LWBK124-392G-FM[i-xviii] 10/17/08 8:16 AM Page i Aptara (PPG-Quark) C LINICAL N EUROANATOMY S E V E N T H E D I T I O N Richard S.Snell, M.R.C.S., L.R.C.P., MB, BS, MD, PhD Emeritus Professor of Anatomy George Washington University School of Medicine and Health Sciences Washington,DC Formerly Associate Professor of Anatomy and Medicine,Yale University Medical School;Lecturer in Anatomy King’s College University of London;and Visiting Professor of Anatomy,Harvard Medical School. LWBK124-392G-FM[i-xviii] 10/17/08 8:16 AM Page ii Aptara (PPG-Quark) Acquisitions Editor:Crystal Taylor Managing Editor:Kelly Horvath Marketing Manager:Emilie Linkins Managing Editor,Production:Eve Malakoff-Klein Designer:Stephen Druding Compositor:Aptara Seventh Edition Copyright © 2010,2006,2001,1997,1992,1987,1980 Lippincott Williams & Wilkins,a Wolters Kluwer business. 351 West Camden Street 530 Walnut Street Baltimore, MD 21201 Philadelphia, PA 19106 Printed in China All rights reserved.This book is protected by copyright.No part of this book may be reproduced or transmitted in any form or by any means,including as photocopies or scanned-in or other electronic copies,or utilized by any information storage and retrieval system without written permission from the copyright owner,except for brief quotations embodied in critical articles and reviews.Materials appearing in this book prepared by individuals as part of their official duties as U.S.government employees are not covered by the above-mentioned copyright.To request permission,please contact Lippincott Williams & Wilkins at 530 Walnut Street,Philadelphia,PA 19106,via email at permis- [email protected],or via website at lww.com (products and services). 9 8 7 6 5 4 3 2 1 Library of Congress Cataloging-in-Publication Data Snell,Richard S. Clinical neuroanatomy / Richard S.Snell.— 7th ed. p.;cm. Includes bibliographical references and index. ISBN 978-0-7817-9427-5 1. Neuroanatomy. I.Title. [DNLM:1. Nervous System—anatomy & histology.WL 101 S671c 2010] QM451.S64 2010 616.8—dc22 2008040897 DISCLAIMER Care has been taken to confirm the accuracy of the information present and to describe gen- erally accepted practices.However,the authors,editors,and publisher are not responsible for errors or omissions or for any consequences from application of the information in this book and make no warranty,expressed or implied,with respect to the currency,completeness,or accuracy of the con- tents of the publication.Application of this information in a particular situation remains the profes- sional responsibility of the practitioner;the clinical treatments described and recommended may not be considered absolute and universal recommendations. 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To purchase additional copies of this book,call our customer service department at (800) 638-3030 or fax orders to (301) 223-2320.International customers should call (301) 223-2300. Visit Lippincott Williams & Wilkins on the Internet:http://www.lww.com.Lippincott Williams & Wilkins customer service representatives are available from 8:30 am to 6:00 pm,EST. LWBK124-392G-FM[i-xviii] 10/17/08 8:16 AM Page iii Aptara (PPG-Quark) P R E F A C E T his book contains the basic neuroanatomical facts will encounter when making a diagnosis and treating a necessary for the practice of medicine. It is suitable patient. It also provides the information necessary to for medical students, dental students, nurses, and understand many procedures and techniques and notes allied health students. Residents fnd this book use- the anatomical “pitfalls” commonly encountered. ful during their rotations. ● Clinical Problem Solving.This section provides the stu- The functional organization of the nervous system has dent with many examples of clinical situations in which a been emphasized and indicates how injury