Biological Psychology an Introduction to Behavioral, Cognitive, and Clinical Neuroscience SEVENTH EDITION

Biological Psychology An Introduction to Behavioral, Cognitive, and Clinical Neuroscience SEVENTH EDITION

S. MARC BREEDLOVE NEIL V. WATSON Michigan State University Simon Fraser University

Sinauer Associates, Inc. Publishers • Sunderland, Massachusetts

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Chapter 1 Biological Psychology: Scope and Outlook 1 PART I Biological Foundations of Behavior 21 Chapter 2 Functional Neuroanatomy: The Nervous System and Behavior 23 Chapter 3 Neurophysiology: The Generation, Transmission, and Integration of Neural Signals 59 Chapter 4 The Chemistry of Behavior: Neurotransmitters and Neuropharmacology 91 Chapter 5 Hormones and the Brain 125 PART II Evolution and Development of the Nervous System 155 Chapter 6 Evolution of the Brain and Behavior 157 Chapter 7 Life-Span Development of the Brain and Behavior 185 PART III Biological Foundations of Behavior 221 Chapter 8 General Principles of Sensory Processing, Touch, and Pain 223 Chapter 9 Hearing, Vestibular Perception, Taste, and Smell 255 Chapter 10 Vision: From Eye to Brain 291 Chapter 11 Motor Control and Plasticity 327 PART IV Regulation and Behavior 359 Chapter 12 Sex: Evolutionary, Hormonal, and Neural Bases 361 Chapter 13 Homeostasis: Active Regulation of the Internal Environment 393 Chapter 14 Biological Rhythms, Sleep, and Dreaming 423 PART V Emotions and Mental Disorders 455 Chapter 15 Emotions, Aggression, and Stress 457 Chapter 16 Psychopathology: Biological Basis of Behavioral Disorders 491 PART VI Cognitive Neuroscience 523 Chapter 17 Learning and Memory 525 Chapter 18 Attention and Higher Cognition 561 Chapter 19 Language and Hemispheric Asymmetry 597

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Biological Psychology: 1 Scope and Outlook 1 Human or Machine? 1 The Brain Is Full of Surprises 2 What Is Biological Psychology? 2 Five Viewpoints Explore the Biology of Behavior 3 BOX 1.1 We Are All Alike, and We Are All Different 5 Three Approaches Relate Brain and Behavior 6 Neuroplasticity: Behavior Can Change the Brain 7 Biological Psychologists Use Several Levels of Analysis 10 A Preview of the Book: Relations between Brain and Behavior 11 Neuroscience Contributes to Our Understanding of Human Disorders 12 Animal Research Makes Vital Contributions 13 The History of Research on the Brain and Behavior Begins in Antiquity 14 BOX 1.2 Bigger Better? The Case of the Brain and Intelligence 17 The Cutting Edge Neuroscience Is Advancing at a Tremendous Rate 19 Visual Summary 20

PART I Biological Foundations of Behavior 21

Functional Neuroanatomy: 2 The Nervous System and Behavior 23 A Stimulating Experience 23 The Nervous System Is Composed of Cells 24

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BOX 2.1 Neuroanatomical Methods Provide Ways The Chemistry of Behavior: to Make Sense of the Brain 26 Neurotransmitters and The Nervous System Consists of Central and 4 Neuropharmacology 91 Peripheral Divisions 34 BOX 2.2 Three Customary Orientations for Viewing The Birth of a Pharmaceutical Problem the Brain and Body 40 Child 91 The Brain Is Described by Both Structure and Many Chemical Neurotransmitters Have Been Function 43 Identifi ed 92 Specialized Support Systems Protect and Nourish Neurotransmitter Systems Form a Complex Array in the Brain 47 the Brain 94 Brain Imaging Techniques Reveal the Structure and The Effects of a Drug Depend on Its Site of Action Function of the Living Human Brain 49 and Dose 98 BOX 2.3 Isolating Specifi c Brain Activity 51 Drugs Affect Each Stage of Neural Conduction and Synaptic Transmission 104 The Cutting Edge Two Heads Are Better Than One 54 Drugs That Affect the Brain Can Be Divided into Functional Classes 107 Visual Summary 56 Drug Abuse Is Pervasive 116 Neurophysiology: The BOX 4.1 The Terminology of Substance-Related Generation, Transmission, Disorders 117 3 and Integration of Neural The Cutting Edge The Needle and the Signals 59 Damage Undone 121 The Laughing Brain 59 Visual Summary 123 Electrical Signals Are the Vocabulary of the Nervous System 60 Hormones and the BOX 3.1 Changing the Channel 67 5 Brain 125 BOX 3.2 Electrical Synapses Work with No Life-Threatening Lethargy 125 Time Delay 71 Hormones Have Many Actions in the Body 125 Synapses Cause Graded, Local Changes in the Postsynaptic Membrane Potential 71 Hormones Have a Variety of Cellular Actions 131 Synaptic Transmission Requires a Sequence of Events 76 BOX 5.1 Techniques of Modern Behavioral Endocrinology 134 Neurons and Synapses Combine to Make Circuits 82 Each Endocrine Gland Secretes Specifi c Hormones 137 Gross Electrical Activity of the Human Brain 84 BOX 5.2 Stress and Growth: Psychosocial The Cutting Edge Optogenetics: Dwarfi sm 143 Using Light to Probe Brain–Behavior Hormones Affect Behavior in Many Different Relationships 87 Ways 149 Visual Summary 88 Hormonal and Neural Systems Interact to Produce Integrated Responses 150 The Cutting Edge Hormones Made By the Brain, for the Brain 152 Visual Summary 154

