxia 9 Hypnogogic hallucinations 13 Sleep apnea 13* Attention-deficit/hyperactivity disorder Idiopathic seizure 16 Sleep paralysis 13 (ADHD) 6*, 7, 16 Insomnia 13 Spinal-cord injury 2, 8, 11, 12 Autism spectrum disorder (ASD) 7, 11*, Ischemia 2*, 16 Split-brain syndrome 15 15* Klüver–Bucy syndrome 12 Stress 2, 6, 7, 8, 12, 16 Autoimmune disease 4*, 16 Korsakoff ’s syndrome 14* Stroke 2*, 6, 16 Bell’s palsy 2* Language disorders 6, 7, 10, 14*, 15*, 16 Substance abuse 8, 12 Bipolar disorder 8, 16 Learning disabilities 6, 7, 14*, 16 Sudden infant death syndrome (SIDS) Blindsight 9* Locked-in syndrome 11*, 13 7*, 13* Brain tumors 3*, 6 Lou Gehrig’s disease (ALS) 4*, 11* Suicide 6, 8*, 16 Carbon monoxide poisoning 9* Major depression 5, 6, 8*, 12, 16 Symptomatic seizure 16 Cataplexy 13 Mania 5, 8, 16 Synesthesia 15* Catatonic posture 16 Memory deficits 6, 14 Tardive dyskinesia 16 Cerebral aneurysm 10* Meningioma 3* Tay–Sachs disease 3, 16 Cerebral palsy 7*, 11 Meningitis 2* Tourette’s syndrome 2, 5, 11* Cognitive disorders 6, 7, 15 Mental retardation 2*, 7, 16 Traumatic brain injury (TBI) 1*, 6, 14, Color-deficient vision 9* Metastatic tumor 3* 16 Coma 1*, 4, 8, 13, 16 Migraine 9* Tumors 3*, 6 Contralateral neglect 15, 16 Minimally conscous state (MCS) 1* Visual-form agnosia 9vi
  8. 8. Contents in BriefPreface xviiCHAPTER 1 What Are the Origins of Brain and Behavior? 1CHAPTER 2 How Does the Nervous System Function? 31CHAPTER 3 What Are the Units of Nervous-System Function? 69CHAPTER 4 How Do Neurons Transmit Information? 103CHAPTER 5 How Do Neurons Communicate and Adapt? 133CHAPTER 6 How Do We Study the Brain’s Structure and Functions? 169CHAPTER 7 How Does the Nervous System Develop and Adapt? 199CHAPTER 8 How Do Drugs and Hormones Influence the Brain and Behavior? 237CHAPTER 9 How Do We Sense, Perceive, and See the World? 279CHAPTER 10 How Do We Hear, Speak, and Make Music? 317CHAPTER 11 How Does the Nervous System Respond to Stimulation and Produce Movement? 353CHAPTER 12 What Causes Emotional and Motivated Behavior? 395CHAPTER 13 Why Do We Sleep and Dream? 443CHAPTER 14 How Do We Learn and Remember? 481CHAPTER 15 How Does the Brain Think? 519CHAPTER 16 What Happens When the Brain Misbehaves? 561Glossary G-1References R-1Name Index NI-1Subject Index SI-1 vii
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  10. 10. Contents Preface xvii Plastic Patterns of Neural Organization 33 Functional Organization of the Nervous System 34 The Brain’s Surface Features 35 CHAPTER 1 THE BASICS: Finding Your Way Around the Brain 36 What Are the Origins of Brain and CLINICAL FOCUS 2-2 Meningitis and Encephalitis 39 Behavior? 1 The Brain’s Internal Features 40 CLINICAL FOCUS 1-1 Living with Brain Injury 2 CLINICAL FOCUS 2-3 Stroke 41 Neuroscience in the Twenty-First Century 2 Why Study Brain and Behavior? 3 Evolutionary Development of the Nervous System 45 What Is the Brain? 3 The Central Nervous System: Mediating Behavior 46 The Spinal Cord 47 RESEARCH FOCUS 1-2 Recovering Consciousness 4 The Brainstem 47 Gross Anatomy of the Nervous System 6 The Forebrain 51 What Is Behavior? 7 The Somatic Nervous System: Transmitting Perspectives on Brain and Behavior 8 Information 56 Aristotle and Mentalism 8 The Cranial Nerves 56 Descartes and Dualism 9 The Spinal Nerves 57 COMPARATIVE FOCUS 1-3 The Speaking Brain 10 CLINICAL FOCUS 2-4 Magendie, Bell, and Darwin and Materialism 11 Bell’s Palsy 59 Evolution of Brain and Behavior 13 The Autonomic Nervous System: Balancing Origin of Brain Cells and Brains 14 Internal Functions 60 Classification of Life 14 Evolution of Animals Having Nervous Systems 15 Ten Principles of Nervous-System Function 62 The Chordate Nervous System 16 Principle 1: The Nervous System’s Function Is to Produce Movement Within a Perceptual World Evolution of the Human Brain and Behavior 18 Created by the Brain 62 Humans: Members of the Primate Order 18 Principle 2: The Details of Nervous-System Functioning Australopithecus: Our Distant Ancestor 19 Are Constantly Changing, an Attribute Called The First Humans 20 Neuroplasticity 62 Relating Brain Size and Behavior 21 Principle 3: Many of the Brain’s Circuits Are Crossed 62 Climate and the Enlarging Hominid Brain 22 Principle 4: The Central Nervous System Functions Why the Hominid Brain Enlarged 23 on Multiple Levels 63 Principle 5: The Brain Is Both Symmetrical and Modern Human Brain Size and Intelligence 25 Asymmetrical 63 Fallacies of Human Brain-Size Comparisons 26 Principle 6: Brain Systems Are Organized Both Culture 27 Hierarchically and in Parallel 64 COMPARATIVE FOCUS 1-4 Evolution and Adaptive Principle 7: Sensory and Motor Divisions Exist Throughout Behavior 28 the Nervous System 64 Principle 8: Sensory Input to the Brain Is Divided for Object Summary 29 Recognition and Motor Control 65 Key Terms 30 Principle 9: Functions in the Brain Are Both Localized and Distributed 66 Principle 10: The Nervous System Works by Juxtaposing CHAPTER 2 Excitation and Inhibition 66 How Does the Summary 67 Nervous System Key Terms 68 Function? 31 CHAPTER 3Jeff Minton RESEARCH FOCUS 2-1 Brain Size and Human What Are the Units of Nervous-System Behavior 32 Function? 69 An Overview of Brain Function and Structure 33 RESEARCH FOCUS 3-1 Programming Behavior 70 ix
  11. 11. x ContentsCells of the Nervous System 71 Voltage-Sensitive Channels and the Action Potential 126Neurons: The Basis of Information Processing 72 Into the Nervous System and Back Out 128Five Types of Glial Cells 76 How Sensory Stimuli Produce Action Potentials 128CLINICAL FOCUS 3-2 Brain Tumors 78 How Nerve Impulses Produce Movement 129CLINICAL FOCUS 3-3 Multiple Sclerosis 80 CLINICAL FOCUS 4-4 Lou Gehrig’s Disease 130THE BASICS: Chemistry Review 84 Summary 131Internal Structure of a Cell 82 Key Terms 132The Cell As a Factory 82The Cell Membrane: Barrier and Gatekeeper 86 CHAPTER 5The Nucleus: Site of Gene Transcription 87The Endoplasmic Reticulum: Site of RNA Synthesis 88 How Do NeuronsProteins: The Cell’s Product 89 CommunicateGolgi Bodies and Microtubules: Protein Packaging and and Adapt? 133 Shipment 89 Katsuyoshi TanakaCrossing the Cell Membrane: Channels, Gates, RESEARCH FOCUS 5-1 The and Pumps 90 Basis of Neural Communication in aGenes, Cells, and Behavior 92 Heartbeat 134Chromosomes and Genes 93Genotype and Phenotype 94 A Chemical Message 134Dominant and Recessive Alleles 94 Structure of Synapses 136Genetic Mutations 95 CLINICAL FOCUS 5-2 Parkinson’s Disease 138Mendel’s Principles Apply to Genetic Disorders 95 Neurotransmission in Four Steps 139CLINICAL FOCUS 3-4 Huntington’s Chorea 97 Varieties of Synapses 141Chromosome Abnormalities 97 Excitatory and Inhibitory Messages 141Genetic Engineering 98 Evolution of Complex Neurotransmission Systems 142RESEARCH FOCUS 3-5 Brainbow: Rainbow Neurons 100 Varieties of Neurotransmitters 143Summary 101 Four Criteria for Identifying Neurotransmitters 143Key Terms 102 Three Classes of Neurotransmitters 145 CLINICAL FOCUS 5-3 Awakening with L-Dopa 147CHAPTER 4 Two Classes of Receptors 149How Do Neurons Transmit Information? 