Biological Psychology

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1 Biological Psychology An Introduction to Behavioral, Cognitive, and Clinical Neuroscience SEVENTH EDITION S. MARC BREEDLOVE Michigan State University NEIL V. WATSON Simon Fraser University Sinauer Associates, Inc. Publishers Sunderland, Massachusetts

2 Brief Contents Chapter 1 Biological Psychology: Scope and Outlook 1 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 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 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 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 V Emotions and Mental Disorders 455 Chapter 15 Emotions, Aggression, and Stress 457 Chapter 16 Psychopathology: Biological Basis of Behavioral Disorders 491 VI Cognitive Neuroscience 523 Chapter 17 Learning and Memory 525 Chapter 18 Attention and Higher Cognition 561 Chapter 19 Language and Hemispheric Asymmetry 597

3 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 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 I 2 Biological Foundations of Behavior 21 Functional Neuroanatomy: The Nervous System and Behavior 23 A Stimulating Experience 23 The Nervous System Is Composed of Cells 24

4 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 BOX 2.2 Three Customary Orientations for Viewing the Brain and Body 40 The Brain Is Described by Both Structure and Function 43 Specialized Support Systems Protect and Nourish the Brain 47 Brain Imaging Techniques Reveal the Structure and Function of the Living Human Brain 49 BOX 2.3 Isolating Specifi c Brain Activity 51 The Cutting Edge Two Heads Are Better Than One 54 Visual Summary 56 3 Neurophysiology: The Generation, Transmission, and Integration of Neural Signals 59 The Laughing Brain 59 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 Synapses Cause Graded, Local Changes in the Postsynaptic Membrane Potential 71 Synaptic Transmission Requires a Sequence of Events 76 Neurons and Synapses Combine to Make Circuits 82 Gross Electrical Activity of the Human Brain 84 The Cutting Edge Optogenetics: Using Light to Probe Brain Behavior Relationships 87 Visual Summary 88 4 The Chemistry of Behavior: Neurotransmitters and Neuropharmacology 91 The Birth of a Pharmaceutical Problem Child 91 Many Chemical Neurotransmitters Have Been Identified 92 Neurotransmitter Systems Form a Complex Array in the Brain 94 The Effects of a Drug Depend on Its Site of Action and Dose 98 Drugs Affect Each Stage of Neural Conduction and Synaptic Transmission 104 Drugs That Affect the Brain Can Be Divided into Functional Classes 107 Drug Abuse Is Pervasive 116 BOX 4.1 The Terminology of Substance-Related Disorders 117 The Cutting Edge The Needle and the Damage Undone 121 Visual Summary Hormones and the Brain 125 Life-Threatening Lethargy 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 Dwarfi sm 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

5 CONTENTS IX II Evolution and Development of the Nervous System 155 Evolution of the Brain and Behavior 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 BOX 6.1 Why Should We Study Particular Species? 163 BOX 6.2 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 177 Evolution Continues Today 179 The Cutting Edge Are Humans Still Evolving? 181 Visual Summary 183 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 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 Experience Is an Important Influence on Brain Development 209 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 Biological Foundations of Behavior 221 III 8 General Principles of Sensory Processing, Touch, and Pain 223 What s Hot? What s Not? 223 Sensory Processing 223 Sensory Receptor Organs Detect Energy or Substances 224 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

6 X CONTENTS Skin Is a Complex Organ That Contains a Variety of Sensory Receptors 235 The Dorsal Column System Carries Somatosensory Information from the Skin to the Brain 238 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 Hearing, Vestibular Perception, Taste, and Smell 255 No Ear for Music 255 Hearing 255 BOX 9.1 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 Brainstem Auditory Systems Are Specialized for Localizing Sounds 266 The Auditory Cortex Performs Complex Tasks in the Perception of Sound 268 Hearing Loss Is a Major Disorder of the Nervous System 270 Vestibular Perception 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 The Human Tongue Discriminates Five Basic Tastes 276 Chemicals in the Air Elicit Odor Sensations 281 The Cutting Edge More Than a Matter of Taste 287 Visual Summary Vision: From Eye to Brain 291 When Seeing Isn t Seeing 291 The Visual System Extends from the Eye to the Brain 291 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 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 Perception of Visual Motion Is Analyzed by a Special System That Includes Cortical Area V5 319 The Many Cortical Visual Areas Are Organized into Two Major Streams 320 Visual Neuroscience Can Be Applied to Alleviate Some Visual Deficiencies 322 The Cutting Edge Seeing the Light 323 Visual Summary Motor Control and Plasticity 327 What You See Is What You Get 327 The Behavioral View 327 The Control Systems View 329