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PART II Evolution and Development of the Nervous System 155 Evolution of the Brain Life-Span Development 6 and Behavior 157 of the Brain and 7 Behavior 185 We Are Not So Different, Are We? 157 How Did the Enormous Variety of Species Arise Overcoming Blindness 185 on Earth? 158 Growth and Development of the Brain Are Orderly Why Should We Study Other Species? 162 Processes 185 BOX 6.1 Why Should We Study Particular Development of the Nervous System Can Be Species? 163 Divided into Six Distinct Stages 187 BOX 6.2 To Each Its Own Sensory World 165 BOX 7.1 Degeneration and Regeneration of Nervous Tissue 191 All Vertebrate Brains Share the Same Basic Structures 167 BOX 7.2 The Frog Retinotectal System Demonstrates Intrinsic and Extrinsic Factors The Evolution of Vertebrate Brains Reﬂ ects in Neural Development 200 Changes in Behavior 169 Developmental Disorders of the Brain Impair Many Factors Led to the Rapid Evolution of a Behavior 202 Large Cortex in Primates 174 Genes Interact with Experience to Guide Brain BOX 6.3 Evolutionary Psychology 177 Development 204 Evolution Continues Today 179 BOX 7.3 Transgenic and Knockout Mice 205 The Cutting Edge Are Humans Experience Is an Important Inﬂ uence on Brain Still Evolving? 181 Development 209 Visual Summary 183 The Brain Continues to Change as We Grow Older 212 Two Timescales Are Needed to Describe Brain Development 216 The Cutting Edge Genetically Reversing an Inherited Brain Disorder 217 Visual Summary 219

PART III Biological Foundations of Behavior 221 General Principles of What Type of Stimulus Was That? 225 Sensory Processing, Sensory Processing Begins in Receptor Cells 226 8 Touch, and Pain 223 Sensory Information Processing Is Selective and Analytical 228 What’s Hot? What’s Not? 223 BOX 8.1 Synesthesia 235 Sensory Processing 223 Touch: Many Sensations Blended Sensory Receptor Organs Detect Energy or Together 235 Substances 224

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Skin Is a Complex Organ That Contains a Variety of The Human Tongue Discriminates Five Basic Sensory Receptors 235 Tastes 276 The Dorsal Column System Carries Somatosensory Chemicals in the Air Elicit Odor Sensations 281 Information from the Skin to the Brain 238 The Cutting Edge More Than a Matter of Pain: An Unpleasant but Adaptive Taste 287 Experience 241 Visual Summary 288 Human Pain Can Be Measured 242 Pain Can Be Diffi cult to Control 247 Vision: From Eye to The Cutting Edge Sticks and Brain 291 Stones… 251 10 When Seeing Isn’t Seeing 291 Visual Summary 253 The Visual System Extends from the Eye to the Brain 291 Hearing, Vestibular BOX 10.1 The Basics of Light 294 Perception, Taste, and 9 Smell 255 Neural Signals Travel from the Retina to Several Brain Regions 299 No Ear for Music 255 BOX 10.2 Eyes with Lenses Have Evolved in Several Phyla 302 Hearing 255 Neurons at Different Levels of the Visual System BOX 9.1 The Basics of Sound 256 Have Very Different Receptive Fields 303 Each Part of the Ear Performs a Specifi c Area V1 Is Organized in Columns 312 Function in Hearing 257 Color Vision Depends on Special Channels from the Auditory System Pathways Run from the Retinal Cones through Cortical Area V4 314 Brainstem to the Cortex 262 BOX 10.3 Most Mammalian Species Have Some Pitch Information Is Encoded in Two Color Vision 316 Complementary Ways 264 Perception of Visual Motion Is Analyzed by a Brainstem Auditory Systems Are Specialized Special System That Includes Cortical Area for Localizing Sounds 266 V5 319 The Auditory Cortex Performs Complex Tasks The Many Cortical Visual Areas Are Organized into in the Perception of Sound 268 Two Major Streams 320 Hearing Loss Is a Major Disorder of the Nervous Visual Neuroscience Can Be Applied to Alleviate System 270 Some Visual Defi ciencies 322 Vestibular Perception 273 The Cutting Edge Seeing the Light 323 The Receptor Mechanisms for the Vestibular Visual Summary 325 System Are in the Inner Ear 273 Nerve Fibers from the Vestibular Portion of the Vestibulocochlear Nerve (VIII) Synapse in the Motor Control and Brainstem 275 11 Plasticity 327 Some Forms of Vestibular Excitation Produce What You See Is What You Get 327 Motion Sickness 276 The Behavioral View 327 The Chemical Senses: Taste and Smell 276 The Control Systems View 329