103 Neurotransmitter Systems and Behavior 151CLINICAL FOCUS 4-1 Epilepsy 104 Neurotransmission in the Somatic Nervous System 151 Two Activating Systems of the AutonomicSearching for Electrical Activity in the Nervous Nervous System 152System 105 Four Activating Systems in the Central Nervous System 153Early Clues That Linked Electricity and Neural Activity 105 CLINICAL FOCUS 5-4 The Case of the Frozen Addict 156THE BASICS: Electricity and Electrical Stimulation 106 Role of Synapses in Three Kinds of LearningTools for Measuring a Neuron’s Electrical Activity 108 and in Memory 157How the Movement of Ions Creates Electrical Charges 110 Habituation Response 158Electrical Activity of a Membrane 113 Sensitization Response 159Resting Potential 113 Long-Term Potentiation and Associative Learning 161Graded Potentials 115 Learning As a Change in Synapse Number 163The Action Potential 117 RESEARCH FOCUS 5-5 Dendritic Spines, SmallRESEARCH FOCUS 4-2 Light-Sensitive Ion Channels 119 but Mighty 165The Nerve Impulse 120 Summary 166Saltatory Conduction and Myelin Sheaths 121 Key Terms 168How Neurons Integrate Information 123Excitatory and Inhibitory Postsynaptic Potentials 123 CHAPTER 6CLINICAL FOCUS 4-3 Myasthenia Gravis 124 How Do We Study the Brain’s StructureSummation of Inputs 125 and Functions? 169
  12. 12. Contents xi RESEARCH FOCUS 6-1 Stimulating the Brain Origins of Neurons and Glia 205 into Action 170 Growth and Development of Neurons 206 Measuring Brain and Behavior 171 CLINICAL FOCUS 7-2 Cerebral Palsy 208 Linking Neuroanatomy and Behavior 171 Glial Development 213 Methods of Behavioral Neuroscience 172 Manipulating and Measuring Brain–Behavior Correlating Behavior with Nervous-System Interactions 176 Development 214 Motor Behaviors 214 Measuring the Brain’s Electrical Activity 179 Language Development 215 EEG Recordings of Graded Potentials 179 Development of Problem-Solving Ability 216 Mapping Brain Function with Event-Related Potentials 180 A Caution about Linking Correlation to Causation 220 Magnetoencephalography 182 Recording Action Potentials from Single Cells 182 Brain Development and the Environment 220 CLINICAL FOCUS 6-2 Mild Head Injury and Experience and Cortical Organization 220 Depression 183 Experience and Neural Connectivity 222 RESEARCH FOCUS 7-3 Increased Cortical Activation for Static Imaging Techniques: CT and MRI 184 Second Languages 223 Dynamic Brain Imaging 186 Critical Periods for Experience and Brain Development 225 Functional Magnetic Resonance Imaging 186 Abnormal Experience and Brain Development 226 Positron Emission Tomography 187 Optical Tomography 189 CLINICAL FOCUS 7-4 Romanian Orphans 227 Hormones and Brain Development 228 Chemical and Genetic Measures of Brain and Injury and Brain Development 230 Behavior 190 Drugs and Brain Development 231 Measuring the Brain’s Chemistry 190 Other Kinds of Abnormal Brain Development 231 Measuring Genes in Brain and Behavior 191 Developmental Disability 232 CLINICAL FOCUS 6-3 Cannabis Use, Psychosis, CLINICAL FOCUS 7-5 Schizophrenia 233 Genetics 192 How Do Any of Us Develop a Normal Brain? 234 Epigenetics 192 Summary 235 Using Animals in Brain–Behavior Research 193 Key Terms 236 Benefits of Creating Animal Models of Disease 194 Animal Welfare and Scientific Experimentation 194 CHAPTER 8 RESEARCH FOCUS 6-4 Attention-Deficit/Hyperactivity Disorder 195 How Do Drugs and Hormones Influence the Brain and Behavior? 237 Summary 197 Key Terms 198 CLINICAL FOCUS 8-1 Addiction 238 Principles of Psychopharmacology 239 Drug Routes into the Nervous System 239 CHAPTER 7© Alessandra Sanguinetti/Magnum Drug Action at Synapses: Agonists and Antagonists 242 How Does the An Acetylcholine Synapse: Examples of Drug Action 242 Nervous System Tolerance 244 Sensitization 245 Develop and Adapt? 199 Classification of Psychoactive Drugs 248 Class I: Antianxiety Agents and Sedative Hypnotics 249 RESEARCH FOCUS 7-1 Linking Serotonin CLINICAL FOCUS 8-2 Fetal Alcohol Syndrome 250 to SIDS 200 Class II: Antipsychotic Agents 251 Three Perspectives on Brain Development 201 Class III: Antidepressants 252 Predicting Behavior from Brain Structure 201 Class IV: Mood Stabilizers 253 Correlating Brain Structure and Behavior 201 CLINICAL FOCUS 8-3 Major Depression 254 Influences on Brain and Behavior 202 Class V: Opioid Analgesics 254 Neurobiology of Development 202 Class VI: Psychomotor Stimulants 256 Gross Development of the Human Nervous System 203 Class VII: Psychedelic and Hallucinogenic Stimulants 258
  13. 13. xii ContentsFactors Influencing Individual Responses The Visual Brain in Action 311to Drugs 259 Injury to the Visual Pathway Leading to the Cortex 311Behavior on Drugs 259 Injury to the “What” Pathway 312Addiction and Dependence 261 Injury to the “How” Pathway 313Sex Differences in Addiction 262 CLINICAL FOCUS 9-4 Carbon Monoxide Poisoning 314Explaining and Treating Drug Abuse 263 Summary 315Wanting-and-Liking Theory 264Why Doesn’t Everyone Abuse Drugs? 265 Key Terms 316Treating Drug Abuse 266Can Drugs Cause Brain Damage? 267 CHAPTER 10 Joe Klamar/APF/Getty ImagesCLINICAL FOCUS 8-4 Drug-Induced Psychosis 269 How Do We Hear,Hormones 270 Speak, and MakeHierarchical Control of Hormones 270 Music? 317Classes and Functions of Hormones 271 RESEARCH FOCUS 10-1Homeostatic Hormones 272 The Evolution of LanguageGonadal Hormones 272 and Music 318Anabolic–Androgenic Steroids 273Stress Hormones 274 Sound Waves: The Stimulus for Audition 319Ending a Stress Response 275 Physical Properties of Sound Waves 319Summary 276 Perception of Sound 323Key Terms 278 Properties of Language and Music As Sounds 324 Functional Anatomy of the Auditory System 325CHAPTER 9 Structure of the Ear 326How Do We Sense, Perceive, and See Auditory Receptors 328the World? 279 Pathways to the Auditory Cortex 329 Auditory Cortex 330CLINICAL FOCUS 9-1 Migraines and a Case of Blindsight 280 Neural Activity and Hearing 332 Hearing Pitch 332The Nature of Sensation and Perception 281 Detecting Loudness 333Sensory Receptors 281 Detecting Location 333Neural Relays 283 Detecting Patterns in Sound 334Sensory Coding and Representation 283Perception 284 Anatomy of Language and Music 335 Processing Language 335Functional Anatomy of the Visual System 285Structure of the Retina 285 RESEARCH FOCUS 10-2 Distinct Cortical Areas for Second Languages 337THE BASICS Visible Light and the Structure of the Eye 286 CLINICAL FOCUS 10-3 Left-Hemisphere Dysfunction 341Photoreceptors 289 CLINICAL FOCUS 10-4 Arteriovenous Malformations 342CLINICAL FOCUS 9-2 Visual Illuminance 290 Processing Music 343Retinal-Neuron Types 291 CLINICAL FOCUS 10-5 Cerebral Aneurysms 344Visual Pathways 292Dorsal and Ventral Visual Streams 294 RESEARCH FOCUS 10-6 The Brain’s Music System 345Location in the Visual World 297 Auditory Communication in NonhumanCoding Location in the Retina 298 Species 346Location in the Lateral Geniculate Nucleus and Cortical Birdsong 346 Region V1 298 Echolocation in Bats 348The Visual Corpus Callosum 300 Summary 350Neuronal Activity 301 Key Terms 352Seeing Shape 301Seeing Color 307 CHAPTER 11RESEARCH FOCUS 9-3 Color-Deficient Vision 308 How Does the Nervous System Respond toNeural Activity in the Dorsal Stream 310 Stimulation and Produce Movement? 353
