NEUROSCIENCE Edited by DALE PURVES GEORGE J. AUGUSTINE DAVID FITZPATRICK WILLIAM C. HALL ANTHONY-SAMUEL LAMANTIA JAMES O. MCNAMARA S MARK WILLIAMS Sinauer associates, Inc. Publishers Sunderland, Massachusetts U.S.A
Edited by DALE PURVES GEORGE J. AUGUSTINE DAVID FITZPATRICK WILLIAM C. HALL ANTHONY-SAMUEL LAMANTIA JAMES O. MCNAMARA S. MARK WILLIAMS NEUROSCIENCE THIRD EDITION Sinauer Associates, Inc. • Publishers Sunderland, Massachusetts U.S.A. Purves3/eFM 5/13/04 12:59 PM Page iii
THE COVER Dorsal view of the human brain Courtesy of S. Mark Williams Neuroscience NEUROSCIENCE: Third Edition Copyright C 2004 by Sinauer Associates, Inc. All rights reserved This book may not be reproduced in whole or in part without permission. Address inquiries and orders to Sinauer Associates, Inc. 23 Plumtree road Sunderland, MA 01375 U.S.A vww. sinauer. com FAX:413-549-1118 orders@sinauer.com publish@sinauer.com Library of Congress Cataloging-in-Publication Data Neuroscience / edited by Dale Purves [et al. ].-3rd ed Includes bibliographical references and index ISBN 0-87893-725-0(casebound alk paper) 1. Neurosciences DNLM: 1. Nervous System Physiology. 2. Neurochemistry WL 102 N50588 2004 I Purves, Dale QP355.2N4872004 6128-dc22 2004003973 Printed in U.S.A 54321
NEUROSCIENCE: Third Edition Copyright © 2004 by Sinauer Associates, Inc. All rights reserved. This book may not be reproduced in whole or in part without permission. Address inquiries and orders to Sinauer Associates, Inc. 23 Plumtree Road Sunderland, MA 01375 U.S.A. www.sinauer.com FAX: 413-549-1118 orders@sinauer.com publish@sinauer.com Library of Congress Cataloging-in-Publication Data Neuroscience / edited by Dale Purves ... [et al.].— 3rd ed. p. ; cm. Includes bibliographical references and index. ISBN 0-87893-725-0 (casebound : alk. paper) 1. Neurosciences. [DNLM: 1. Nervous System Physiology. 2. Neurochemistry. WL 102 N50588 2004] I. Purves, Dale. QP355.2.N487 2004 612.8—dc22 2004003973 Printed in U.S.A. 5 4 3 2 1 THE COVER Dorsal view of the human brain. (Courtesy of S. Mark Williams.) Purves3/eFM 5/13/04 12:59 PM Page iv
Contributo George J. Augustine, Ph. D Dona M. Chikaraishi, Ph Michael d. ehlers, M. D. Ph.D Gillian einstein ph D David Fitzpatrick, ph D William C. Hall, ph. D Erich Jarvis, Ph. Lawrence C. Katz, Ph D Julie Kauer, Ph. D Anthony-Samuel LaMantia, Ph D ames O. McNamara, MD Richard D. Mooney, Ph D Miguel A. L Nicolelis, M. D, Ph D Dale Purves, M.D Peter H. Reinhart. Ph. D Sidney A. Simon, Ph. D J H. Pate Skene, Ph D ames Voyvodic, ph. D Leonard E White. Ph. D S Mark Williams, Ph. D UNIT EDITORS UNIT I: George J. Augustine UNIT I: David Fitzp UNIT III: William C UNIT IV: Anthony-Samuel LaMantia UNIT V: Dale purves
George J. Augustine, Ph.D. Dona M. Chikaraishi, Ph.D. Michael D. Ehlers, M.D., Ph.D. Gillian Einstein, Ph.D. David Fitzpatrick, Ph.D. William C. Hall, Ph.D. Erich Jarvis, Ph.D. Lawrence C. Katz, Ph.D. Julie Kauer, Ph.D. Anthony-Samuel LaMantia, Ph.D. James O. McNamara, M.D. Richard D. Mooney, Ph.D. Miguel A. L. Nicolelis, M.D., Ph.D. Dale Purves, M.D. Peter H. Reinhart, Ph.D. Sidney A. Simon, Ph.D. J. H. Pate Skene, Ph.D. James Voyvodic, Ph.D. Leonard E. White, Ph.D. S. Mark Williams, Ph.D. UNIT EDITORS UNIT I: George J. Augustine UNIT II: David Fitzpatrick UNIT III: William C. Hall UNIT IV: Anthony-Samuel LaMantia UNIT V: Dale Purves Contributors Purves3/eFM 5/13/04 12:59 PM Page v
