INTERNATIONAL EDITION NEIL R. CARLSONSIXTH EDITION Foundations of Physiological Psychology University of Massachusetts, Amherst+s Editor: Kelly May Marketing Manager: Karen N; ‘Composition and Prepress Buyer: Linda Cox Cover Administrator: Lincla Knowles Editorial-Production Coordinator: Mary Beth Finch, Copyright © 2005, Inwernet: www.ablongman.com All rights reserved. No part of the ms reproduced or wilized in any form or by any m including photocopying, recording, or by any without iritten permission from the copyright owner, cans, electro ‘To obtain permission(s) to use to Allyn and Bacon, Permissions Depat fr fax your request 10 617-848-7320, 75 Arlington Bens have el 1 the time Web si |. Also the ation is gathe scription of URLs ‘corrected in subsequent editions Library of Congress Cataloging in-Publication Data Carlson, Neil R al protected by this copy jc or mechanical, cdl and published, son n result in typographical errors. The publisher would appreciate notification where these occur so th Photo Researcher: Helane M Editorial Production Service: Barbara 1 berg, Copseditor: Barbara Willette ator: Jay Ales Electronic Composit en ‘Omegatype Typography, 2002, 1999, 1995,1992, 1988 Pearson Education, Ine information storage and retrieval system, erial from this work, please submit a writen request treet, Boston, MA 02116 they may be Foundations of physiological psychology / Neil R. Carlson.—6th ed. p. em Includes bibliog: ISBN 0.205-197255 L. Psychophysiology. 1 QP360.6 ical references and index. Title Photo Credits: p. 1, Martin Harvev/Peter Arnold, Inc. p-2, © Rachel Epstein, PhotoEdit; p. 6, photo courtesy of Neil R. Carlson; p. 17, © Michael Schwarz/The Image Works; p. 26, fim Karageorge /Taxi/© Gettylmages, 10, © Clay Wiseman / Animals Animals; p. 51, © R. 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Dactnm The Image Works p. 311, © Robert Brenner/ PhotoFdit; p. 821, Alexandra Grablewski /BoodPix/ Get © Laswvence Migdale /Stone; p. 349, © Earl Dotter; p 356, Art Montes de Oca/Taxi/O Geuylmages, Inc. p. 371, © Elizabeth Grews/The Image Works; p. 376, De Boissary/Taxi/ © GeuyImages, Inc: p. 885, Elyse Levin / Image Bank/© Genylmage, lc p. 398, Hide Fup fe Bank/© Geutylmages, Inc: p. 401, © Spencer or 1/ Photokdit; p. 416, © B, Daemmrich/The Image Works; p. 422, © Cameramann/The Image Works; p. 132, © James Shaffer /PhotoFdit; p. 438, © Paul Conklin, PhotoEdit; p. 444, © Ellen Senisi/The Image Works: 60, Color Day Production /Image Bank/ © Gettylmages, p 467, © Robert Brennet/PhotoFd ©1993 Bric Nelson / woek Photo: p. 484, © Photo Rese p. 495, Kamil Vojnar © Geutylmages, Inc. reout/Momatinn, © Cindy Charles/Phowodit; p. 5! Alistock/© Gettylmages, Inc‘in memory of Paul Wayne JohnsonBRIEF CONTENTS 1. origins of physiological Psychology 1 2 structure and Functions of Cells of the Nervous System 26 3. structure of the Nervous System 61 4 Psychopharmacology 94 5 methods and strategies of Research 122 6 vi 7 audition, the Body Senses, and the Chemical Senses 188 mn 154 8& steep and Biological Rhythms 226 QD Reproductive Behavior 260 10 Emotion 292 11 ingestive Behavior 321 12 Learning and Memory 356 13> Human communication 398 14 neurotogical Disorders 432 15, schizophrenia, Affective Disorders, and Anxiety Disorders 460 16 autistic, attenti -Deficit/Hyperactivity, Stress, and Substance Abuse Disorders 495Preface sib Origins of Physiological Psychology 1 PROLOGUE: fen’ Inspiration 2 Understanding Human Consciousness: A Physiological Approach 3 Split Brains 3 LINTERIM SUMMARY 5 ‘The Nature of Physiological Psychology 6 The Goals of Rese Biological Roots of Physiological Psychology 7 MLINTERIM SUMMARY 10 Natural Selection and Evaluation 10 Functionalism and the Inheritance of Traits 11 Fyolution of the Hun wn Species 13 1s Feolution of Large Brat MLINTERIM SUMMARY 16 Ethical Issues in Research with Animals 17 Careers in Neuroscience 19 MLINTERIM SUMMARY Strategies for Learning 21 EPILOGUE: Model of Brain Functions 23 Key Concepts 24 Suggested Readings 24 Suggested Web sites 24 SB oh ns Structure and Functions of Cells of the Nervous System 26 = PROLOGUE: Unresponsive Muscles 27 Calls of the Nervous System 28 Neurons 28 Supporting Cells 3 The Blood-Brain Barrier 85 MLINTERIM SUMMARY 35 ‘Communication Within a Neuron N Measuring Hlectrical Potentials of Asons 38 21 Com An Overview 36 Balance of Two Forces 41 The Membrane Potent The Aetion Povential 43 Concluetion of the Action Potential 45 INTERIM SUMMARY 47 Communication Between Neurons 48 Structure of Semapses 48. Release of the Neurotransmitter 50) Act om of Receptors 50 Postsynaptie Potentials 51 Ter tion of Postsynaptic Pote Effects of Postsynaptic Poten Autorece Nonsynaptic Communication: Newromodulators and Hormones INTERIM SUMMARY 56 EPILOGUE: Myasthenia Gravis 57 Key Concepts 59 Suggested Readings 59 Suggested Web Sites 59 SIRT Structure of the Nervous System 61 | PROLOGUE: TheLeit Is Gone 62 Basic Features of the Nervous System 62 An Overview 64 nd Produc Cerebrospinal Fluid 66contents HLINTERIM SUMMARY 68 “The Central Nervous System 68 Development of the Central Nervous System 68 The Forebs The Mid The Hinds The Spinal Gord 8t interim summary 86 ‘The Peripheral Nervous System 86 se Cranial Nerves 86 rves 87 he Autonomic Nervous Syst INTERIM SUMMARY 91 89 EPILOGUE: Unilateral Neglect 91 Key Concepts 