SIXTH EDITION Companion Website oup-arc.com/access/purves-6e The Neuroscience, Sixth Edition companion website offers review and study tools to help you master the material presented in the textbook. Access to the site is free of charge. The site includes: n Chapter Outlines: The complete heading structure of each chapter. n Chapter Summaries: Concise overviews of the important topics covered in each chapter. n Animations: Detailed animations present concepts in a dynamic manner to help you visualize and better understand many of the complex processes of neuroscience. n Web Topics: New for this edition, these provide novel or historical topics for special discussion. n Flashcards: Flashcard activities help you master the extensive vocabulary of neuroscience. n Glossary: The complete glossary, including all the bold terms from the textbook. Sylvius 4 Online by S. Mark Williams and Leonard E. White An Interactive Atlas and Visual Glossary of Human Neuroanatomy Each new copy of Neuroscience, Sixth Edition includes access to Sylvius—a unique digital learning environment for exploring and understanding the structure of the human central nervous system. Sylvius features fully annotated surface views of the human brain, as well as interactive tools for dissecting the central nervous system and viewing annotated cross-sections of preserved specimens and living subjects imaged by magnetic resonance. See next page for details and access instructions. NeuroSci6E_Endpapers.indd 2-3 9/8/17 10:48 AM Sylvius Components and Features n Surface Anatomy Atlases: Photographic, Magnetic Resonance Image, Brainstem Model SIXTH EDITION n Sectional Anatomy Atlases: Photographic, Magnetic Resonance Image, Brainstem and Spinal Cord n Pathways: Allow you to follow the flow of information in several important long-tract pathways of the central nervous system n Visual Glossary: Searchable glossary that provides visual representations, concise anatomical and functional definitions, and audio pronunciation of neuroanatomical structures n Over 500 neuroanatomical structures and terms are identified and described Access Instructions To access Sylvius, follow these instructions: 1. Retrieve your Sylvius 4 Online registration code from your ebook bookshelf. Click here for directions. 2. Go to https://sylvius.sinauer.com. 3. Click “Register using a registration code”, and follow the instructions to create an account. (Requires a valid email address.) 4. Log in at https://sylvius.sinauer.com using your email address and new password. New codes can be purchased online at: http://register.dashboard.oup.com. NeuroSci6E_Endpapers.indd 4-5 9/8/17 10:48 AM SIXTH EDITION 00_FM_PurvesNS_6e.indd 1 9/14/17 4:25 PM 00_FM_PurvesNS_6e.indd 2 9/14/17 4:25 PM EDITORS Dale Purves • George J. Augustine David Fitzpatrick • William C. Hall • Anthony-Samuel LaMantia Richard D. Mooney • Michael L. Platt • Leonard E. White Sinauer Associates NEW YORK OXFORD OXFORD UNIVERSITY PRESS 00_FM_PurvesNS_6e.indd 3 9/14/17 4:25 PM NEUROSCIENCE, Sixth Edition Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide. Oxford is a registered trade mark of Oxford University Press in the UK and certain other countries. Published in the United States of America by Oxford University Press 198 Madison Avenue, New York, NY 10016, United States of America © 2018 Oxford University Press Sinauer Associates is an imprint of Oxford University Press. For titles covered by Section 112 of the US Higher Education Opportunity Act, please visit www.oup.com/us/he for the latest information about pricing and alternate formats. All rights reserved. No part of this publication may be reproduced, stored in a retrieval sys- tem, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, by license, or under terms agreed with the appropriate reproduction rights organization. Cover image Courtesy of Inquiries concerning reproduction outside the scope of the above should be sent to the Rights Nan-Kuei Chen and Allen W. Song Department, Oxford University Press, at the address above. You must not circulate this work in any other form and you must impose this same condition on any acquirer. Address editorial correspondence to: Sinauer Associates 23 Plumtree Road Sunderland, MA 01375 USA [email protected] Address orders, sales, license, permissions, and translation inquiries to: Oxford University Press USA 2001 Evans Road Cary, NC 27513 USA Orders: 1-800-445-9714 Library of Congress