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 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 00_FM_Breedlove.indd III 3/1/13 1:51 PM Brief Contents Chapter 1 PART I Chapter 2 Chapter 3 Chapter 4 Chapter 5 Biological Psychology: Scope and Outlook 1 Biological Foundations of Behavior 21 Functional Neuroanatomy: The Nervous System and Behavior 23 Neurophysiology: The Generation, Transmission, and Integration of Neural Signals 59 The Chemistry of Behavior: Neurotransmitters and Neuropharmacology 91 Hormones and the Brain 125 PART II Chapter 6 Chapter 7 Evolution and Development of the Nervous System 155 Evolution of the Brain and Behavior 157 Life-Span Development of the Brain and Behavior 185 PART III Chapter 8 Chapter 9 Chapter 10 Chapter 11 Biological Foundations of Behavior 221 General Principles of Sensory Processing, Touch, and Pain Hearing, Vestibular Perception, Taste, and Smell 255 Vision: From Eye to Brain 291 Motor Control and Plasticity 327 PART IV Chapter 12 Chapter 13 Chapter 14 Regulation and Behavior 359 Sex: Evolutionary, Hormonal, and Neural Bases 361 Homeostasis: Active Regulation of the Internal Environment Biological Rhythms, Sleep, and Dreaming 423 PART V Chapter 15 Chapter 16 Emotions and Mental Disorders 455 Emotions, Aggression, and Stress 457 Psychopathology: Biological Basis of Behavioral Disorders PART VI Chapter 17 Chapter 18 Chapter 19 Cognitive Neuroscience 523 Learning and Memory 525 Attention and Higher Cognition 561 Language and Hemispheric Asymmetry 223 393 491 597 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 00_FM_Breedlove.indd VI 3/4/13 3:52 PM Contents 1 Biological Psychology: 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 BOX 1.1 3 We Are All Alike, and We Are All Different Three Approaches Relate Brain and Behavior 5 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 2 Biological Foundations of Behavior 21 Functional Neuroanatomy: The Nervous System and Behavior A Stimulating Experience 23 23 The Nervous System Is Composed of Cells 24 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 00_FM_Breedlove.indd VII 3/1/13 1:51 PM VIII CONTENTS BOX 2.1 Neuroanatomical Methods Provide Ways to Make Sense of the Brain 26 The Nervous System Consists of Central and Peripheral Divisions 34 4 The Chemistry of Behavior: Neurotransmitters and Neuropharmacology 91 BOX 2.2 Three Customary Orientations for Viewing the Brain and Body 40 The Birth of a Pharmaceutical Problem Child 91 The Brain Is Described by Both Structure and Function 43 Many Chemical Neurotransmitters Have Been Identified 92 Specialized Support Systems Protect and Nourish the Brain 47 Neurotransmitter Systems Form a Complex Array in the Brain 94 Brain Imaging Techniques Reveal the Structure and Function of the Living Human Brain 49 The Effects of a Drug Depend on Its Site of Action and Dose 98 BOX 2.3 Drugs Affect Each Stage of Neural Conduction and Synaptic Transmission 104 Isolating Specific Brain Activity The Cutting Edge Than One 54 Visual Summary 51 Two Heads Are Better Drugs That Affect the Brain Can Be Divided into Functional Classes 107 56 Drug Abuse Is Pervasive 3 Neurophysiology: The Generation, Transmission, and Integration of Neural Signals 59 The Laughing Brain Electrical Signals Are the Vocabulary of the Nervous System 60 BOX 3.1 Changing the Channel 67 BOX 3.2 Electrical Synapses Work with No Time Delay 71 Synaptic Transmission Requires a Sequence of Events 76 Neurons and Synapses Combine to Make Circuits 82 Gross Electrical Activity of the Human Brain The Cutting Edge Optogenetics: Using Light to Probe Brain–Behavior Relationships 87 88 The Cutting Edge The Needle and the Damage Undone 121 5 123 Hormones and the Brain 125 Life-Threatening Lethargy Synapses Cause Graded, Local Changes in the Postsynaptic Membrane Potential 71 Visual Summary BOX 4.1 The Terminology of Substance-Related Disorders 117 Visual Summary 59 116 84 125 Hormones