92 Suggested Readings 93 Suggested Web Sites 93 aie Se Psychopharmacology 94 PROLOGUE: A Contaminated Drug 95 Principles of Psychopharmacology 95 Pharin kinetics 96 Drug Effectiveness 98 Effects of Repeated Administation Placebo Eifeets 100 MLINTERIM SUMMARY 101 Sites of Drug Action 101 Effects on Production of Newt 1 Release of Effects on Receptors 108 Effects on Storage wz Neuroiransmitters 104 MLINTERIM SUMMARY _ 105 ‘Neurotransmitters and Neuromodulators 105 Acetylcholine 106 The Monosmines 108 Amino Acids 113 Peptides 116 Lipids 117 Nucleosides 117 wvuswablongman.com/earlsoné= Soluble Gases 118 MLINTERIM SUMMARY 118 EPILOGUE: Helpful Hints from a Tragedy 120 Key Concepts 120 Suggested Readings 121 Suggested Web Sites 121 Methods and Strategies of Research 122 PROLOGUE: Heart Repaired, Brain Damaged 123, Experimental Ablatio Evahuatin Brain Dam Producing Brain Lesions 125 Stereo je Surgery 126 Histological Methods 128 Tracing N sal Connections 130 a Bs acy of the Living F 1s MLINTERIM SUMMARY 136 Recording and Stimulating Neural Activity 136 Recording Newr Activity 136 Recording the Brain's Metabolic and Symapii Activiey 130 ‘Seer Jewral Activity 142 Measuring the B ons ML Behavioral Effects of Electrical Brain Stimulation 144 WLINTERIM SUMMARY 145 Neurochemical Methods 146 Finding Neurons That Produce Particular Newrochemicals 117 Localizing Particular Receptors M48 MLINTERIM SUMMARY 149 Genetic Methods 150 “Twin Studies 150 Adoption Studies EPILOGUE: Watch the Brain Waves 152 Key Concepts 152 Suggested Readings 153 Suggested Web Sites 153Vision 154 | PROLOGUE: Seeing with His Hands 155 ‘The Stimulus 156 Anatomy of the Visual System 156 The Exes Photoreeeptors 164 Connections Beaseen Eye and Brain 161 MLINTERIM SUMMARY. 162 ‘Coding of Visual Information in the Retina 163 g of Lightand Dark 163 Coding of Color 164 MLINTERIM SUMMARY 168 Analysis of Visual Information: Role of the Striate Cortex 168 Anatomy of the Strfate C 168 Orientation and Movement 169 “ey 170 y 7 Spatial Frequ Retinal Disp Modular Organization of the Strate Cortex 172 MLINTERIM SUMMARY 173 Analysis of Visual Information: Role of the Visual Association Cortex 173 Two Streams of Visual Analysis 173 Perception of Color 175 Analysis of Form 176 nt 180 Location 182 Perception of Mow Perception of Sp MLINTERIM SUMMARY 183 EPILOGUE: Case Studies 185 Key Concepts 186 Suggested Readings 186 Suggested Web Sites 187 Audition, the Body Senses, and the Chemical Senses 188 PROLOGUE: Allin Her Head? 189 Contents Audition 189 The Stimulus 190 Anatomy of the Ear 190 Auditory Hair Cells and the Transduction ‘of Auditory Information 193. ‘The Auditory Pathway 194 Perception of Pte 196 Pereeption of Timbre 199) Perception of Spatial Location 200) Functions of the iory System 201 MLINTERIM SUMMARY 202 Vestibular System 203 Anatomy of the Vestibular Apparatus 203 ‘The Receptor Cells 204 The Vestibular Pathway 205 MLINTERIM SUMMARY 208 Somatosenses 206 Thes Anatoniy of the Skin and Ls Receptive Organs 207 Perception of Cx ali cous Stimulation 208 0 The Somatosensory Pathways Perception of Pain 211 INTERIM. SUMMARY._213 Gustation 214 The Stimuli 2 Anatomy of the Taste Buds and Gustatory Cells 214 Perception of Gustatory Information 215 ‘The Gustatory Pathway MLINTERIM SUMMARY 218 Olfaction 219 “The Stimulus 219 Anatomy of the Olfactory Apparatus 219 ransduction of Olfietory Infor Perception of Specific Odors & MLINTERIM SUMMARY 222 EPILOGUE: Natural Analgesia 223 Key Concepts 224 Suggested Readings 225 Suggested Web Sites 225,Contents ene SS Sleep and Biological Rhythms 226 | PROLOGUE: Waking Nightmares 227 A Physiological and Behavioral Description of Sleep 227 MLINTERIM SUMMARY 231 Disorders of Sleep 231 231 Narcolepsy 2 REM Steep Behavior Disorder 234 Problems Aswciated ist SlowWave Sleep 235, MLINTERIM SUMMARY 235 Why Do We Sleep? 236 Functions of Sow Wine Sleep 236 Functions of REM Sleep 258 MLINTERIM SUMMARY. 240 Physiological Mechanisms of Sleep and Waking 240 Chemical Control of Sleep 240 Newal Control of Arousal 241 Neural Conta of Slow-Wave Ste Neutal Control of REM Sleep INTERIM suMMARY 250 Biological Clocks 251 Gread 5 Rhythmsand Zeiigebers Role of the Suj Control of Seaso ‘and Melatonin wlian Rhythms: Shift Work asmatie Nuelews 1 Rhythms: The Pineal Gland HLINTERIM SUMMARY 257 EPILOGUE: Functions of Dreams 258 Key Concepts 258 Suggested Readings 259 Suggested Web Sites 259 Reproductive Behavior 260 = PROLOGUE: From Boy 0 Gi! 261 ‘Sexual Development 261 Production of Gametes and Fertilization 261 www ablongman.com/earisonGe Development ofthe Sex Organs 262 Sexual Maturation 265 MLINTERIM SUMMARY. 266 Hormonal Control of Sexual Behavior 268 e Rep Hormonal Control of Fe Cycles 268 oductive Hormonal Control of Sex Laboratory Animals 26 Behavior of iational Effects of Androgens on Beh Etfects of Pheromones 271 Human Sexual Behavior Sextial O ion 276 MLINTERIM SUMMARY 280 Neural Control of Sexual Behavior 281 Males 281 Females 289 INTERIM SUMMARY 284 Parental Behavior 285 Maternal Behavior of Rodents 2 Hormonal Contiol of Maternal Behavior 2 Neural Control of Maternal Behavior 288 ernal Behavior 288 MLINTERIM SUMMARY 289 Neural Control of P EPILOGUE: From Boy to Girl and Back Again 289 Key Concepts 290, Suggested Readings 291 Suggested Web Sites 291 TER Emotion 292 | PROLOGUE: [niellect and Emotion 293 Emotions as Response Patterns 293 Fear Hormonal Control of Aggressive Behavior | 303 INTERIM SUMMARY 307 Communication of Emotions 309 Facial Expression of Emotions: Innate Responses 309) Neural Basis of dhe Communi of Emotions: Recognition S10Neural Basis of the Communication of Emotions Expression 313 INTERIM suMMARY 316 Feelings of Emotions 316 The ge Theory 316 INTERIM SUMMARY 318 EPILOGUE: Mi.