Contents in brief 1. Studying the Nervous Systems of Humans and Other Animals UNIT NEURAL SIGNALING 2. Electrical Signals of Nerve Cells 31 3. Voltage-Dependent Membrane Permeability 4 4 Channels and Transporters 69 5. Synaptic Transmission 93 6. Neurotransmitters, Receptors, and Their Effects 129 7. Molecular Signaling within Neurons 165 UNIT/ SENSATION AND SENSORY PROCESSING 8. The Somatic Sensory System 189 ain 10. Vision: The Eye 229 11. Central Visual Pathways 259 12. The Auditory System 283 13. The Vestibular System 315 14. The Chemical Senses 337 UNIT I MOVEMENT AND /TS CENTRAL CONTROL 15. Lower Motor Neuron Circuits and motor Control 371 16. Upper Motor Neuron Control of the Brainstem and Spinal Cord 393 17. Modulation of Movement by the Basal Ganglia 417 18 Modulation of Movement by the Cerebellum 435 19. Eye Movements and Sensory Motor Integration 453 20. The Visceral Motor System 469 UNIT/V THE CHANGING BRAIN 21. Early Brain Development 501 22. Construction of Neural Circuits 521 23. Modification of Brain Circuits as a Result of Experience 557 24. Plasticity of Mature Synapses and Circuits 575 UNITV COMPLEX BRAIN FUNCTIONS 25. The Association Cortices 613 26. Language and Speech 637 27. Sleep and Wakefulness 659 28. Emotions 687 29. Sex, Sexuality, and the Brain 711 30. Memory 733 APPENDIX A THE BRAINSTEM AND CRANIAL NERVES 755 APPENDIX B VASCULAR SUPPLY, THE MENINGES, AND THE VENTRICULAR SYSTEM 763
1. Studying the Nervous Systems of Humans and Other Animals 1 UNIT I NEURAL SIGNALING 2. Electrical Signals of Nerve Cells 31 3. Voltage-Dependent Membrane Permeability 47 4. Channels and Transporters 69 5. Synaptic Transmission 93 6. Neurotransmitters, Receptors, and Their Effects 129 7. Molecular Signaling within Neurons 165 UNIT II SENSATION AND SENSORY PROCESSING 8. The Somatic Sensory System 189 9. Pain 209 10. Vision: The Eye 229 11. Central Visual Pathways 259 12. The Auditory System 283 13. The Vestibular System 315 14. The Chemical Senses 337 UNIT III MOVEMENT AND ITS CENTRAL CONTROL 15. Lower Motor Neuron Circuits and Motor Control 371 16. Upper Motor Neuron Control of the Brainstem and Spinal Cord 393 17. Modulation of Movement by the Basal Ganglia 417 18. Modulation of Movement by the Cerebellum 435 19. Eye Movements and Sensory Motor Integration 453 20. The Visceral Motor System 469 UNIT IV THE CHANGING BRAIN 21. Early Brain Development 501 22. Construction of Neural Circuits 521 23. Modification of Brain Circuits as a Result of Experience 557 24. Plasticity of Mature Synapses and Circuits 575 UNIT V COMPLEX BRAIN FUNCTIONS 25. The Association Cortices 613 26. Language and Speech 637 27. Sleep and Wakefulness 659 28. Emotions 687 29. Sex, Sexuality, and the Brain 711 30. Memory 733 APPENDIX A THE BRAINSTEM AND CRANIAL NERVES 755 APPENDIX B VASCULAR SUPPLY, THE MENINGES, AND THE VENTRICULAR SYSTEM 763 Contents in Brief Purves3/eFM 5/13/04 12:59 PM Page vii