Have Many Actions in the Body 125 Hormones Have a Variety of Cellular Actions 131 BOX 5.1 Techniques of Modern Behavioral Endocrinology 134 Each Endocrine Gland Secretes Specific Hormones 137 BOX 5.2 Stress and Growth: Psychosocial Dwarfism 143 Hormones Affect Behavior in Many Different Ways 149 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 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 00_FM_Breedlove.indd VIII 3/1/13 1:51 PM IX CONTENTS PART II 6 Evolution and Development of the Nervous System Evolution of the Brain and Behavior 157 We Are Not So Different, Are We? 157 How Did the Enormous Variety of Species Arise on Earth? 158 Why Should We Study Other Species? 162 To Each Its Own Sensory World 165 All Vertebrate Brains Share the Same Basic Structures 167 The Evolution of Vertebrate Brains Reflects Changes in Behavior 169 Many Factors Led to the Rapid Evolution of a Large Cortex in Primates 174 BOX 6.3 Evolutionary Psychology Evolution Continues Today Life-Span Development of the Brain and Behavior 185 Overcoming Blindness 185 Growth and Development of the Brain Are Orderly Processes 185 Development of the Nervous System Can Be Divided into Six Distinct Stages 187 BOX 6.1 Why Should We Study Particular Species? 163 BOX 6.2 7 155 177 179 BOX 7.1 Degeneration and Regeneration of Nervous Tissue 191 BOX 7.2 The Frog Retinotectal System Demonstrates Intrinsic and Extrinsic Factors in Neural Development 200 Developmental Disorders of the Brain Impair Behavior 202 Genes Interact with Experience to Guide Brain Development 204 BOX 7.3 Transgenic and Knockout Mice 205 The Cutting Edge Are Humans Still Evolving? 181 Experience Is an Important Influence on Brain Development 209 Visual Summary The Brain Continues to Change as We Grow Older 212 183 Two Timescales Are Needed to Describe Brain Development 216 The Cutting Edge Genetically Reversing an Inherited Brain Disorder 217 Visual Summary 219 PART III 8 Biological Foundations of Behavior General Principles of Sensory Processing, Touch, and Pain 223 What’s Hot? What’s Not? Sensory Processing 223 223 Sensory Receptor Organs Detect Energy or Substances 224 221 What Type of Stimulus Was That? 225 Sensory Processing Begins in Receptor Cells 226 Sensory Information Processing Is Selective and Analytical 228 BOX 8.1 Synesthesia 235 Touch: Many Sensations Blended Together 235 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 00_FM_Breedlove.indd IX 3/1/13 1:51 PM X CONTENTS Skin Is a Complex Organ That Contains a Variety of Sensory Receptors 235 The Human Tongue Discriminates Five Basic Tastes 276 The Dorsal Column System Carries Somatosensory Information from the Skin to the Brain 238 Chemicals in the Air Elicit Odor Sensations Pain: An Unpleasant but Adaptive Experience 241 Human Pain Can Be Measured 242 Pain Can Be Difficult to Control 247 The Cutting Edge Sticks and Stones… 251 Visual Summary The Cutting Edge Taste 287 Visual Summary 10 More Than a Matter of 288 Vision: From Eye to Brain 291 When Seeing Isn’t Seeing 253 281 291 The Visual System Extends from the Eye to the Brain 291 9 Hearing, Vestibular Perception, Taste, and Smell 255 No Ear for Music Hearing BOX 9.1 255 BOX 10.1 The Basics of Light 294 Neural Signals Travel from the Retina to Several Brain Regions 299 BOX 10.2 Eyes with Lenses Have Evolved in Several Phyla 302 255 The Basics of Sound 256 Each Part of the Ear Performs a Specific Function in Hearing 257 Auditory System Pathways Run from the Brainstem to the Cortex 262 Pitch Information Is Encoded in Two Complementary Ways 264 Neurons at Different Levels of the Visual System Have Very Different Receptive Fields 303 Area V1 Is Organized in Columns 312 Color Vision Depends on Special Channels from the Retinal Cones through Cortical Area V4 314 BOX 10.3 Most Mammalian Species Have Some Color Vision 316 Brainstem Auditory Systems Are Specialized for Localizing Sounds 266 Perception of Visual Motion Is