\, Revisted 318 Key Concopts 319 Suggested Readings 320 Suggested Web sites 320 Ingestive Behavior 321 PROLOGUE: Not Her Fault? 322 Physiological Regulatory Mechanisms 323 Drinking 323 Some Faets About Fluid Balance 323, Two Types of Thitst 3 Neural Mechanisms of Thirst 327 INTERIM SUMMARY. 328 Eating and Metabolism 329 MLINTERIM SUMMARY 332 What Starts a Meal? 932 and Environmental Factors 332 Physiological Hunger Signals 333, MLINTERIM SUMMARY 334 ‘What Stopsa Meal? 3: Gastric Factors 835 336 Liver Factors 387 Satiety During the Absoptive Ph Role of Inst Brain Stem 339 Hypothalamus 310 IM SUMMARY 344 Contents Eating Disorders 345 Obesity 5 Anorexia Nervosa/Bulimia Nervosa 348 INTERIM SUMMARY 351 EPILOGUE: Treatment of Obesity 353 Key Concepts 354 Suggested Readings 354 Suggested Web Sites 355 Learning and Memory 356 PROLOGUE: Every Day Alone 357 ‘The Nature of Learning 357 ‘INTERIM SUMMARY 361 Learning and Synaptic Plasticity 361 Induction of Role of NMDA Receptors 363 ng:Term Potentiation 302 Mechanisms of Synaptic Plasticity: 365, Long-Term Depression 370 MLINTERIM SUMMARY 370 Perceptual Learning 371 MLINTERIM SUMMARY 373 Classical Conditioning 373 INTERIM. SUMMARY 375 Instrumental Conditioning 875 Role of the Basal Ganglia 875 Reinfore MLINTERIM SUMMARY 381 Relational Learning Human Anterograde Basic Deseription 38 Spared Learning Abilities 384 Declarative and Nondeclarative Memories 385 Anatomy of Anterograde Amnesia 885 Anterograde Amnesia Failure of Relational Learning 389 Role th Spi of Hippocampal Memory 389)Contents Relational Learning in Labormory Animals 390 MLINTERIM SUMMARY 395 1 EPILOGUE: What, Exactly, Does the Hippocampus Do? 396 Key Concepts 396 Suggested Reat Human Communication 398 PROLOGUE: Can't Hear Words 399, Speech Production and Comprehension: Brain Mechanisms 399 Lateralization 100 fon 401 Speech Comprehension 405 The Bilingual Brain 415 Prosoxly: Rhythm, Tone and Emphasis in Speech 416 MLINTERIM. SUMMARY. 417 Disorders of Reading and Writing 419 Relationship to Aphasia 419 Pare Alexia 419 Speech Produc Toward an Understanding of Reading 422 ing of Writing 424 Developmental Dystexias 126 MLINTERIM SUMMARY 428 1 EPILOGUE: Analyss of Speech Sounds 429 Key Concepts 430 Suggested Readings 430 Suggested Web Sites 431 ‘Toward an Underss, Neurological Disorders 432 PROLOGUE: It Started with Her Foot 433 Tumors 433 Seizure Disorders 435 Cerebrovascular Accidents 438 MLINTERIM SUMMARY 449 wwrwablongman.com/carlsonse Disorders of Development 441 Toxie Chemicals 41 Inherited M Down Syndr MLINTERIM SUMMARY. 444 Degenerative Disorders 444 abolie Disorders 442 443 jmnissible Spongiform Eneephalopathies 445 Parkinson's Discase 46 Huntington's Disease 449 Aleheimer’s Disease 450 Muiliple Sclerosis 454 MLINTERIM SUMMARY 454 Disorders Caused by Infections Diseases 455 MLINTERIM SUMMARY. 457 EPILOGUE: Seizute Surgery 457 Key Concepts 458 Suggested Readings 458 Suggested Web Sites 459 Schizophrenia, Affective Disorders, and Anxiety Disorders 460 fm PROLOGUE: Sie Effects 461 Schizophrenia 462 Description 462 Hertabily 468 Pharmacology of Schizophrenia: The Do Hypothesis 463 166, Schieophrenia ss a Neurological Disorder MLINTERIM SUMMARY 473 Major Affective Disorders 475 Description 475 Heritability Physiological Treatments 476 The Role of Monoamine 479 Evidence for Brain Abnormalities 480 Role of Grcadian Rhythms 481 MLINTERIM SUMMARY 484 Anxiety Disorders 485 Panic Disorder 4185 Obsessive Compubive Disorder 487MLINTERIM SUMMARY 491 EPILOGUE: Tardive Dyskinesia 492 Key Concepts 493 Suggested Readings 493 Suggested Web Sites 494 Autistic, Attention-Deficit/Hyperactivity, Stress, and Substance Abuse Disorders 495 fm PROLOGUE: A Sudden Crving 436 er 406 16 Posie Cases. 408 ‘MLINTERIM SUMMARY 500 Atention-Defict Hyperactivity Disorders 500 Deseription 500 Possible Causes 501 Stress Disorders 502 Physiology of the Stress Response 503 Contents Health Eifeets of Long Te Posttraumatic Suess Disorder The Coping Response 508 Suress and Infcetious Diseawes MLINTERIM SUMMARY 510 Substance Abuse Disorders 511 What Is Addiction: Common Features of Addiction 514 Commonly Abused Drugs 517 Heredity and Deng Abuse 52 ‘Therapy for Drug Abuse 525 MLINTERIM SUMMARY 527 EPILOGUE: Classically Conditioned Craving 529 Key Concepts 530 Suggested Readings 530 Suggested Web Sites 530 References 531 Name Index 558 Subject Index 563When I was a box, [took apart alarm clocks, digs, my mother's sewing machine A I my life [ have wanted to know how things work, nd other teresting gadgets, 10 sce what was inside, Much to my parents’ relief, Loutgrew that habit (or at least got bet ter at putting things back together), but my euriosity is, still with me. Since my college days, I have been trying to find out all Tean about the wo ticate piece of mach wwe know of the human The field of neuroscience rest and productive one today. A larg are trying (o understand the physiology of beh nd more advanced