  14. 14. Contents xiii RESEARCH FOCUS 11-1 The Brain–Computer Biology, Evolution, and Environment 401 Interface 354 Evolutionary Influences on Behavior 402 The Chemical Senses 403 Hierarchical Control of Movement 355 Environmental Influences on Behavior 407 The Forebrain and Initiation of Movement 356 Inferring Purpose in Behavior: To Know a Fly 409 The Brainstem and Species-Typical Movement 358 The Spinal Cord and Executing Movement 360 Neuroanatomy of Motivated Behavior 410 Regulatory and Nonregulatory Behavior 411 CLINICAL FOCUS 11-2 Autism Spectrum Disorder 361 The Regulatory Function of the Hypothalamic Circuit 412 CLINICAL FOCUS 11-3 Spinal-Cord Injury 362 The Organizing Function of the Limbic Circuit 417 The Executive Function of the Frontal Lobes 418 Organization of the Motor System 363 The Motor Cortex 363 CLINICAL FOCUS 12-2 Agenesis of the Frontal Lobe 420 Corticospinal Tracts 365 Stimulating Emotion 421 Motor Neurons 366 Explanations for Emotion 421 Control of Muscles 367 The Amygdala and Emotional Behavior 423 The Motor Cortex and Skilled Movement 368 The Prefrontal Cortex and Emotional Behavior 424 Control of Skilled Movement in Nonhuman Species 369 Emotional Disorders 425 How Motor-Cortex Damage Affects Skilled CLINICAL FOCUS 12-3 Anxiety Disorders 426 Movement 369 Control of Regulatory Behavior 427 The Basal Ganglia and the Cerebellum 371 Controlling Eating 428 The Basal Ganglia and Movement Force 371 CLINICAL FOCUS 12-4 Weight-Loss Strategies 429 CLINICAL FOCUS 11-4 Tourette’s Syndrome 372 Controlling Drinking 432 The Cerebellum and Movement Skill 373 Control of Nonregulatory Behavior 433 Organization of the Somatosensory System 376 Effects of Sex Hormones on the Brain 433 Somatosensory Receptors and Perception 377 Dorsal-Root Ganglion Neurons 379 CLINICAL FOCUS 12-5 Androgen-Insensitivity Syndrome Somatosensory Pathways to the Brain 380 and the Androgenital Syndrome 434 Spinal Reflexes 381 The Hypothalamus, the Amygdala, and Sexual Behavior 435 Feeling and Treating Pain 382 Sexual Orientation, Sexual Identity, and Brain RESEARCH FOCUS 11-5 Phantom Limb Pain 383 Organization 436 Cognitive Influences on Sexual Behavior 437 The Vestibular System and Balance 386 Reward 438 Exploring the Somatosensory Cortex 387 The Somatosensory Homunculus 388 Summary 441 Effects of Damage to the Somatosensory Cortex 389 Key Terms 442 RESEARCH FOCUS 11-6 Tickling 390 CHAPTER 13 The Somatosensory Cortex and Complex Movement 391 Summary 393 Why Do We Sleep and Dream? 443 Key Terms 394 COMPARATIVE FOCUS 13-1 The Variety of Biological Rhythms 444 CHAPTER 12 A Clock for All Seasons 444 What Causes Origins of Biological Rhythms 445 Biological Clocks 446 Emotional and Biological Rhythms 446John Henley/Corbis Motivated Free-Running Rhythms 448 Behavior? 395 Zeitgebers 449 RESEARCH FOCUS 12-1 CLINICAL FOCUS 13-2 Seasonal Affective Disorder 450 The Pain of Rejection 396 Neural Basis of the Biological Clock 452 Identifying the Causes of Behavior 397 Suprachiasmatic Rhythms 452 Behavior for Brain Maintenance 397 Dual Clocks 453 Drives and Behavior 399 Immortal Time 453 Neural Circuits and Behavior 400 What Ticks? 454 The Nature of Behavior: Why Cats Kill Birds 400 Pacemaking Circadian Rhythms 454
  15. 15. xiv ContentsPacemaking Circannual Rhythms 454 Neural Circuit for Implicit Memories 500 Neural Circuit for Emotional Memories 500RESEARCH FOCUS 13-3 Synchronizing Biorhythms at the Molecular Level 455 Structural Basis of Brain Plasticity 501 Measuring Synaptic Change 502Sleep Stages and Dreaming 456 Enriched Experience and Plasticity 504Measuring How Long We Sleep 456 Sensory or Motor Training and Plasticity 505Measuring Sleep in the Laboratory 457 Plasticity, Hormones, Trophic Factors, and Drugs 508Stages of Waking and Sleeping 457A Typical Night’s Sleep 458 RESEARCH FOCUS 14-5 Movement, Learning, andNREM Sleep and REM Sleep 459 Neuroplasticity 509CLINICAL FOCUS 13-4 Restless Legs Syndrome 460 Some Guiding Principles of Brain Plasticity 512Dreaming 461 Recovery from Brain Injury 513What We Dream About 461 Donna’s Experience with Traumatic BrainWhat Does Sleep Accomplish? 465 Injury 514 Three-Legged Cat Solution 515Sleep As a Passive Process 465 New-Circuit Solution 515Sleep As a Biological Adaptation 465 Lost-Neuron-Replacement Solution 516Sleep As a Restorative Process 466Sleep and Memory Storage 468 Summary 517 Key Terms 518Neural Bases of Sleep 469Reticular Activating System and Sleep 470Neural Basis of EEG Changes Associated with Waking 471 CHAPTER 15 Group, University of Louisiana at Lafayette, New Iberia Research. Courtesy of Cognitive EvolutionNeural Basis of REM Sleep 472 How Does the BrainSleep Disorders 473 Think? 519Disorders of Non-REM Sleep 473 COMPARATIVE FOCUS 15-1Disorders of REM Sleep 474 Animal Intelligence 520CLINICAL FOCUS 13-5 Sleep Apnea 475 The Nature ofWhat Does Sleep Tell Us about Thought 521Consciousness? 477 Characteristics of Human Thought 522Summary 478 The Neural Unit of Thought 523Key Terms 480 Cognition and the Association Cortex 525 Knowledge about Objects 526CHAPTER 14 Multisensory Integration 527 Spatial Cognition 527How Do We Learn and Remember? 481 Attention 529CLINICAL FOCUS 14-1 Remediating Dyslexia 482 Planning 531 Imitation and Understanding 532Connecting Learning and Memory 483 RESEARCH FOCUS 15-2 Consequences of Mirror-NeuronStudying Learning and Memory in the Laboratory 483 Dysfunction 533Two Categories of Memory 485What Makes Explicit and Implicit Memory Different? 487 Cognitive Neuroscience 534What Is Special about Personal Memories? 489 CLINICAL FOCUS 15-3 NeuropsychologicalDissociating Memory Circuits 490 Assessment 535Disconnecting Explicit Memory 491 Cerebral Asymmetry in Thinking 537CLINICAL FOCUS 14-2 Patient Boswell’s Amnesia 492 Anatomical Asymmetry 537 Functional Asymmetry in Neurological Patients 538Disconnecting Implicit Memory 492 Functional Asymmetry in the Normal Brain 540Neural Systems Underlying Explicit and Implicit The Split Brain 540 Explaining Cerebral Asymmetry 543Memories 493 The Left Hemisphere, Language, and Thought 544Neural Circuit for Explicit Memories 493 Variations in Cognitive Organization 545CLINICAL FOCUS 14-3 Alzheimer’s Disease 496 Sex Differences in Cognitive Organization 545CLINICAL FOCUS 14-4 Korsakoff’s Syndrome 499 Handedness and Cognitive Organization 549