7 CONTENTS XI The Neuroscience View 330 Movements Are Controlled at Several Nervous System Levels 337 BOX 11.1 Cortical Neurons Can Guide a Robotic Arm 341 Extrapyramidal Systems Also Modulate Motor Commands 345 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 IV Regulation and Behavior 359 Sex: Evolutionary, Hormonal, and Neural Bases Homeostasis: Active Regulation of the Internal Environment 393 Genitals and Gender: What Makes Us Male and Female? 361 Sexual Behavior 361 Reproductive Behavior Can Be Divided into Four Stages 362 The Neural Circuitry of the Brain Regulates Reproductive Behavior 365 Pheromones Guide Reproductive Behavior in Many Species 367 The Hallmark of Human Sexual Behavior Is Diversity 369 For Many Vertebrates, Parental Care Determines Offspring Survival 372 Sexual Differentiation 373 Sex Determination and Sexual Differentiation Occur Early in Development 373 How Should We Define Gender by Genes, Gonads, Genitals, or the Brain? 378 Gonadal Hormones Direct Sexual Differentiation of the Brain and Behavior 378 BOX 12.1 The Paradoxical Sexual Differentiation of the Spotted Hyena 381 Do Fetal Hormones Masculinize Human Behaviors in Adulthood? 386 The Cutting Edge Sex on the Brain 390 Visual Summary 391 A Love-Hate Relationship with Food 393 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 Water Moves between Two Major Body Compartments 400 Two Internal Cues Trigger Thirst 401 Food and Energy Regulation 405 Nutrient Regulation Helps Prepare for Future Needs 405 Insulin Is Crucial for the Regulation of Body Metabolism 408 The Hypothalamus Coordinates Multiple Systems That Control Hunger 409 Obesity Is Difficult to Treat 415 BOX 13.2 Body Fat Stores Are Tightly Regulated, Even after Surgical Removal of Fat 416 Eating Disorders Are Life-Threatening 418 The Cutting Edge A Rumbling in the Belly 419 Visual Summary 421

8 XII CONTENTS 14 Biological Rhythms, Sleep, and Dreaming 423 When Sleep Gets Out of Control 423 Biological Rhythms 423 Many Animals Show Daily Rhythms in Activity 423 The Hypothalamus Houses a Circadian Clock 425 Some Biological Rhythms Are Longer or Shorter than a Day 429 Sleep and Waking 430 Human Sleep Exhibits Different Stages 430 Different Species Provide Clues about the Evolution of Sleep 434 Our Sleep Patterns Change across the Life Span 435 Manipulating Sleep Reveals an Underlying Structure 437 BOX 14.1 Sleep Deprivation Can Be Fatal 438 What Are the Biological Functions of Sleep? 439 At Least Four Interacting Neural Systems Underlie Sleep 443 Sleep Disorders Can Be Serious, Even Life-Threatening 448 The Cutting Edge Can Individual Neurons Be Sleepy? 451 Visual Summary 453 Emotions and Mental Disorders 455 V Emotions, Aggression, and Stress Trouble in Paradise 457 What Are Emotions? 458 Broad Theories of Emotion Emphasize Bodily Responses 458 BOX 15.1 Lie Detector? 461 How Many Emotions Do We Experience? 462 Emotions from the Evolutionary Viewpoint 465 Do Distinct Brain Circuits Mediate Emotions? 468 Neural Circuitry, Hormones, and Synaptic Transmitters Mediate Violence and Aggression 476 Stress Activates Many Bodily Responses 479 Stress and Emotions Affect the Immune System 482 The Cutting Edge Synaptic Changes during Fear Conditioning 487 Visual Summary 489 Psychopathology: Biological Basis of Behavioral Disorders 491 My Lobotomy 491 The Toll of Psychiatric Disorders Is Huge 492 Schizophrenia Is the Major Neurobiological Challenge in Psychiatry 492 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 513 BOX 16.3 Tics, Twitches, and Snorts: The Unusual Character of Tourette s Syndrome 516 Neurosurgery Has Been Used to Treat Psychiatric Disorders 517 Abnormal Prion Proteins Destroy the Brain 518 The Cutting Edge Are Abnormal Eye Movements an Endophenotype for People at Risk for Schizophrenia? 519 Visual Summary 521

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

10 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 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 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 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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