and disease can knowledge of neuroanatomy is necessary to solve clinical result in neurologic deficits. The amount of factual infor- problems and to institute treatment; solutions to the prob- mation has been strictly limited to that which is clini- lems are provided at the end of the chapter. cally important. ● Review Questions. The purpose of the questions is In this edition, the content of each chapter has been threefold: to focus attention on areas of importance, to reviewed, obsolete material has been discarded, and new enable students to assess their areas of weakness, and to material added. provide a form of self-evaluation when questions are Each chapter is divided into the following categories: answered under examination conditions. Some of the questions are centered around a clinical problem that requires a neuroanatomical answer. Solutions to the prob- ● Clinical Example. A short case report that serves to lem are provided at the end of each chapter. dramatize the relevance of neuroanatomy introduces each chapter. In addition to the full textfrom the book, an interactive ● Chapter Objectives.This section details the material that Review Test, including over 450 questions, is provided is most important to learn and understand in each chapter. online. ● Basic Neuroanatomy. This section provides basic infor- The book is extensively illustrated. The majority of the fig- mation on neuroanatomical structures that are of clinical ures have been kept simple and are in color. As in the previ- importance. Numerous examples of normal radiographs, ous edition, a concise Color Atlas of the dissected brain is CT scans, MRIs, and PET scans are also provided. Many included prior to the text. This small but important group of cross-sectional diagrams have been included to stimulate colored plates enables the reader to quickly relate a particu- students to think in terms of three-dimensional anatomy, lar part of the brain to the whole organ. which is so important in the interpretation of CT scans and References to neuroanatomical literature are included MRI images. should readers wish to acquire a deeper knowledge of an ● Clinical Notes. This section provides the practical appli- area of interest. cation of neuroanatomical facts that are essential in clini- cal practice. It emphasizes the structures that the physician R.S.S. iii LWBK124-392G-FM[i-xviii] 10/17/08 8:16 AM Page iv Aptara (PPG-Quark) A C K N O W L E D G M E N T S I am greatly indebted to the following colleagues who grateful to Dr. G. Size of the Department of Radiology at Yale provided me with photographic examples of neu- University Medical Center for examples of CT scans and MRI roanatomical material: Dr. N. Cauna, Emeritus images of the brain. I also thank Dr. H. Dey, Director of the Professor of Anatomy, University of Pittsburgh School PET Scan Unit of the Department of Radiology, Veterans of Medicine; Dr. F. M. J. Fitzgerald, Professor of Anatomy, Affairs Medical Center, West Haven, Connecticut, for several University College, Galway, Ireland; and Dr. A. Peters, examples of PET scans of the brain. I thank the medical pho- Professor of Anatomy, Boston University School of Medicine. tographers of the Department of Radiology at Yale for their My special thanks are owed to Larry Clerk, who, as a sen- excellent work in reproducing the radiographs. ior technician in the Department of Anatomy at the George As in the past, I express my sincere thanks to Myra Washington University School of Medicine and Health Feldman and Ira Grunther, AMI, for the preparation of the Sciences, greatly assisted me in the preparation of neu- very fine artwork. roanatomical specimens for photography. Finally, to the staff of Lippincott Williams & Wilkins, I I am also grateful to members of the Department of again express my great appreciation for their