  16. 16. Contents xv CLINICAL FOCUS 15-4 The Sodium Amobarbital RESEARCH FOCUS 16-2 Treating Behavioral Disorders Test 550 with TMS 573 Synesthesia 551 Understanding and Treating Neurological CLINICAL FOCUS 15-5 A Case of Synesthesia 551 Disorders 577 Traumatic Brain Injury 577 Intelligence 552 Stroke 579 The Concept of General Intelligence 553 Epilepsy 581 Multiple Intelligences 553 Multiple Sclerosis 583 Divergent and Convergent Intelligence 554 Neurodegenerative Disorders 584 Intelligence, Heredity, Environment, and the Synapse 555 Are Parkinson’s and Alzheimer’s Aspects of One Disease? 590 Age-Related Cognitive Loss 590 Consciousness 556 Why Are We Conscious? 556 Understanding and Treating Behavioral What Is the Neural Basis of Consciousness? 557 Disorders 592 Summary 558 Psychotic Disorders 592 Key Terms 560 Mood Disorders 595 RESEARCH FOCUS 16-3 Antidepressant Action in CHAPTER 16 Neurogenesis 597Gordon Parks/Time & Life Pictures/ What Happens Anxiety Disorders 598 When the Brain Is Misbehavior Always Bad? 599 Misbehaves? 561 Summary 600 RESEARCH FOCUS 16-1 Key Terms 601Getty Images Posttraumatic Stress Disorder 562 Glossary G-1 Multidisciplinary Research on Brain and Behavioral Disorders 564 References R-1 Causes of Abnormal Behavior 564 Investigating the Neurobiology of Behavioral Disorders 565 Name Index NI-1 Classifying and Treating Brain and Behavioral Disorders 568 Subject Index SI-1 Identifying and Classifying Behavioral Disorders 568 Treatments for Disorders 570
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  18. 18. Preface he Third Edition of An Introduction to Brain and Behavior continues to reflect theT evolution of the field of behavioral neuroscience. Perhaps the two biggest develop-ments are the rapid emergence of noninvasive imaging to study cognitive processing inhuman and nonhuman subjects as well as the burgeoning evidence of the plasticity ofbrain structure and function. We have thus added more imaging and plasticity conceptsand research to the text and in Focus boxes. The biggest change in the book has been the addition of a chapter dedicated to meth-ods. Chapter 6, “How Do We Study the Brain’s Structure and Functions?” was a responseto requests from instructors to have a dedicated discussion showcasing the range ofmethods used by behavioral neuroscientists to study brain and behavior. The chapterincludes both traditional methods as well as newer techniques such as optical tomog-raphy and epigenetics. Material from the Second Edition’s Appendix on animal welfarehas been incorporated in Chapter 6 in the context of how we develop animal modelsof human disorders. Expanded discussions of other techniques appear in other chap-ters where appropriate. For example, Chapter 16 includes a discussion of the new meth-ods of virtual-reality therapies for anxiety-related disorders. Several new features have been added throughout the book. We had originally in-cluded background material such as anatomical orientation (Chapter 2) and basic chem-istry (Chapter 3) within the main text. This material is now separated into boxes called“The Basics.” So the material is still there for readers who are not familiar with it, andreaders familiar with the information can easily skip it. Another new feature is margin notes that increase the reader’s ease in finding re-lated information elsewhere in the book. This should be especially helpful when con-cepts are introduced early in the text and then elaborated on in later chapters. Readerscan return quickly to the earlier discussion to refresh their knowledge of the basic con-cept. The margin notes should also allow instructors to move through the book to pre-view later discussions. Much of the book remains familiar. Throughout, we continue to examine the nerv-ous system with a focus on function, on how our behavior and our brains interact. Westructured the First Edition of An Introduction to Brain and Behavior on key questionsthat students and neuroscientists ask about the brain:• Why do we have a brain?• How is the nervous system organized?• How do drugs affect our behavior?• How does the brain learn?• How does the brain think?As it was then, our goal in the new edition is to bring coherence to a vast subject byhelping students understand the big picture. Asking fundamental questions about thebrain has another benefit: it piques students’ interest in the subject and challenges themto join us on the journey of discovery that is brain science. Scientific understanding of the brain and human behavior continues to grow at anexponential pace. We want to communicate the excitement of recent breakthroughs inbrain science and to relate some of our own experiences in studying the brain for thepast 40 years, both to make the field’s developing core concepts and latest revelationsunderstandable and meaningful and to take uninitiated students to the frontiers of phys-iological psychology. xvii
  19. 19. xviii PREFAC E Chapter-by-Chapter Updates in the Third Edition Chapter 1 now opens with a Clinical Focus on traumatic brain injury (TBI) and its treatments (Research Focus 1-2) and is refocused, briefer, and more inviting. Chapter 2 includes a streamlined emphasis on neuroplasticity and nervous-system anatomy and function, providing a road map for the rest of the book. Chapter 3 streamlines the coverage of the nervous system and adds a “The Basics” feature on its most referenced material. Clinical Focus 3-3 provides updated coverage of multiple sclerosis, and new Research Focus 3-5 covers Brainbow imaging. Chapter 4 improves the presentation of electrical activity at the cell membrane, including back propagation. It also features new Research Focus 4-2 on light-sensitive ion channels. Chapter 5 expands coverage of neural activating systems and streamlines coverage of learning at the synapse. Chapter 6 showcases a state-of-the-art range of techniques and topics including optimal tomography and epigenetics and includes new Research Focus 6-4 on ADHD. Chapter 7 begins with new Research Focus 7-1 on SIDS, and new Research Focus 7-3 discusses cortical representation of second languages. Chapter 8 includes new Clinical Focus 8-1 on addiction and a new section on sex differences in and treatments for drug abuse. It also includes a new section on anabolic steroids. Chapter 9 expands coverage of sensation and perception. The