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The central nervous system : structure and function PDF

532 Pages·2004·101.917 MB·English
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The Central Nervous System This page intentionally left blank THE CENTRAL NERVOUS SYSTEM Structure and Function THIRD EDITION PER BRODAL Institute of Basic Medical Sciences University of Oslo OXPORD UNIVERSITY PRESS 2004 OXFORD UNIVERSITY PRESS Oxford New York Auckland Bangkok Buenos Aires Cape Town Chennai Dar es Salaam Delhi Hong Kong Istanbul Karachi Kolkata Kuala Lumpur Madrid Melbourne Mexico City Mumbai Nairobi Sao Paulo Shanghai Taipei Tokyo Toronto Copyright © 1992, 1998, 2004 by Oxford University Press, Inc. Published by Oxford University Press, Inc. 198 Madison Avenue, New York, New York, 10016 http://www.oup-usa.org Oxford is a registered trademark of Oxford University Press All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of Oxford University Press. Library of Congress Cataloging-in-Publication Data Brodal, Per. The central nervous system : structure and function / Per Brodal.—3rd ed. p. cm. Includes bibliographical references and index. ISBN 0-19-516560-8 1. Central nervous system. I. Tide. QP370.B76 2003 612.8'2—dc21 2003042924 987654321 Printed in the United States of America on acid-free paper Preface This book is intended primarily for use by students of medicine, physical therapy, and psychology—that is, for use in neuroscience or neuroanatomy courses by stu- dents who need knowledge of the nervous system as a basis for later clinical study and practice. The need for a third edition since 1998 attests to the rapid pace of development in the neurosciences and related fields. In this edition I have tried to incorporate new findings by rewriting rather than just adding new material. In this way, the text has not been significantly ex- panded. Many chapters have been reorganized, and two new chapters deal with the vestibular system (Chapter 9) and control of eye movements (Chapter 15). New illustrations have been added, and most others have been redrawn to im- prove their clarity. My intentions remain the same as those of my father, Alf Brodal, when he wrote the Norwegian forerunner of this book more than 50 years ago: to stimulate un- derstanding rather than memorization of isolated facts, while at the same time fos- tering a realistic attitude toward our still-limited ability to explain the marvels of the human brain. The book aims to present the difficult subject of neuroscience so that those ap- proaching it for the first time can understand it. Therefore, many details are left out that might be of great interest to the specialist but would merely obscure the essentials for the beginner. Everyday experiences and clinical examples are inte- grated throughout the text to help students link the new material with their prior knowledge and future profession. The nervous system, however, is exceedingly com- plex, both structurally and functionally, and much remains to be learned before we can answer many fundamental questions. Thus, while an undergraduate course can provide only partial insights, no one is served by a presentation that avoids con- troversial issues and areas of ignorance. Indeed, pointing out what we do not know is sometimes better than presenting an oversimplified version. For this reason I have also discussed how the data were obtained and the limitations inherent in the various methods. The main challenge—for both the student and the scientist—is to understand how the nervous system solves its multifarious tasks. This requires an integrated ap- proach, drawing on data from all fields of neurobiology, as well as from psychol- ogy and clinical research. Textbooks sharing this goal nevertheless differ markedly in how they present the material and where they put the emphasis. Perhaps be- cause my own field of research is the wiring patterns of the brain, I strongly feel that knowledge of how the nervous system is built—in particular, how the various parts are interconnected to form functional systems—is a prerequisite for proper understanding of data from other fields. A fair knowledge of brain anatomy is es- pecially important for sound interpretations of the symptoms of brain disease. Text- books of neuroanatomy often overwhelm the reader with details that are not strictly relevant for either functional analysis or clinical thinking. Neither does a strong emphasis on cellular mechanisms at the expense of the properties of neural sys- tems seem the right choice if the aim is to help readers understand how the brain V vi Preface performs its tasks and how the site of a disease process relates to a patient's symp- toms. Therefore, neither anatomical nor cellular and molecular details are included in this book if they cannot in some way be related to function. My hope is that the book presents a balance of cellular and neural systems material that is right for stu- dents. In-depth sections and more advanced clinical material are placed in boxes so that they should not disturb reading of the main text. Since the needs of readers differ, however, they are encouraged to read selectively and pick the material they find most relevant and interesting from their perspective, regardless of