continued Radiology for the loan of radiographs and CT scans that have enthusiasm and support throughout the preparation of this been reproduced in different sections of this book. I am most book. To My Students—Past,Present,and Future This book is designed so that the information is presented without masses of confusing detail involving complicated neural connections.The arrangement permits the students and future health providers to quickly recall the essential features necessary for the diagnosis and treatment of patients. iv LWBK124-392G-FM[i-xviii] 10/17/08 8:16 AM Page v Aptara (PPG-Quark) C O N T E N T S Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .iv Color Atlas of Brain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .xi CHAPTER 1 Introduction and Organization of the Nervous System 1 Chapter Objectives 2 Central and Peripheral Nervous Systems 2 Major Divisions of the Central Nervous System 2 Major Divisions of the Peripheral Nervous System 10 Early Development of the Nervous System 14 Clinical Notes 17 Clinical Problem Solving 28 Answers and Explanations to Clinical Problem Solving 29 Review Questions 30 Answers and Explanations to Review Questions 31 Additional Reading 32 CHAPTER 2 The Neurobiology of the Neuron and the Neuroglia 33 Chapter Objectives 34 Definition of a Neuron 34 Varieties of Neurons 34 Structure of the Neuron 34 Definition of Neuroglia 53 Astrocytes 53 Oligodendrocytes 54 Microglia 57 Ependyma 58 Extracellular Space 59 Clinical Notes 61 Clinical Problem Solving 63 Answers and Explanations to Clinical Problem Solving 64 Review Questions 65 Answers and Explanations to Review Questions 67 Additional Reading 69 CHAPTER 3 Nerve Fibers, Peripheral Nerves, Receptor and Effector Endings, Dermatomes, and Muscle Activity 70 Chapter Objectives 71 Nerve Fibers 71 Peripheral Nerves 80 Conduction in Peripheral Nerves 84 Receptor Endings 86 v LWBK124-392G-FM[i-xviii] 10/17/08 8:16 AM Page vi Aptara (PPG-Quark) vi Contents Effector Endings 95 Segmental Innervation of Skin 100 Segmental Innervation of Muscles 100 Muscle Tone and Muscle Action 101 Summation of Motor Units 103 Muscle Fatigue 104 Posture 104 Clinical Notes 107 Clinical Problem Solving 120 Answers and Explanations to Clinical Problem Solving 123 Review Questions 126 Answers and Explanations to Review Questions 129 Additional Reading 131 CHAPTER 4 The Spinal Cord and the Ascending and Descending Tracts 132 Chapter Objectives 133 A Brief Review of the Vertebral Column 133 Gross Appearance of the Spinal Cord 137 Structure of the Spinal Cord 138 The Ascending Tracts of the Spinal Cord 143 Anatomical Organization 143 Functions of the Ascending Tracts 144 The Descending Tracts of the Spinal Cord 153 Anatomical Organization 153 Functions of the Descending Tracts 154 Corticospinal Tracts 155 Reticulospinal Tracts 157 Tectospinal Tract 158 Rubrospinal Tract 159 Vestibulospinal Tract 159 Olivospinal Tract 160 Descending Autonomic Fibers 160 Intersegmental Tracts 161 Reflex Arc 162 Influence of Higher Neuronal Centers on the Activities of Spinal Reflexes 164 Renshaw Cells and Lower Motor Neuron Inhibition 164 Clinical Notes 165 Clinical Problem Solving 177 Answers and Explanations to Clinical Problem Solving 178 Review Questions 181 Answers and Explanations to Review Questions 183 Additional Reading 185 CHAPTER 5 The Brainstem 186 Chapter Objectives 187 A Brief Review of the Skull 187 The Cranial Cavity 192 Introduction to the Brainstem 196 Gross Appearance of the Medulla Oblongata 197 Internal Structure 198 Gross Appearance of the Pons 206 Internal Structure of the Pons 208 Gross Appearance of the Midbrain 210 Internal Structure of the Midbrain 210 Clinical Notes 217 Clinical Problem Solving 221 Answers and Explanations to Clinical Problem Solving 222 Review Questions 224 Answers and Explanations to Review Questions 