coverage of vision is more streamlined than it was in the previous edition for more balance with the other senses. New Research Focus 9-3 covers color-deficient vision. Chapter 10 includes new coverage of “bilingual” brains and a new Research Focus on the neural basis of musical ability. The chapter also includes a new example from Oliver Sacks’s Musicophilia. Chapter 11 includes new Research Focus 11-1 on the computer–brain interface (BCI) and new Research Focus 11-5 on treating phantom limb pain. Clinical Focus 11-2 on autism spectrum disorder has been updated. Chapter 12 includes a new section on unconscious olfactory processing in humans and updated coverage on weight-loss strategies based on results from comparative research on diets. Chapter 13 features new information on why some of us are morning people and others evening people. It also presents expanded discussion of how genes influence daily biorhythms and of a neural basis for cataplexy, and it updates clinical coverage of SAD, RLS, and sleep apnea. Chapter 14 emphasizes seven principles of neuroplasticity. Clinical Focus 14-1 includes a technique, featured in The New Science of Learning on PBS, for remediating dyslexia. Coverage of patient H. M. is updated and includes his photo, which was released after his death. Also illustrated is dramatic cortical healing of a stroke lesion. Chapter 15 discusses research on the role of the human mirror-neuron system in social cognition and the implications of its dysfunction in autism spectrum disorder. The chapter includes a new section on multisensory integration, and it updates coverage of the binding problem and incidence of synesthesia. Chapter 16 includes research on the neural basis of PTSD; success of virtual reality simulations as therapy; a new section on age- related cognitive loss; updated treatment therapies using TMS, DBS, growth factors, and stem cells. Emphasis is on the efficacy of talking therapies—including real-time fMRI and cognitive-behavioral therapy—used in conjunction with drugs or in lieu of them. We feature the relation between the brain and behavior in every chapter. For ex- ample, when we first describe how neurons communicate in Chapter 5, we also describe how plasticity in connections between neurons can serve as the basis of learning. Later, in Chapter 14, which is directed to the question of how we learn and remember, we explore how interactions among different parts of the brain enable our more complex behaviors. Emphasis on Evolution, Genetics, Plasticity and Psychopharmacology To make sure that this book conveys the excitement of current neuroscience as re- searchers understand it, we emphasize evolution, genetic research, neural plasticity, and psychopharmacology throughout. After all, evolution results from the complex inter-
  20. 20. Preface xixplay of genes and environment. This ongoing interplay influences a person’s brain andbehavior from earliest infancy through old age. We cover the evolution of the brain in depth in Chapters 1 and 2 and return to this per-spective of neuroscience in an evolutionary context in almost every chapter. Examples:• Evolution of the synapse in Chapter 5• Evolution of the visual pathways in Chapter 9• How natural selection might promote overeating in Chapter 12• Evolutionary theories of sleeping and dreaming in Chapter 13• Evolution of sex differences in spatial cognition and language in Chapter 15• Links between our evolved reactions to stress and the development of anxiety disorders in Chapter 16 The foundations of genetic research, including knockout technology, are introducedin depth in Chapter 3. Chapter 5 includes discussions of metabotropic receptors andDNA and of learning and genes. The role of genes in development is integral to Chap-ter 7, as is the topic of genes and drug action in Chapter 8. Chapter 9 explains the ge-netics of color vision, and the genetics of sleep disorders is a major topic in Chapter 13.Chapter 16 considers the role of genetics in understanding the causes of behavioraldisorders. We have updated and expanded the Research Focus boxes to showcase new and in-teresting developments on topics relevant to each chapter. In Chapter 6, for example,we explore research on attention-deficit/hyperactivity disorder (ADHD) in the contextof developing animal models of disease. In Chapter 12 we investigate brain activationin response to physical and emotional pain and find their effects to be surprisinglysimilar. We have added special emphasis in this edition on common behavioral disor-ders such as traumatic brain injury (TBI), addiction, Alzheimer’s disease, Parkinson’sdisease, autism spectrum disorder (ASD), and posttraumatic stress disorder (PTSD). InChapter 16, for example, we review new approaches to the treatment of PTSD in vet-erans of the wars in Iraq and Afghanistan. Chapter 8 deals with drugs and behavior, and we turn to this topic in many otherchapters as well. You will find coverage of drugs and information transfer in Chapter4, drugs and cellular communication in Chapter 5, drugs and motivation in Chapter12, drugs and sleep disorders in Chapter 13, neuronal changes with drug use in Chap-ter 14, and drugs as treatments for a range of disorders in Chapter 16. Neural plasticity continues as a hallmark of the book. We introduce the concept inChapter 2 and expand on it throughout, including an elaboration of basic principles inChapter 14. Scientific Background ProvidedWe describe the journey of discovery that is neuroscience in a way that students justbeginning to study the brain and behavior can understand. We have found that thiscourse can sometimes be daunting, largely because understanding brain function re-quires information from all the basic sciences. If we are ultimately interested in how thebrain understands language or music, for example, the properties of sound waves andthe functioning of the ear become relevant because the ear is where auditory informa-tion enters the brain. These encounters with basic science can be both a surprise and a shock to intro-ductory students, who often come to the course without the necessary background.Our approach provides all the background students require to understand an