Analyzed by a Special System That Includes Cortical Area V5 319 The Auditory Cortex Performs Complex Tasks in the Perception of Sound 268 The Many Cortical Visual Areas Are Organized into Two Major Streams 320 Hearing Loss Is a Major Disorder of the Nervous System 270 Visual Neuroscience Can Be Applied to Alleviate Some Visual Deficiencies 322 Vestibular Perception The Cutting Edge 273 The Receptor Mechanisms for the Vestibular System Are in the Inner Ear 273 Nerve Fibers from the Vestibular Portion of the Vestibulocochlear Nerve (VIII) Synapse in the Brainstem 275 Some Forms of Vestibular Excitation Produce Motion Sickness 276 The Chemical Senses: Taste and Smell 276 Visual Summary 11 Seeing the Light 323 325 Motor Control and Plasticity 327 What You See Is What You Get The Behavioral View 327 327 The Control Systems View 329 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 00_FM_Breedlove.indd X 3/1/13 1:51 PM CONTENTS The Neuroscience View 330 Movements Are Controlled at Several Nervous System Levels 337 BOX 11.1 Cortical Neurons Can Guide a Robotic Arm 341 XI Disorders of Muscle, Spinal Cord, or Brain Can Disrupt Movement 347 The Cutting Edge Cerebellar Glia Play a Role in Fine Motor Coordination 354 Visual Summary 356 Extrapyramidal Systems Also Modulate Motor Commands 345 PART IV 12 Regulation and Behavior Sex: Evolutionary, Hormonal, and Neural Bases 361 13 Genitals and Gender: What Makes Us Male and Female? 361 Sexual Behavior 359 361 Reproductive Behavior Can Be Divided into Four Stages 362 Homeostasis: Active Regulation of the Internal Environment 393 A Love-Hate Relationship with Food Homeostasis Maintains a Consistent Internal Environment: The Example of Thermoregulation 394 BOX 13.1 Integrated Physiological and Behavioral Thermoregulation Helps Young Animals to Survive 399 The Neural Circuitry of the Brain Regulates Reproductive Behavior 365 Water Moves between Two Major Body Compartments 400 Pheromones Guide Reproductive Behavior in Many Species 367 Two Internal Cues Trigger Thirst 401 The Hallmark of Human Sexual Behavior Is Diversity 369 Food and Energy Regulation For Many Vertebrates, Parental Care Determines Offspring Survival 372 Nutrient Regulation Helps Prepare for Future Needs 405 Sexual Differentiation Insulin Is Crucial for the Regulation of Body Metabolism 408 373 Sex Determination and Sexual Differentiation Occur Early in Development 373 How Should We Define Gender—by Genes, Gonads, Genitals, or the Brain? 378 BOX 12.1 The Paradoxical Sexual Differentiation of the Spotted Hyena 381 Do Fetal Hormones Masculinize Human Behaviors in Adulthood? 386 Visual Summary Sex on the Brain 405 The Hypothalamus Coordinates Multiple Systems That Control Hunger 409 Obesity Is Difficult to Treat Gonadal Hormones Direct Sexual Differentiation of the Brain and Behavior 378 The Cutting Edge 393 415 BOX 13.2 Body Fat Stores Are Tightly Regulated, Even after Surgical Removal of Fat 416 Eating Disorders Are Life-Threatening The Cutting Edge Belly 419 Visual Summary 418 A Rumbling in the 421 390 391 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 00_FM_Breedlove.indd XI 3/1/13 1:51 PM XII CONTENTS 14 Biological Rhythms, Sleep, and Dreaming 423 When Sleep Gets Out of Control Biological Rhythms Our Sleep Patterns Change across the Life Span 435 Manipulating Sleep Reveals an Underlying Structure 437 423 423 BOX 14.1 Sleep Deprivation Can Be Fatal 438 Many Animals Show Daily Rhythms in Activity 423 What Are the Biological Functions of Sleep? 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 and Waking 430 Human Sleep Exhibits Different Stages 439 Sleep Disorders Can Be Serious, Even Life-Threatening 448 The Cutting Edge Can Individual Neurons Be “Sleepy”? 451 430 Different Species Provide Clues about the Evolution of Sleep 434 Visual Summary 453 PART V 15 Emotions and Mental Disorders Emotions, Aggression, and Stress 457 Trouble in Paradise What Are Emotions? 