227 Additional Reading 229 LWBK124-392G-FM[i-xviii] 10/17/08 8:16 AM Page vii Aptara (PPG-Quark) Contents vii CHAPTER 6 The Cerebellum and Its Connections 230 Chapter Objective 231 Gross Appearance of the Cerebellum 231 Structure of the Cerebellum 231 Cerebellar Cortical Mechanisms 236 Cerebellar Afferent Fibers 237 Cerebellar Efferent Fibers 240 Functions of the Cerebellum 242 Clinical Notes 243 Clinical Problem Solving 245 Answers and Explanations to Clinical Problem Solving 246 Review Questions 247 Answers and Explanations to Review Questions 249 Additional Reading 250 CHAPTER 7 The Cerebrum 251 Chapter Objectives 252 Subdivisions of the Cerebrum 252 Diencephalon 252 General Appearance of the Cerebral Hemispheres 257 Main Sulci 258 Lobes of the Cerebral Hemisphere 260 Internal Structure of the Cerebral Hemispheres 262 Clinical Notes 271 Clinical Problem Solving 277 Answers and Explanations to Clinical Problem Solving 278 Review Questions 279 Answers and Explanations to Review Questions 281 Additional Reading 283 CHAPTER 8 The Structure and Functional Localization of the Cerebral Cortex 284 Chapter Objective 285 Structure of the Cerebral Cortex 285 Mechanisms of the Cerebral Cortex 287 Cortical Areas 288 Cerebral Dominance 295 Clinical Notes 296 Clinical Problem Solving 298 Answers and Explanations to Clinical Problem Solving 299 Review Questions 300 Answers and Explanations to Review Questions 302 Additional Reading 303 CHAPTER 9 The Reticular Formation and the Limbic System 304 Chapter Objective 305 Reticular Formation 305 Limbic System 307 Clinical Notes 312 Clinical Problem Solving 312 Answers and Explanations to Clinical Problem Solving 313 Review Questions 313 Answers and Explanations to Review Questions 314 Additional Reading 315 CHAPTER 10 The Basal Nuclei (Basal Ganglia) and Their Connections 316 Chapter Objective 317 Terminology 317 LWBK124-392G-FM[i-xviii] 10/17/08 8:16 AM Page viii Aptara (PPG-Quark) viii Contents Corpus Striatum 317 Amygdaloid Nucleus 319 Substantia Nigra and Subthalamic Nuclei 319 Claustrum 319 Connections of the Corpus Striatum and Globus Pallidus 319 Connections of the Corpus Striatum 319 Connections of the Globus Pallidus 319 Functions of the Basal Nuclei 320 Clinical Notes 322 Clinical Problem Solving 327 Answers and Explanations to Clinical Problem Solving 327 Review Questions 327 Answers and Explanations to Review Questions 329 Additional Reading 329 CHAPTER 11 The Cranial Nerve Nuclei and Their Central Connections and Distribution 331 Chapter Objective 332 The 12 Cranial Nerves 332 Organization of the Cranial Nerves 332 Olfactory Nerves (Cranial Nerve I) 335 Optic Nerve (Cranial Nerve II) 336 Oculomotor Nerve (Cranial Nerve III) 340 Trochlear Nerve (Cranial Nerve IV) 340 Trigeminal Nerve (Cranial Nerve V) 341 Abducent Nerve (Cranial Nerve VI) 344 Facial Nerve (Cranial Nerve VII) 346 Vestibulocochlear Nerve (Cranial Nerve VIII) 348 Glossopharyngeal Nerve (Cranial Nerve IX) 350 Vagus Nerve (Cranial Nerve X) 352 Accessory Nerve (Cranial Nerve XI) 354 Hypoglossal Nerve (Cranial Nerve XII) 356 Clinical Notes 358 Clinical Problem Solving 363 Answers and Explanations to Clinical Problem Solving 364 Review Questions 365 Answers and Explanations to Review Questions 368 Additional Reading 369 CHAPTER 12 The Thalamus and Its Connections 371 Chapter Objective 372 General Appearances of the Thalamus 372 Subdivisions of the Thalamus 372 Connections of the Thalamus 375 Function of the Thalamus 375 Clinical Notes 378 Clinical Problem Solving 378 Answers and Explanations to Clinical Problem Solving 378 Review Questions 379 Answers and Explanations to Review Questions 380 Additional Reading 381 CHAPTER 13 The Hypothalamus and Its Connections 382 Chapter Objectives 383 The Hypothalamus 383 Hypothalamic Nuclei 383 Afferent Nervous Connections of the Hypothalamus 385