methods, yielding more and more interesting results, Their findings provide me ‘with much to write about. [admire their dedication and, hard work, and I thank them for gi say, Without the book. Twwrote the first edition of this book at the request of my colleagues who teach the course, and who wanted a briefer version of Physiology of Behavior with more em- phasis on research related to hum: this book is concerned with foundations: the history of the field, the struc roanatomy, psychoph: ings of the most Wg me something 0 efforts I could not have written this he sensory systems, The third part deals with what might be called “motivated” behavior: sleep, reprod tion, and ingestion, The fourth part d and with verbal communication. The final part deals, with neurological and mental disorders. search, The second part is concerned with input Is with le Content Changes teresting research results to describe in my book. The problem is always to inelude the interesting new mater ial without letting the length of the book get out of hand. Like the previous editions, this edition contains a chapter on neurological disorders, whieh is not found in, Physiology of Behavior The following list includes some of the information| that is nev to this edition: ‘+ Funetional imaging studies on perception of form from motion * New resea plification of vibrations of the basilar membr: * New research streams in the h on the role of outer hair cells in am- n the presence of dorsal uditory system nd vent = New research on th knockout mice += *Olfaerotopie” representation in olfactory cortex * Deep brain stimulation for Parkinson’s disease * The discovery that marcolepyy is a neurodegenerse tive disease of hypocretinergic neurons + New arch on the role of adenosine as a sleep- promoting chemical + The discovery of two families of pheromone recep: tor proteins + New research on the effects of estradiol on women’s sexual interest New researeh on the hu tional memory * New research on the role of the prefrontal cortex in decision making and moral judgments nygelaka andnd aggression * New research on the mechanisms of long: potentiation and its role in learning c cells in spatial memory aging studies of the human * Anew section 0 the bilingual brain * The role of languages with irregular orthography in the prevalence of developmental dyslexia + Evidence for loss of cerebral gray matter in schizophrenia New research on the ini frontal cortex, ven accumbens i Evidence that depression may activity of the amygdala and and hypoat cortex Lack of a fusiform dults ctions between pre- | tegmental area, and nucleus the development of schizophre area in the brains of autistic = A new section on attention-deficit /hyperactivity disorder The role of increased sensitivity of brain glucoc ticoid receptors in posttraumatic stress disorder * The role of basolateral amygdala in classically con- ditioned drug eraving rbitofrontal cortex and involved in drug cra + Fvidence that the rior cingulate cor te ng + Evidence that stress early in life can increase sus ceptibility wo drug addiction There are some important differences bewween this book and Piysiolagy of Behavior. The text of this book is ot simply a shorter and denser ver sor. [kept the illustrative examples, especially those deal ing with human disorders, and added explanations of phenomena to be sure that students without march back ground in biology could understand what I was saying. Although I have simplified some of the detailed expli- nations I have retained the important principles. n of its predeces- Strategies for Learning This theme, which runs throughout the book, was created to help apply physiological psychology to daily fe. You will find a “Strategies for ng” heading Chapter 1, and a chapter entitled “Methods and Strategics of Research.” This chapter dees not co a bewildering list of research methods; instead, reader is led through a set of hypothetical investigations, Praface organized the way that a research project might pro- ced. Each step illustrates a particular procedure in th context in which it would be applied in an ongoing, progrs * Learning Objectives. Fach chapter begins with a list, of learning objectives, which also serve as the f work for the study guide that accompanies this Prologue. A Prologue, which contains the desctip- tion of an episode involving a neurological disorder ‘ran issue in neuroscience, opens each chapter * Epilogue. An Ep solves the issues raised ‘them in terms of what the re chapter, or introduces a rela atthe end of the chapter he prologue, discussing, jer has learned in the ed topic. Interim Summary follows each major sec book. They not only provide useful reviews, but also break each chapter into n ble chunks. * Thought Questions follow most interim s and provide an opportunity to think about what has been learned in the previous sec * Definitions of Key Terms are pri rear the pla Promu cepts, whieh provide a quick review: A list of Sug- gested Readings and Suggested Web Sites provide more information about the topies discussed in thi chapter: Full-Color Art The illustrations in this hook were prepared by Jay Alexander, of Hua Graphies. Jay also works in the Psy: chology Deparunent at the University of Massachusetts and he and I have been working togethes fr several years. I think the result of our collaboration