  21. 21. xx PREFAC E introduction to brain science. We provide the philosophical and anatomical back- ground needed to understand modern evolutionary theory, offer a short introduc- tion to chemistry before describing the chemical activities of the brain, and briefly discuss electricity before exploring the brain’s electrical activity. Because some of this information is very familiar to students with an extensive science background, we have separated some basic material from the main text into boxes called “The Ba- sics.” Readers already comfortable with the material can easily skip it, and less expe- rienced readers can still review it. Similarly, we review basic psychological facts such as behavioral development in Chapter 7 and the forms of learning and memory in Chapter 14. In these ways, students can tackle brain science with greater confidence. Students in social science disciplines often remark on the amount of biology and chemistry in the book, whereas an equal number of students in biological sciences remark on the amount of psychology. More than half the students enrolled in our Bachelor of Science in Neuroscience program have switched from biochemistry or psychology majors after taking this course. We must be doing something right! Clinical Focus Maintained We repeatedly emphasize that neuroscience is a human science—that everything in this book is relevant to our lives. Neuroscience helps us understand how we learn, how we develop, and how we can help people who suffer from sometimes deadly and destruc- tive brain and behavioral disorders. We have found that emphasizing the clinical as- pects of neuroscience is especially useful in motivating introductory students and in demonstrating to them the relevance of our field. Clinical material helps to make neurobiology particularly relevant to those who are going on to careers in psychology, social work, or other professions related to men- tal health, as well as to students pursuing careers in the biological sciences. For this reason, we not only integrate clinical information throughout this textbook and fea- ture it in Clinical Focus boxes but also expand on it in Chapter 16 at the end of the book. As in the First Edition, the placement of some topics in An Introduction to Brain and Behavior is novel relative to traditional treatments. For example, we include brief de- scriptions of brain diseases close to discussions of basic processes that may be associ- ated with the diseases. The integrated coverage of Parkinson’s disease in Chapter 5 is an example. This strategy helps first-time students repeatedly see the close links between what they are learning and real-life problems. The integration of real-life problems, especially behavioral disorders, into every chapter is further accomplished by our many Clinical Focus boxes. These boxes typically describe interesting case studies to highlight spe- cific disorders that are related to a chapter’s content. In this new edition, the range of disorders we cover has expanded to more than 130 disorders, all indexed on page vi, opposite the Contents in Brief. Our capstone Chapter 16 expands on the nature of neuroscience research and the multidisciplinary treatment methods described in the preceding chapters’ coverage of neurological and psychiatric disorders, including a discussion of causes and classifications of abnor- mal behavior. Certainly the most common questions from students who contact us, sometimes years after they have completed our courses, concern the major behavioral disorders that people are likely to encounter in their lifetimes that are reviewed in Chapter 16. Another area of emphasis in this book is questions that relate to the biological basis of behavior. For us, the excitement of neuroscience is in understanding how the brain
  22. 22. Preface xxiexplains what we do, whether it is talking, sleeping, seeing, or learning. Readers willtherefore find nearly as many illustrations about behavior as there are illustrations aboutthe brain. This emphasis on explaining behavior’s biological foundation is another rea-son that we have included both Clinical Focus and Research Focus boxes throughoutthe text. Abundant Chapter PedagogyIn addition to the innovative teaching devices just described, you will find numerousother in-text pedagogical aids. Every chapter begins with a chapter outline and a Focusbox that draws students into the topic by connecting brain and behavior to relevantclinical or research experience. Within the chapters, end-of-section Reviews help stu-dents remember major points, summary illustrations help them visualize or review con-cepts, and terms are defined in the margins to reinforce their importance. Each chapter ends with a Summary, consisting of topical key questions and theiranswers, and a list of Key Terms, which includes the page numbers on which the termsare defined. In contrast to the previous editions, Review Questions and For FurtherThought can be found on the Companion Web Site at www.worthpublishers.com/kolb.Other material on the Web site will broaden students’ understanding of chapter topics. Superb Visual ReinforcementOur most important learning aid, which you can see by simply paging through the book,continues to be an expansive and, we believe, exceptional set of illustrations. Hand inhand with our words, these illustrations describe and illuminate the world of the brain.In response to advice from instructors and readers, important anatomical diagramshave been enlarged to ease perusal. The illustrations in every chapter are consistent and reinforce one another. For ex-ample, throughout the book, we consistently color-code diagrams that illustrate eachaspect of the neuron and those that depict each structural region in the brain. We ofteninclude micrographic images to show what a particular neural structure actually lookslike when viewed through a microscope. You will also find these images on our Power-Point presentations and integrated as labeling exercises in our study guide and testingmaterials. Teaching Through Metaphors, Examples, and PrinciplesWe’ve developed this book in a style that students enjoy because, if a textbook is notenjoyed, it has little chance of teaching well. We heighten students’ interest in the ma-terial they are reading through abundant use of metaphors and examples. Students readabout patients whose brain injuries are sources of insight into brain function, and weexamine car engines, robots, and prehistoric flutes for the same purpose. Frequent com-parative biology examples and representative Comparative Focus boxes help studentsunderstand how much we humans have in common with creatures as far distant fromus as sea slugs. We also facilitate learning by reemphasizing main points and by distilling sets ofprinciples about brain function that can serve as a framework to guide students’ think-ing. Thus, Chapter 2 concludes by introducing ten key principles that explain howthe various parts of the nervous system work together. Chapter 14 summarizes sevenguiding principles of neuroplasticity, a concept fundamental to understandingbrain–behavior relationships. These principles form the basis of many discussionsthroughout the book, and marginal notes remind readers when they encounter theprinciples again.