Cataloging-in-Publication Data Names: Purves, Dale, editor. Title: Neuroscience / editors, Dale Purves, George J. Augustine, David Fitzpatrick, William C. Hall, Anthony-Samuel LaMantia, Richard D. Mooney, Michael L. Platt, Leonard E. White ; contributors, Ben Hayden, Fan Wang. Description: Sixth edition. | Sunderland, Massachusetts : Oxford University Press, [2018] | Includes bibliographical references and index. Identifiers: LCCN 2017028094 | ISBN 9781605353807 (casebound) Subjects: LCSH: Neurosciences. Classification: LCC QP355.2 .N487 2018 | DDC 612.8--dc23 LC record available at https://lccn.loc.gov/2017028094 9 8 7 6 5 4 3 2 1 Printed in the United States of America 00_FM_PurvesNS_6e.indd 4 9/14/17 4:25 PM Contents in Brief 1 Studying the Nervous System 1 UNIT I Neural Signaling 31 2 Electrical Signals of Nerve Cells 33 3 Voltage-Dependent Membrane Permeability 49 4 Ion Channels and Transporters 65 5 Synaptic Transmission 85 6 Neurotransmitters and Their Receptors 113 7 Molecular Signaling within Neurons 145 8 Synaptic Plasticity 169 UNIT II Sensation and Sensory Processing 191 9 The Somatosensory System: Touch and Proprioception 193 10 Pain 213 11 Vision: The Eye 233 12 Central Visual Pathways 261 13 The Auditory System 281 14 The Vestibular System 305 15 The Chemical Senses 323 UNIT III Movement and Its Central Control 355 16 Lower Motor Neuron Circuits and Motor Control 357 17 Upper Motor Neuron Control of the Brainstem and Spinal Cord 381 18 Modulation of Movement by the Basal Ganglia 407 19 Modulation of Movement by the Cerebellum 427 20 Eye Movements and Sensorimotor Integration 447 21 The Visceral Motor System 465 U NIT IV The Changing Brain 489 22 Early Brain Development 491 23 Construction of Neural Circuits 521 24 Circuit Development: Intrinsic Factors and Sex Differences 551 25 Experience-Dependent Plasticity in the Developing Brain 571 26 Repair and Regeneration in the Nervous System 595 UNIT V Complex Brain Functions and Cognitive Neuroscience 625 27 Cognitive Functions and the Organization of the Cerebral Cortex 627 28 Cortical States 643 29 Attention 667 30 Memory 681 31 Emotion 703 32 Thinking, Planning, and Deciding 725 33 Speech and Language 745 34 Development and Evolution of Cognitive Functions 767 Appendix A1 Atlas: The Human Central Nervous System AT1 00_FM_PurvesNS_6e.indd 5 9/14/17 4:25 PM Contributors George J. Augustine, Ph.D. David Fitzpatrick, Ph.D. William C. Hall, Ph.D. Ben Hayden, Ph.D. Anthony-Samuel LaMantia, Ph.D. Richard D. Mooney, Ph.D. Michael L. Platt, Ph.D. Dale Purves, M.D. Fan Wang, Ph.D. Leonard E. White, Ph.D. Unit Editors UNIT I: George J. Augustine UNIT II: David Fitzpatrick and Richard D. Mooney UNIT III: Leonard E. White and William C. Hall UNIT IV: Anthony-Samuel LaMantia UNIT V: Dale Purves and Michael L. Platt 00_FM_PurvesNS_6e.indd 6 9/14/17 4:25 PM Contents CHAPTER 1 Studying the Nervous System 1 Structural Analysis of Neural Systems 19 Functional Analysis of Neural Systems 21 Overview 1 Analyzing Complex Behavior 22 Genetics and Genomics 1 Imaging the Living Human Brain 22 Cellular Components of the Nervous System 2 Early Brain Imaging Using X-Rays 22 Neurons 5 Early Functional Mapping Using Cortical Surface Glial Cells 7 Stimulation and Electroencephalography 23 Cellular Diversity in the Nervous System 9 Computerized Tomography 25 Neural Circuits 10 Magnetic Resonance Imaging 25 Other Ways to Study Neural Circuits 11 Functional Brain Imaging 27 Organization of the Human Nervous System 13 Summary 28 Neural Systems 16 Additional Reading 29 Genetic Analysis of Neural Systems 16  BOX 1A Model Organisms in Neuroscience 17 Neural Signaling 31 UNIT I CHAPTER 2 Electrical Signals of Nerve  BOX 2A The Remarkable Giant Nerve Cells of Squid 45 Cells 33 The Ionic Basis of Action Potentials 46 Overview 33  BOX 2B Action Potential Form and Electrical Signals of Nerve Cells 33 Nomenclature 47 Long-Distance Transmission of Electrical Summary 47 Signals 35 Additional Reading 48  CLINICAL APPLICATIONS Anesthesia and Neuronal Electrical Signaling 37 CHAPTER 3 How Ion Movements Produce Electrical Signals 39 Forces That Create Membrane Potentials 41 Voltage-Dependent Membrane Electrochemical Equilibrium in an Environment with Permeability 49 More Than One Permeant Ion 42 Overview 49 The Ionic Basis of the Resting Membrane Ion Currents across Nerve Cell Membranes 49 Potential 44  BOX 3A The Voltage Clamp Method 50 00_FM_PurvesNS_6e.indd 7 9/14/17 4:25 PM