isa set of clear, consistent, and al on my books active illustrations. Strategies for Learning Supplements Ihave prepared a revised CD-ROM, which contains the Newroncience Animations and. the Computeriee Study Guide, The animations demonsty of the most importar principles of neuroscience through movement and inte action. The animations have been substantially revised and expanded from the previous version, modules on neurophysiology (Neural €Pretace ¢ Action Potential, Synapses, and Posisynaptic Poters (Phe Rotatable Brain, Brain Slices, ancl Meninges and CSF), psychopharmacology, research, methods, audition, memory, and verbal communicatio The modules on research methods include three new videos demo de, aecessible through contains a set of self-tests that include multiple-choice questions and an online review of terms, definitions. The questions and list of terms and deli- ns present questionsand keep track of your progress, presenting missed items until you have answered all of «d study guide also in- igrams from the book that will help students learn cermsand concepts. This CD- ROM is included free with the purchase of a new book A Study Guide, whieh my wife and [ wrote, is lable. This workbook provides a framework for study behavior. It promotes a thorough und standing of the principles of physiological psychology through active on in the learning process nsa set of Concept Cards, An i portant part of learning about physiological psychology nd the concept cards will ec!_on one side of ed on the other this book, Allyn & Bacon, hosts a Web site for this text: wwwablongman.com/ carlson6e. This site contains additional multiple-choice test questions for nized by chapter. This forum allows you wo further pr ‘aking. The Web site also provides hot links to other relevant sites of interest and research updates, provided by P man, Professor of Psychology at Texas A\ them correetly. The computeri these Supplements for Instructors ‘eral supplements are available for instructors who adopt this book. © Instructor’s Manual. Writt sity of Peunsytvan by Bill Meil, Indiana this is an exellent toot preparation and ent. Each n ALAGlance Grid, with detailed pedagogical information linking to other available supplements, teaching objectives, lecture material, demonstrations ities, and an updated list, of video, media, suggested readings, and web re- sources. In addition, the appendix includes a com prehensive list of student handouts. ‘+ Test Bank. Written by Paul Wellman, Texas A&M University, this resource is filled with challenging, \vawwvablongman.com/carisonée question yet key concepts. Each chapter has approximately 100 questions, i 1g multiple- choice, true /lalse, shortanswer, and essay, each with, an answer justification, ces, x difficulty rating, and type designation. In addition, the ap- pendix includes a sample ope quiz. This product is also available in TestGen 5.5 compu ized version, for use in creating tests in the € + Powerpoint Presentation. An interactive tool for ts in the elassroom, was ereated by Grant MeLaret Edinboro University of Pennsylvania, Fach chapter Includes images from the textbook, with demon- strations, In add ROM contains the Instruct ual files, ‘* Transparencies for Physiological Psychology, © 2005. Completely updated. There are approximately 100 fulkeolor acetates to enhance classroom lec discussion, This package includes im and Bacon’s major physiological psychology texts. * Digital Media Archive for Physiological Psychology, © 2004. This iy a comprehensive source for video and animation; text images, including charts, graphs, tables, ures; and PowerPoint lecture presentations physiological psychology texts, Its a powe for customized classroom prese! + Physiological Psychology VideoWorkshop, Instruc- tor’s Teaching Guide. A CD-ROM. modules that offei ing VideoWorkshop into your course. It also cludes correlation grids for individual physiologi psychology texts, stu for each video clip, critical thinking questions, multiple-choice ques tions, web links, and an answer key for the Suxdent Learning Guide. For your review, the CD-ROM and Student Learning Guide is also included in the hing Guide, giving you the complete program in one cay reference + CourseCompass. Powered by Blickboard, course management system uses & powerful tools so instruct for any course. In Conclusion Trying to keep up with the rapid progress being made in neuroscience research poses 31 challenge and textbook writers. [fa student simply memorizes what ive believe at the time to he facts, he or she is left with, knowledge that quickly becomes obsolete, In this book 1 have tried to provide enough background material andpugh knowledge of basic physiologi that the reader can revise what he or she when research provides us with new information, I designed this text to be interesting and informa tive. I have endeavored to provide a solid foundation for further study. Students who will not take subsequent, courses in this or related fields should receive the satis tion of a much bet acing of the havior. Also, they will have a greater appreciat forthcoming advances in medical practices related to disorders that affect a person’s perception, mood, or be- hasior. | hope that people who carefully yead this book will henceforth perceive human behavior in a new light. runders own