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The Neuroscience View 330 Disorders of Muscle, Spinal Cord, or Brain Can Disrupt Movement 347 Movements Are Controlled at Several Nervous System Levels 337 The Cutting Edge Cerebellar Glia Play a BOX 11.1 Cortical Neurons Can Guide a Robotic Role in Fine Motor Coordination 354 Arm 341 Visual Summary 356 Extrapyramidal Systems Also Modulate Motor Commands 345

PART IV Regulation and Behavior 359 Sex: Evolutionary, Homeostasis: Active Hormonal, and Neural Regulation of the Internal 12 Bases 361 13 Environment 393 Genitals and Gender: What Makes Us Male A Love-Hate Relationship with Food 393 and Female? 361 Homeostasis Maintains a Consistent Sexual Behavior 361 Internal Environment: The Example of Thermoregulation 394 Reproductive Behavior Can Be Divided into Four Stages 362 BOX 13.1 Integrated Physiological and Behavioral Thermoregulation Helps Young Animals to The Neural Circuitry of the Brain Regulates Survive 399 Reproductive Behavior 365 Water Moves between Two Major Body Pheromones Guide Reproductive Behavior in Compartments 400 Many Species 367 Two Internal Cues Trigger Thirst 401 The Hallmark of Human Sexual Behavior Is Diversity 369 Food and Energy Regulation 405 For Many Vertebrates, Parental Care Determines Nutrient Regulation Helps Prepare for Offspring Survival 372 Future Needs 405 Sexual Differentiation 373 Insulin Is Crucial for the Regulation of Body Metabolism 408 Sex Determination and Sexual Differentiation Occur Early in Development 373 The Hypothalamus Coordinates Multiple Systems That Control Hunger 409 How Should We Deﬁ ne Gender—by Genes, Gonads, Genitals, or the Brain? 378 Obesity Is Difﬁ cult to Treat 415 Gonadal Hormones Direct Sexual Differentiation of BOX 13.2 Body Fat Stores Are Tightly Regulated, Even after Surgical Removal of Fat 416 the Brain and Behavior 378 Eating Disorders Are Life-Threatening 418 BOX 12.1 The Paradoxical Sexual Differentiation of the Spotted Hyena 381 The Cutting Edge A Rumbling in the Do Fetal Hormones Masculinize Human Behaviors Belly 419 in Adulthood? 386 Visual Summary 421 The Cutting Edge Sex on the Brain 390 Visual Summary 391

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Biological Rhythms, Sleep, Our Sleep Patterns Change across the Life 14 and Dreaming 423 Span 435 Manipulating Sleep Reveals an Underlying When Sleep Gets Out of Control 423 Structure 437 Biological Rhythms 423 BOX 14.1 Sleep Deprivation Can Be Fatal 438 Many Animals Show Daily Rhythms in Activity 423 What Are the Biological Functions of Sleep? 439 The Hypothalamus Houses a Circadian Clock 425 At Least Four Interacting Neural Systems Underlie Sleep 443 Some Biological Rhythms Are Longer or Shorter than a Day 429 Sleep Disorders Can Be Serious, Even Life-Threatening 448 Sleep and Waking 430 The Cutting Edge Can Individual Neurons Human Sleep Exhibits Different Stages 430 Be “Sleepy”? 451 Different Species Provide Clues about the Visual Summary 453 Evolution of Sleep 434