16 457 Lie Detector? Psychopathology: Biological Basis of Behavioral Disorders 491 “My Lobotomy” 458 Emotions from the Evolutionary Viewpoint 462 465 Do Distinct Brain Circuits Mediate Emotions? Neural Circuitry, Hormones, and Synaptic Transmitters Mediate Violence and Aggression 476 Stress Activates Many Bodily Responses 492 Schizophrenia Is the Major Neurobiological Challenge in Psychiatry 492 461 How Many Emotions Do We Experience? 491 The Toll of Psychiatric Disorders Is Huge Broad Theories of Emotion Emphasize Bodily Responses 458 BOX 15.1 455 468 BOX 16.1 Long-Term Effects of Antipsychotic Drugs 502 Mood Disorders Are a Major Psychiatric Category 506 BOX 16.2 The Season to Be Depressed 511 There Are Several Types of Anxiety Disorders 479 513 BOX 16.3 Tics, Twitches, and Snorts: The Unusual Character of Tourette’s Syndrome 516 Stress and Emotions Affect the Immune System 482 Neurosurgery Has Been Used to Treat Psychiatric Disorders 517 The Cutting Edge Synaptic Changes during Fear Conditioning 487 Abnormal Prion Proteins Destroy the Brain Visual Summary 489 518 The Cutting Edge Are Abnormal Eye Movements an Endophenotype for People at Risk for Schizophrenia? 519 Visual Summary 521 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 00_FM_Breedlove.indd XII 3/1/13 1:51 PM XIII CONTENTS PART VI 17 Cognitive Neuroscience Learning and Memory 525 Trapped in the Eternal Now 523 18 525 Attention and Higher Cognition 561 One Thing at a Time Functional Perspectives on Memory 525 There Are Several Kinds of Memory and Learning 526 Attention 561 561 Attention Selects Stimuli for Processing 561 Attention May Be Endogenous or Exogenous Memory Has Temporal Stages: Short, Intermediate, and Long 530 BOX 18.1 Output Successive Processes Capture, Store, and Retrieve Information in the Brain 532 Electrophysiological Techniques Trace Rapid Changes of Brain Activity 570 BOX 17.1 Many Brain Regions Are Involved in Processes of Attention 574 Emotions and Memory 534 Different Brain Regions Process Different Aspects of Memory 536 Neural Mechanisms of Memory Storage 542 565 Reaction-Time Responses, from Input to 566 Two Cortical Networks Collaborate to Govern Attention 578 Disorders Provide Clues about the Organization of Attention 581 Memory Storage Requires Neuronal Remodeling 542 Consciousness 583 Invertebrate Nervous Systems Show Plasticity 545 Consciousness Is the Most Mysterious Property of the Nervous System 583 Synaptic Plasticity Can Be Measured in Simple Hippocampal Circuits 547 The Frontal Lobes Govern Our Most Complex Behavior 588 Some Simple Learning in Mammals Relies on Circuits in the Cerebellum 551 BOX 18.2 In the Adult Brain, Newly Born Neurons May Aid Learning 554 Learning and Memory Change as We Age The Cutting Edge Engram 557 Visual Summary 555 Phineas Gage The Cutting Edge YouTube 593 Visual Summary 590 Putting the You in 595 Artificial Activation of an 559 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 00_FM_Breedlove.indd XIII 3/1/13 1:51 PM XIV CONTENTS 19 Language and Hemispheric Asymmetry 597 Putting a Name to a Face 597 Language Has Both Learned and Innate Components 597 Language Disorders Result from Region-Specific Brain Damage BOX 19.1 Williams Syndrome Offers Clues about Language BOX 19.2 The Wada Test 605 600 605 Competing Models Describe Left-Hemisphere Language Specializations 608 Reading Skills Are Difficult 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 Deficits in Spatial Perception Follow Right-Hemisphere Damage Following Some Injuries, the Brain Can Recover Function BOX 19.3 The Amazing Resilience of a Child’s Brain BOX 19.4 Contact Sports Can Be Costly 624 626 627 628 The Cutting Edge Studying Connectivity in the Living Brain 630 Visual Summary Appendix Glossary 632 A–1 G–1 Illustration Credits References IC–1 R–1 Author Index Subject Index AI–1 SI–1 © Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 00_FM_Breedlove.indd XIV 3/1/13 1:51 PM