be- Acknowledgements Although I must accept the blame for any shorteomings of the book, I want to thank colleagues who helped me. thank Bill Meil for his work on the instructor's manual and Paul Wellman for his work on the test bank, I wat to thank colleagues who helped me with the present ed tion of this book and the latest edition of Physiology of Be- havior by sending reprints of their work, suggesting. topics that 1 should cover, sending photographs that have been reproduced in this book, and pointing out deficiencies in the previous edition, I thank ing reviewers for their comments on this edition: 1¢ follow Adriana Alcantara, University of Texas at Austin Janis L. And John F Axelson, Holy Gross College Norris R, Bancroft, Weber State Uni son, University of New Mexico Mark E, Basham, Metro State College of Denver Jon E, the, Monme th College Talso want to thank the people at Allyn and Baco Kelly May; my editor, provided assistance, support, and encouragem nd Adam Whitehurst, editorial assis- tant, who helped to gather comments and suggestions. from colleagues who have read the book. Erin ‘oversaw the production of the ancillary material: study guide, Animation CD-ROM, instructor’s manual, Prolace parencies, Videoworkshop, PowerPoint, and digital me- dia archive. Mary Beth Finch, the production editor, as sembled the team that designed and produced the book. Carol Somberg designed the book—which looks. pretty good, I think. Barbara Gracia, of Woodstock Pub: lishers’ Services, demonstrated her masterful skills of or- the book's production, She got thing done on time, despite an extremely tight schedule. Few people realize what a difficult, demand ing, and time-consuming job a production editor has. with a project such as this, with hundreds of illustrations and an author who tends to pi e, but Ido, and thank her forall she has done, Barbara Willette served as copy editor. She gave ice 10 fix my errors be- fore anyone else saw them in print I must also thank my wife Mary for her support Writing isa lonely pursuit, because one must be alone with one’s thoughts for many hours of the day. I thank, her for giving me the time to read, reflect, and wi feeling that I was neglecting her too much To the Reader I hope that in reading this book you will come not only 1 more about the brain but also to appreciate it The b most remarkable thing 10 understand it vl myselF talk wl for the marvelous organ it complex, and perhaps swe are able to use i While working on this book, 1 ing with students, telling them interesting stories about the findings of clinicians and research scientists, Imag: in nice made the task of writing a litle lonely. | hope that the dialogue will continue. Please write to me and tell me what you like and dislike abo the book. My address is: Department of Psychology, To- bin Hall, chusetts 01003. My e-mail address is mn@psych.umassodi, Ifyou write to me (or send me an e-mail), we can make the conversation a nwoaway exchange ng your pres ess NRGCHAPTER OUTLINE Understanding Human Consciousness: Origins of Physiological, ‘=~ Psychology Sanaa The Goals of Resear Biological Roots Physiological Ps Natural Selection and Evolution Functionatism and the Inhentance of Tras Evolution of the Humay Species Evolution of Large Brains Ethical Issues in Research with Animals Careers in Neuroscience Strategies for Learning 1. Describe the behavior of people with split brains and explain what this phenomenon contributes to our understanding of self-awareness. . Describe the goals of scientific research. . Describe the biological roots of physiological psychology, J. Describe the role of natural selection in the evolution of behavioral traits. . Describe the evolution of the human species. . Discuss the value of research with animals and ethical issues concerning their care. . Describe career opportunities in neuroscience. evaweun Outline the strategies that will help you learn as much as possible from this book.René’s Inspiration René, a lonely and intelligent young gurgling of water but at frst could see operating the machinery. He spent the ‘man of eighteen years, had secluded Nothing in the darkness. Ashis eyes be- rest ofthe afternoon wandering through himself in Saint-Germain, a village to ‘came accustomed to the gloom, he ‘the grottoes, listening to the musicand, the west of Paris.Hehad recently suf- could make outa figure illuminated by __being entertained by the statues fered a nervous breakdown and chose _a flickering torch, He approached the During his stay in Saint-Germain, the retreat to recover. Even before com- figure, which he soon recognized as that René visited the rayal gardens again ing to Seint-Germain, he had heard of of ayoung woman. Ashe drew closer, __and again. He had been thinking about ‘the fabulous royal gardens built for he saw thatshe was actually abronze _the relationship between the move: Henri lV and Marie de Médicis, andone statue of Diana, bathing ina pool of ‘ments of animate and inanimate ob- sunny day he decided to visit them. The water. Suddenly, the Greek goddess fled jects, which had concerned philosophers ‘guard stopped him at the gate, but and hd behind a bronze rosebush. As for some time, He thought he saw in the ‘when he identified himself asa student René pursued her, an imposing statue of apparently purposeful, but obviously at the