PART V Emotions and Mental Disorders 455 Emotions, Aggression, Psychopathology: Biological 15 and Stress 457 Basis of Behavioral 16 Disorders 491 Trouble in Paradise 457 What Are Emotions? 458 “My Lobotomy” 491 Broad Theories of Emotion Emphasize Bodily The Toll of Psychiatric Disorders Is Huge 492 Responses 458 Schizophrenia Is the Major Neurobiological BOX 15.1 Lie Detector? 461 Challenge in Psychiatry 492 How Many Emotions Do We Experience? 462 BOX 16.1 Long-Term Effects of Antipsychotic Drugs 502 Emotions from the Evolutionary Viewpoint 465 Mood Disorders Are a Major Psychiatric Do Distinct Brain Circuits Mediate Emotions? 468 Category 506 Neural Circuitry, Hormones, and Synaptic BOX 16.2 The Season to Be Depressed 511 Transmitters Mediate Violence and There Are Several Types of Anxiety Disorders 513 Aggression 476 BOX 16.3 Tics, Twitches, and Snorts: The Unusual Stress Activates Many Bodily Responses 479 Character of Tourette’s Syndrome 516 Stress and Emotions Affect the Immune Neurosurgery Has Been Used to Treat Psychiatric System 482 Disorders 517 The Cutting Edge Synaptic Changes Abnormal Prion Proteins Destroy the Brain 518 during Fear Conditioning 487 The Cutting Edge Are Abnormal Eye Visual Summary 489 Movements an Endophenotype for People at Risk for Schizophrenia? 519 Visual Summary 521

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PART VI Cognitive Neuroscience 523 Learning and Attention and Higher 17 Memory 525 18 Cognition 561 Trapped in the Eternal Now 525 One Thing at a Time 561 Functional Perspectives on Memory 525 Attention 561 There Are Several Kinds of Memory and Attention Selects Stimuli for Processing 561 Learning 526 Attention May Be Endogenous or Exogenous 565 Memory Has Temporal Stages: Short, Intermediate, BOX 18.1 Reaction-Time Responses, from Input to and Long 530 Output 566 Successive Processes Capture, Store, and Retrieve Electrophysiological Techniques Trace Rapid Information in the Brain 532 Changes of Brain Activity 570 BOX 17.1 Emotions and Memory 534 Many Brain Regions Are Involved in Processes Different Brain Regions Process Different Aspects of Attention 574 of Memory 536 Two Cortical Networks Collaborate to Govern Neural Mechanisms of Memory Attention 578 Storage 542 Disorders Provide Clues about the Organization Memory Storage Requires Neuronal of Attention 581 Remodeling 542 Consciousness 583 Invertebrate Nervous Systems Show Consciousness Is the Most Mysterious Property Plasticity 545 of the Nervous System 583 Synaptic Plasticity Can Be Measured in Simple The Frontal Lobes Govern Our Most Complex Hippocampal Circuits 547 Behavior 588 Some Simple Learning in Mammals Relies on BOX 18.2 Phineas Gage 590 Circuits in the Cerebellum 551 The Cutting Edge Putting the You in In the Adult Brain, Newly Born Neurons May Aid YouTube 593 Learning 554 Visual Summary 595 Learning and Memory Change as We Age 555 The Cutting Edge Artiﬁ cial Activation of an Engram 557 Visual Summary 559

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Language and Hemispheric 19 Asymmetry 597 Putting a Name to a Face 597 Language Has Both Learned and Innate Components 597 Language Disorders Result from Region-Specifi c Brain Damage 605 BOX 19.1 Williams Syndrome Offers Clues about Language 600 BOX 19.2 The Wada Test 605 Competing Models Describe Left-Hemisphere Language Specializations 608 Reading Skills Are Diffi cult to Acquire and Frequently Impaired 611 Brain Stimulation Provides Information about the Organization of Language in the Brain 614 Functional Neuroimaging Tracks Activity in the Brain’s Language Zones 616 The Left Brain Is Different from the Right Brain 618 Defi cits in Spatial Perception Follow Right-Hemisphere Damage 624 Following Some Injuries, the Brain Can Recover Function 626 BOX 19.3 The Amazing Resilience of a Child’s Brain 627 BOX 19.4 Contact Sports Can Be Costly 628 The Cutting Edge Studying Connectivity in the Living Brain 630 Visual Summary 632

Appendix A–1 Glossary G–1 Illustration Credits IC–1 References R–1 Author Index AI–1 Subject Index SI–1

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