  23. 23. xxii PREFAC E Big-Picture Emphasis One challenge in writing an introductory book on any topic is deciding what to include and what to exclude. We decided to organize discussions to focus on the bigger picture. A prime example is the discussion of general principles of nervous-system function in- troduced in Chapter 2 and echoed throughout the book. Although any set of princi- ples may be a bit arbitrary, it nevertheless gives students a useful framework for understanding the brain’s activities. Similarly, in Chapters 7 through 16, we tackle behavioral topics in a more general way than most contemporary books do. For instance, in Chapter 12, we revisit the ex- periments and ideas of the 1960s as we try to understand why animals behave in the way they do, after which we consider emotional and motivated behaviors as diverse as eating and anxiety attacks in humans. Another example of our focus on the larger pic- ture is the discussion of learning and memory in Chapter 14, which is presented along- side a discussion of recovery from brain damage. We believe that this broader focus helps students grasp the larger problems that be- havioral neuroscience is all about. Broadening our focus, however, has required us to leave out some details that might be found in other textbooks. But we think that dis- cussions of larger problems and issues in the study of brain and behavior are of greater interest to students who are new to this field and are more likely to be remembered. As in preceding editions, we have been selective in our citation of the truly massive literature on the brain and behavior because we believe that numerous citations can disrupt the flow of a textbook and distract students from the task of mastering what they read. We provide citations to classic works by including the names of the researchers and by mentioning where the research was performed. In areas where there is contro- versy or new breakthroughs, we also include detailed citations to current papers, citing papers from the years 2006 to 2009 when possible. A References section at the end of the book lists all the literature used in developing the book. Supplements Materials of various kinds, for students and instructors and thoroughly checked for accuracy, are available to supplement our book. For Students New! Online Neuroscience Tool Kit The Neuroscience Tool Kit gives instructors and students access to a dozen, high-quality, interactive tutorials covering important brain and nervous-system structures and important functions (e.g., eating, language, move- ment). Clips from the Worth Neuroscience Video Series are integrated into the Tool Kit to provide a real-world context for the material discussed in the tutorials. Revised! An Introduction to Brain and Behavior, Third Edition, Companion Web Site Created by Joe Morrissey, Binghamton University, and available at www.worth- publishers.com/kolb, the companion Web site is an online educational setting for stu- dents that provides a virtual study guide, 24 hours a day, 7 days a week. Best of all, the resources are free and do not require any special access codes or passwords. Tools on the site include chapter outlines and summaries; learning objectives; annotated Web links; interactive flashcards; research exercises; selections from PsychSim 5.0 by Thomas Ludwig, Hope College; and online quizzes with immediate feedback and in- structor notification. For the instructor, the site offers online quizzing (with access to a quiz gradebook for viewing student results), a syllabus posting service, PowerPoint presentation files,
  24. 24. Preface xxiiielectronic versions of illustrations in the book (through Worth Publishers’ Image andLecture Gallery), and links to additional tools including the WebCT E-pack and Black-board course cartridge.Revised! Study Guide Written by Terrence J. Bazzett, State University of New York atGeneseo, the revised Study Guide is carefully crafted, with the use of a variety of en-gaging exercises and tools, to enhance student interest in the material presented in thetextbook. To aid learning and retention, each chapter includes a review of key conceptsand terms, practice tests, short-answer questions, illustrations for identification andlabeling, Internet activities, and crossword puzzles. Students who have completed thetests and exercises can better organize and apply what they have studied.Scientific American Explores the Hidden Mind: A Collector’s Edition On request,this reader is free of charge when packaged with the textbook. In the past decade, wehave learned more about the brain and how it creates the mind than we have in the en-tire preceding century. In a special collector’s edition, Scientific American provides amust-have compilation of updated feature articles that explore and reveal the mysteri-ous inner workings of our minds and brains.Improving the Mind and Brain: A Scientific American Special Issue On request, thisreader is free of charge when packaged with the textbook. This new single-topic issuefrom Scientific American magazine features the latest findings from the most distin-guished researchers in the field.Ronald J. Comer’s Psychological Disorders: A Scientific American Reader On request,this reader is free of charge when packaged with the textbook. Drawn from ScientificAmerican, this full-color collection of articles enhances coverage of the disorders cov-ered in the physiological psychology course. The selections have been hand-picked byWorth author and clinical psychologist Ronald Comer, who provides a preview and dis-cussion questions for each article.Scientific American: Clinical Neuroscience Reader On request, this reader is free ofcharge when packaged with the textbook. Drawn from Scientific American and Scien-tific American Mind, the reader offers 15 full-color articles on cutting-edge science inclinical neuroscience. The selections have been hand-picked by neuroscientists activelyteaching the biological psychology course.For InstructorsRevised! Instructor’s Resources Revised by Debora Baldwin, University of Tennessee,Knoxville, the resources include chapter-by-chapter learning objectives and chapteroverviews, detailed lecture outlines, thorough chapter summaries, chapter key terms,in-class demonstrations and activities, springboard topics for discussion and debate,ideas for research and term-paper projects, homework assignments and exercises, andsuggested readings for students (from journals and periodicals). Course-planning sug-gestions and a guide to videos and Internet resources also are included.Assessment ToolsRevised! Test Bank Prepared by Robert Sainsbury, University of Calgary, the revisedTest Bank includes a battery of more than 1500 multiple-choice and short-answer testquestions as well as diagram exercises tied to the major illustrations in each chapter.Each item is keyed to the page in the textbook on which the answer can be found. The
  25. 25. xxiv PREFAC E wide variety of applied, conceptual, and factual questions have been thoroughly re- viewed and edited for accuracy. Diploma Computerized Test Bank The Test Bank is also available on a dual-platform CD-ROM. Instructors are guided step-by-step through the process of creating a test and can quickly add an unlimited number of questions, edit, scramble, or resequence items, and format a test. The accompanying gradebook enables them to record students’ grades throughout the course and to sort student records and view detailed analyses of test items, curve tests, generate reports, and add weights to grades. The CD-ROM is the ac- cess point for Diploma Online Testing, allowing instructors to create and administer secure exams over a network and over the Internet, as well as Blackboard- and WebCT- formatted versions of each item in the Test Bank. Online Quizzing—Powered by Questionmark This supplement is accessed through the companion Web site at www.worthpublishers.com/kolb. Instructors can easily and securely quiz students online by using prewritten multiple-choice questions for each chapter. Students receive instant feedback and can take the quizzes multiple times. Using the online quiz gradebook, instructors can view results by quiz, student, or question, or they can obtain weekly results by e-mail. Presentation Chapter Illustrations and Outline PowerPoint Slides Available at www.worthpub- lishers.com/kolb, these PowerPoint slides can be either used directly or customized to fit an instructor’s needs. There are two prebuilt, customizable slide sets for each chap- ter of the book—one featuring chapter section headings and the other featuring all chapter figures, tables, and illustrations. Worth Publishers’ Image and Lecture Gallery Available at www.worthpublishers. com/ilg, the Image and Lecture Gallery is a convenient way to access electronic ver- sions of lecture materials. Instructors can browse, search, and download illustrations from every book published by Worth Publishers, as well as prebuilt PowerPoint pre- sentations that contain all chapter illustrations or chapter section headings in text form. Users can also create personal folders on a personalized home page for easy or- ganization of the materials. Video Worth Publishers’ Neuroscience Video Collection Edited by Ronald J. Comer, Prince- ton University, and available on VHS, DVD, and CD-ROM (in MPEG format), this video collection consists of dozens of video segments, each from 1 to 10 minutes in length. Clinical documentaries, television news reports, and archival footage constitute only a small part of the exciting source material for each video. Each segment has been cre- ated to provide illustrations that can help bring a lecture to life, engaging students and enabling them to apply neuroscience theory to the real world. This collection offers powerful and memorable demonstrations such as the links between the brain and be- havior, neuroanatomical animations, cutting-edge neuroscience research, brain assess- ment in action, important historical events, interviews, and a wide sampling of brain phenomena and brain dysfunction. A special cluster of segments reveals the range of research methods used to study the brain. The accompanying Faculty Guide offers a de- scription of each segment so that instructors can make informed decisions about how to best use the videos to enhance their lectures.