King’s School at La Fléche, he was Neptune rose in front of him, barring _ inanimate, mavernents of the statues an permitted to enter. The gardens con- ‘the way with his trident. ‘answer to some important questions sisted of a series of six large terraces Rend was delighted. He had heard ‘about the relationship between the ‘overlooking the Seine, planted in the about the hydraulically operated me- ‘mind and the body. Even after he left ‘symmetrical, orderly fashion so loved by chanical organs and the moving statues, Saint-Germain, René Descartes revisited ‘the French. Grottoes were cutiinto the bute had not expected such realism. As the grottoes in his memory; and he limestone hillside at the end of each ter- he walked back toward the entrance to went so far as to name his daughter race; René entered one of them. He ‘the grotto, hesaw the plates buried in Francine ater their designers, the heard eerie music accompanied by the _the ground that controlled the valves Francini brothers of Florence. The human nervous sistem makes possible all that we ean do, al that we can and all studying it and unde undertaken. T he last frontier in this world—and perhaps the greatest one—lies within ws we can experience. Is complexity is immense, and the task of tanding it dwarfs all previous explorations our species has One of the most universal of all human characteristics is, curiosity. We want to explain what makes things happen. In times, people believed that natural phenomena were sed by animating spirits. All moving objects—animals, the wind and tides, the sum, moon, and stars—were assumed 10 have spirits that caused them to move. For example, sto fell when they were dropped because 1s spit | with Mother Ear incestors ated and learned more about nature wanted to bi As our bee they abandoned this approach (which we call animism) in fi vor of physical explanations for inanimate moving objects Buc they still used spirits explain human behavior. From the earliest historical times, people have believed that they possessed something intangible that animated them—a mind, a soul, ora spirit. This belief stems from the fact that each of us is aware of his or her own existence. When we thi is thinking or deciding to act. But what is the nature of the huma move them and sensory organs such as eyes and ears that ne more sophis kor act, we feel as though something inside us nindl? We have physical bodies with muscles that ‘Most physiological psychologists believe thet by understanding the workings of the nervous system we eventually will Understand how we think, remember, and act—and will even _ perceive information about the world around us. Within our Understand the nature of our own self-awareness bodies the nervous system plays a central role, receiving iUnderstanding Human Consciousness: A Physiological Approach EE wg the movements of the muscles. GUNA il what role does it play? Does it contol the nervous 9s TAT Reset ee body, completely understand its own formation fre But what is the mi Isita part ofthe nervous system? Is it physical and, ible, like the rest of ‘or is ita spirit that will always remain hidden? workings? A sixteenth-century ictical approach to the study weodeut trom the first edition non produced by of De humani corporis fabrica {On the Workings ofthe Human Body) by Andreas Vesalus Physiolo of hun 1c activity of the n U psychologists take nature, Most of us believe that the mind is a phenom vous system, We believe that o cand the workings of the human bed)—in particular, the workings of the nervous system—we will be able to explain how we perceive, how we think, how we remember, and how we act ‘We will even be able to explain the nature of our own selfawareness, OF course, Wwe are far fiom: understanding the workings of the nervous system, so only time will tell whether this belief is justified. isness? First, let's define variety of concepts, in- ay write about an experiment us- the rats were awake and not How can physiological psychologists study human conscio ‘our terms. The word consciousness can be used to refer to cluding simple wakefulness. Thus, a researcher ing “conscious rats,” referring to the fact th. anesthetized. By consciousness, Lam referring to something else: the fact that we hi mans are aware of—and can tell others about—our thoughts, perceptions, memo- ries, and feelings. ‘We know that bra cause consciousness can be brain, we may hypothesize that consciousness is a physiological fi havior is, We can even speculate about the origins of this selfawareness. Conscio ness and the abil seem (© go hand in hand, Our species, with its complex social structure and enormous capacity for learning, is well served by our ability to communicate: to express intentions to one another and 10 make requests of one another, Verbal communication makes cooperation possible and permits us to establish customs and laws of behavior. Perhaps the evolution of this ability iswhat has given tise to the phenom ousness. That is, our ability to send and receive messages with other people enables us to send and receive our own mes and to be aware of our own existence, (See Fig: damage or drugs € ndly affect consciousness, Be- in the structure or chemistry of the sages—in other words, to th ure 1.) Split Brains Studies of humans who have undergone a particular surgical procedure