whether it is placed in the main text or in boxes. The frequent subheadings should facilitate such selective reading. Difficult or complex subjects are preceded with an overview that may suffice for some readers, while other readers will want to read all the text, including the boxes. During the preparation of the former and the present editions, I have received help from several colleagues, for which I am truly grateful. Jan Bjaalie, Niels Chris- tian Danbolt, Paul Heggelund, Jan Jansen, Harald Kryvi, Kirsten Osen, Ole Petter Ottersen, Eric Rinvik, and Jon Storm-Mathisen have all provided constructive crit- icism and advice. I also gratefully acknowledge the expert help of Kari Ruud, who prepared most of the drawings, and of Gunnar Lothe and Carina Ingebrigtsen, who produced the photographic work. Oslo, Norway P. B. Contents A Brief Survey, xi Studying the Structures and Function of the Nervous System, xiii Animal Experiments Crucial for Progress, xiii Ethics and Animal Experiments, xiv Sources of Error in All Methods, xiv Revising Scientific Truths from Time to Time, xv I MAIN FEATURES OF STRUCTURE AND FUNCTION 1. Cellular Elements of Nervous Tissue, 3 Structure of the Neuron, 3 Coupling of Neurons: Pathways for Signals, 10 The Cytoskeleton and Axonal Transport, 15 dial Cells, 17 2. Functional Properties of Neurons, 26 Basis of Excitability and Impulse Propagation, 26 Synaptic Transmission, 35 Neurotransmitters and Their Receptors, 47 3. Parts of the Nervous System, 66 The Spinal Cord, 66 The Brain Stem, 74 The Cerebrum, 83 The Cerebellum, 88 The Coverings of the Brain, 89 The Cerebral Ventricles and the Cerebrospinal Fluid, 90 The Blood Supply of the Central Nervous System, 94 4. Development, Aging, and Plasticity of the Nervous System, 103 Development of the Nervous System, 103 The Nervous System and Aging, 122 Restitution of Function, 125 II SENSORY SYSTEMS 5. Sensory Receptors in General, 135 Sensory Units and Their Receptive Fields, 135 Transduction: The Translation of Stimuli to Action Potentials, 135 vii viii Contents Receptor Properties, 137 Classification on the Basis of Receptor Properties, 137 Receptors and Subjective Sensory Experience, 138 6. The Somatosensory System, 139 Exteroceptors: Cutaneous Sensation, 139 Proprioceptors: Deep Sensation, 145 The Sensory Fibers and the Dorsal Roots, 155 The Central Somatosensory Pathways, 161 The Somatosensory Cortical Regions, 172 Pain, 175 7. The Visual System, 183 The Eyeball and the Refracting Media, 183 The Retina, 185 Organization of the Visual Pathways, 194 The Visual Cortex and the Final Processing of Visual Information, 200 8. The Auditory System, 208 The Cochlea, 208 The Auditory Pathways, 215 The Auditory Cortex, 219 9. The Sense of Equilibrium, 221 Structure and Function of the Vestibular Apparatus, 221 Connections of the Vestibular Nuclei, 224 Vestibular Reflexes, 226 Receptors and Reflexes of Importance for Body Balance, 228 Cortical Processing of Vestibular Signals, 231 10. Olfaction and Taste, 233 The Olfactory System, 233 The Sense of Taste, 237 III MOTOR SYSTEMS 11. The Peripheral Motor Neurons and Reflexes, 243 Motoneurons and Muscles, 243 Reflexes, 251 Muscle Tone, 258 Injury of Peripheral Motor Neurons and Regeneration, 261 12. The Central Motor Pathways, 265 The Pyramidal Tract (The Corticospinal Tract), 265 Other Descending Pathways to the Spinal Cord, 271 Control of Automatic Movements, 274 Motor Cortical Areas and Control of Voluntary Movements, 276 Symptoms Caused by Interruptions of Central Motor Pathways (Upper Motor Neurons), 281 Contents ix 13. The Basal Ganglia, 286 Structure and Connections of the Basal Ganglia, 286 The Ventral Striatum, 296 Functions of the Basal Ganglia, 297 Diseases of the Basal Ganglia, 299 14. The Cerebellum, 303 Subdivisions and Afferent Connections of the Cerebellum, 303 The Cerebellar Cortex and the Mossy and Climbing Fibers, 309 Efferent Connections of the Cerebellum, 312 Cerebellar Functions and Symptoms in Disease, 315 15. Control of Eye Movements, 321 The Eye Muscles and Movements of the Eyes, 321 Central Control of Eye Movements, 323 IV THE BRAIN STEM AND THE CRANIAL NERVES 16. Reticular Formation, 333 Structure and Connections of the Reticular Formation, 333 Functions of the Reticular Formation, 340 Consciousness, 342 Sleep, 345 17. The Cranial Nerves, 349 General Organization of the Cranial Nerves, 349 The Hypoglossal Nerve, 353 The Accessory Nerve, 354 The Vagus Nerve, 355 The Glossopharyngeal Nerve, 358 The Vestibulocochlear Nerve, 359 The Facial and Intermediate Nerves, 359 The Trigeminal Nerve, 361 The Abducens, Trochkar, and Oculomotor Nerves, 364 V THE AUTONOMIC NERVOUS SYSTEM 18. The Peripheral Autonomic Nervous System, 371 General Organization of the Autonomic System, 371 Peripheral Parts of the Sympathetic System, 374 Peripheral Parts of the Parasympathetic System, 380 The Enteric Nervous System, 382 Functional Aspects of the Autonomic Nervous System, 382 Neurotransmitters in the Autonomic Nervous System, 386 Sensory Innervation of Visceral Organs and Visceral Reflexes, 389 19. The Central Autonomic System: Hypothalamus, 397 Centers in the Brain Stem for Coordination of Behavior, 397 Structure and Connections of the Hypothalamus, 398

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