Contents Preface xvi Acknowledgments xvii Supplements to Accompany NEUROSCIENCe xviii Chapter 1 Studying the Nervous Systems of humans and other Animals 1 ervie Overall Organization of the Human Nervous Genetics, Genomics, and the brain 1 The Cellular Components of the Nervous System 2 Neuroanatomical Terminology 16 Neurons 4 The Subdivisions of the Central Nervous System 18 Neuroglial Cells 8 Organizational Principles of Neural Systems 20 Cellular Diversity in the Nervous System 9 Functional Analysis of Neural Systems 2 Neural Circuits 1 Analyzing Complex Behavior 24 Box A Brain Imaging Techniq Unit NEURAL SIGNALING Chapter 2 Electrical Signals Chapter 3 Voltage-Dependent Membrane of Nerve Cells 31 Permeability 47 Overview 31 Overview 47 Electrical Potentials across Nerve Cell Membranes 31 Ionic Currents Across Nerve Cell Membranes 4 How Ionic Movements Produce Electrical Signals 34 Box A The Voltage Clamp Method 48 The Forces That Create Membrane Potentials 36 Two Types of Voltage-Dependent Ionic Current 49 Electrochemical Equilibrium in an Environment with Two Voltage-Dependent Membrane Conductances 52 More Than one permeant ion 38 Reconstruction of the Action Potential 54 The lonic Basis of the Resting Membrane Potential 40 Long-Distance Signaling by Means of Action Box A The remarkable giant Nerve cells Potentials 56 of Squid Box B Threshold 57 The Ionic Basis of Action Potentials 43 Box C Passive Membrane Properties 60 Box B Action potential form The Refractory Period 61 and Nomenclature 44 ncreased Conduction Velocity as a Result Summary 45 of Myelination 63 Summary 65 Box D Multiple Sclerosis 66
Chapter 1 Studying the Nervous Systems of Humans and Other Animals 1 Overview 1 Genetics, Genomics, and the Brain 1 The Cellular Components of the Nervous System 2 Neurons 4 Neuroglial Cells 8 Cellular Diversity in the Nervous System 9 Neural Circuits 11 Overall Organization of the Human Nervous System 14 Neuroanatomical Terminology 16 The Subdivisions of the Central Nervous System 18 Organizational Principles of Neural Systems 20 Functional Analysis of Neural Systems 23 Analyzing Complex Behavior 24 BOX A Brain Imaging Techniques 25 Summary 26 Contents Unit I NEURAL SIGNALING Chapter 2 Electrical Signals of Nerve Cells 31 Overview 31 Electrical Potentials across Nerve Cell Membranes 31 How Ionic Movements Produce Electrical Signals 34 The Forces That Create Membrane Potentials 36 Electrochemical Equilibrium in an Environment with More Than One Permeant Ion 38 The Ionic Basis of the Resting Membrane Potential 40 BOX A The Remarkable Giant Nerve Cells of Squid 41 The Ionic Basis of Action Potentials 43 BOX B Action Potential Form and Nomenclature 44 Summary 45 Chapter 3 Voltage-Dependent Membrane Permeability 47 Overview 47 Ionic Currents Across Nerve Cell Membranes 47 BOX A The Voltage Clamp Method 48 Two Types of Voltage-Dependent Ionic Current 49 Two Voltage-Dependent Membrane Conductances 52 Reconstruction of the Action Potential 54 Long-Distance Signaling by Means of Action Potentials 56 BOX B Threshold 57 BOX C Passive Membrane Properties 60 The Refractory Period 61 Increased Conduction Velocity as a Result of Myelination 63 Summary 65 BOX D Multiple Sclerosis 66 Preface xvi Acknowledgments xvii Supplements to Accompany NEUROSCIENCE xviii Purves3/eFM 5/13/04 12:59 PM Page viii