demon- strate dramatically how disconnecting parts of the brain that are involved with per ceptions from parts that are involved with verbal behavior also disconnects them from consciousness. These results suggest that the parts of the brain involved in ver= bal behavior may be the ones responsible for consciousness ‘The surgical procedure is one that has been used for people with very severe epilepsy that cannot be controlied by drugs. In these people, nerve cells in one side of the brain become overactive, and the overactivity is transmitted to the other sie ofthe brain bya structure called the corpus callosum, The corpus eallosum isi large bundle of nerve fibers that connec! corresponding paris of one side of the brain with those of the other. Both sides of the brain then engage ‘each other, causing a generalized epileptic seizure. Th times each day, preventing the person from leading a normal life. Neurosurgec corpus callosum (ore pus ka low sum) large tunel of nerve tibers that connect cote sponding parts of one se ofthe bain with those of the ther n will activity ancl stimCHAPTER 1: Orgs ofPysclogkalPychoogy ‘wwwablongman com/earlzonée “The split-orain operation, A “window” has been opened in ‘the side of the brain so that we can see the corpus callosum being cut atthe midline ofthe brain a Corpus callosum PASS. Front split-brain operation &a ‘surgery that occasional oer formed to teat a form of epilepsy, the surgeon cuts the cor ps callosum, which connects the | wo hemspheres ofthe brain. cerebral hemispheres The 190 symmetical halves ofthe bran, constitute the major part of the ban, discovered that cutting the corpus callosum (ihe splitbrain operation) greatly reduced the frequency of the epileptic Figure 1 ‘ ain operation. We | down the middle, from front to back, dividing it into its two symmetrical halves. A “window” has been opened in the left side of the brain so that we can see the co Cutting device (See Figure 1.2.) Sperry (1966) ancl Gazzaniga and his associates (Gauzzaniga, 1970; Gazzaniga and LeDoux, 1978) have studied these patien extensively. The largest part of the brain consists of two syn metrical parts, called the cerebral hemispheres, which receive for fiom the opposite sides of the body. They also control movements of the opposite sides. The corpus callo- suum enables the two hemispheres to share information so that each side knows what t side is perceiving and doing. AF ter the splitbrain operation is performed, the two hemispheres e disconnected and operate independently. Their sensory mechanisms, memories, and motor ystems can no longer € a pus callosum being cut by the neurosurgeon’s special knife. change informat c effects of these disconnections are not obvious to the casual observer, for the simple reason that only fone hemisphiere—in most people, the lefi—controls speech. 'e of an epileptic person with a split b onably The right hemisph appears to be able to understand ve vel, but itis incapable of producing speech Because only one side of the brain can talk about what itis experiencing, peo: ple who speak with a person with a split brain are conversing with only one hem sphere: the left. The operations ofthe right hemisphere are more difficult to detect. Even the patient's left hemisphere has to learn about the indep the right hemisphere, One of the first things that these patients say they notice after the operation is that their left hand seems to have a “mind of its own.” For example, patients may find themselves putting down a book held in the left hand, even if they have been reading it with great imterest, This conflict occurs because the right hem spher ntrols the left hand, cannot read and therefore finds the book bor ing. Atother times, these patients surprise themselves by making obscene gestures (with the left hand) when they had not intended! to. A psyehologist once reported that a man with a split brain had attempted to beat his wife with one hand and pro- tect her with the other. Did he really want to hurt her? Yes and no, [ guess. “ ok factory system. That is, when a person snifls a flower through the left nostril, only the left brain receives a sensation of the odor. Thus, if the right nostil of a patient with a split brain is closed, leaving only the left nostril open, the patient will be able w tell us what the odors are (Gordon and Sperry, 1969). However, ifthe odor enters the right nostil, the patient will say that he or she smells nothing. But, in fact, the right brain has perceived the odor and can identify it. To show that this is $0, we ask the patient to smell an odor with the right nostril and then reach for some objects that are hidden from view by a partition, If asked to use the left hand, controlled by hemisphere that decected the smell, the patient will select the object that corre sponds to the odor—a plastie lower for a floral odor, a toy fish for a fishy odo model tree for the odor of pine, and so forth. But ifasked to use the right hand, the patient fails the test because the right hand is connected to the left hemisph which did not smell the odor. (See Figure 1.3.) The effects of cutting the corpus callosum reinforce the conclusion that we be- ich the p whieh e« 1¢ exception to the crossed representation of sensory information is come conscious of something only if information about itis able to 1