  26. 26. Preface xxvThe Brain and Behavior Video Segments Available on VHS, this special collection in-cludes targeted selections from the revised edition of the highly praised Scientific Amer-ican Frontiers series. Hosted by Alan Alda, these video clips provide instructors with anexcellent tool for showing how neuroscience research is conducted. The 10- to 12-minutemodules focus on the work of Steve Sumi, Renée Baillargeon, Car Rosengren, LauraPettito, Steven Pinker, Barbara Rothbaum, Bob Stickgold, Irene Pepperberg, MarcHauser, Linda Bartoshuk, and Michael Gazzaniga.The Mind—Video Teaching Modules, Second Edition Edited by Frank J. Vattano, Col-orado State University, in consultation with Charles Brewer, Furman University, andDavid Myers, in association with WNET, these 35 brief, engaging video clips dramaticallyenhance and illustrate lecture topics and are available on VHS and DVD. This collectionof short clips contains segments on language processing, infant cognitive development,heredity factors in alcoholism, and living without memory (featuring a dramatic inter-view with Clive Wearing). The accompanying Faculty Guide offers descriptions for eachmodule and suggestions for incorporating them into class presentations.The Brain—Video Teaching Modules, Second Edition Edited by Frank J. Vattano,Thomas L. Bennet, and Michelle Butler, all of Colorado State University, and availableon VHS and DVD, this collection is a great source of classroom discussion ideas. It in-cludes 32 short clips that provide vivid examples of brain development, function, dis-orders, and research. Individual segments range from 3 to 15 minutes in length,providing flexibility in highlighting specific topics. The accompanying Faculty Guideoffers descriptions for each module and suggestions for incorporating them into classpresentations.Course Management AidsOnline Course Materials (WebCT and Blackboard) As a service for adopters whouse WebCT or Blackboard course management systems, the various resources for thistextbook are available in the appropriate format for their systems. The files can be cus-tomized to fit specific course needs or they can be used as is. Course outlines, prebuiltquizzes, links, and activities are included, eliminating hours of work for instructors. Formore information, please visit our Web site at www.worthpublishers.com/mediaroomand click “course management.” AcknowledgmentsAs in the past with this text and Fundamentals of Human Neuropsychology, we have aspecial debt to Barbara Brooks, our development editor. She has learned how to extractthe best from each of us by providing a firm guiding hand to our thinking. While wedon’t always initially agree with her (or our wives), we have learned to listen carefullyand discover that she is usually right. Her sense of humor is infectious and her com-mitment to excellence has again left a strong imprint on the entire book. Thank you,Barbara. We must sincerely thank the many people who contributed to the development ofthis edition. The staff at Worth and W. H. Freeman and Company are remarkable andmake the revision a joy to do. We thank our sponsoring editor, Charles Linsmeier, ourlong-time project editor Georgia Lee Hadler, and our production manager Sarah Segal.Our manuscript editors Patricia Zimmerman and Penelope Hull ensured the clarity andconsistency of the text. We also thank Vicki Tomaselli for the inviting and accessibledesign and Ted Szczepanski for coordinating the photo research and the various
  27. 27. xxvi PREFAC E specialists who found photographs and other illustrative materials that we would not have found on our own. We remain indebted to the illustrators at Dragonfly Media and Northeastern Graphic for their excellent work in updating the artwork. Our colleagues, too, helped in the development of this edition. We are especially in- debted to those reviewers who provided extensive comments on selected chapters and illustrations: Chana Akins, University of Kentucky; Michael Anch, Saint Louis Univer- sity; Maura Mitrushina, California State University, Northridge; Paul Wellman, Texas A & M University; and Ilsun White, Morehead State University. New chapters are always a challenge because they are inserted into an existing storyline and must fit in seam- lessly. The new methods chapter had the additional challenge of taking what could read like a seed catalogue and making it engaging to readers. We therefore are indebted to Margaret G. Ruddy, The College of New Jersey, and Ann Voorhies, University of Wash- ington, for providing extensive advice on Chapter 6. We’d also like to thank those that contributed their thoughts to the Second Edition: Barry Anton, University of Puget Sound; R. Bruce Bolster, University of Winnipeg; James Canfield, University of Washington; Edward Castañeda, University of New Mexico; Darragh P. Devine, University of Florida; Kenneth Green, California State University, Long Beach; Eric Jackson, University of New Mexico; Sheri Mizumori, University of Wash- ington; Michael Nelson, University of Missouri, Rolla; Joshua S. Rodefer, University of Iowa; Charlene Wages, Francis Marion University; Doug Wallace, Northern Illinois Uni- versity; Patricia Wallace, Northern Illinois University; and Edie Woods, Madonna Uni- versity. Sheri Mizumuri deserves special mention here for reading the entire manuscript for accuracy and providing fresh ideas that proved invaluable. Finally, we must thank our tolerant wives for putting up with us during the revi- sion of An Introduction to Brain and Behavior immediately after completing the revi- sion of Fundamentals of Human Neuropsychology. We also thank our graduate students, technicians, and postdoctoral fellows, who kept our research programs moving forward when were engaged in revising the book. Bryan Kolb and Ian Q. Whishaw
  28. 28. CHAPTERWilliam West/AFP/Getty Images 1 What Are the Origins of Brain and Behavior? CLINICAL FOCUS 1-1 Living with Brain Injury Evolution of Animals Having Nervous Systems The Chordate Nervous System Neuroscience in the Twenty-First Century Why Study Brain and Behavior? Evolution of the Human Brain and Behavior What Is the Brain? Humans: Members of the Primate Order RESEARCH FOCUS 1-2 Recovering Consciousness Australopithecus: Our Distant Ancestor Gross Anatomy of the Nervous System The First Humans What Is Behavior? Relating Brain Size and Behavior Climate and the Enlarging Hominid Brain Perspectives on Brain and Behavior Why the Hominid Brain Enlarged Aristotle and Mentalism Descartes and Dualism Modern Human Brain Size and Intelligence Fallacies of Human Brain-Size Comparisons COMPARATIVE FOCUS 1-3 The Speaking Brain Culture Darwin and Materialism COMPARATIVE FOCUS 1-4 Evolution and Adaptive Evolution of Brain and Behavior Behavior Origin of Brain Cells and Brains Classification of Life 1

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