BARR’S The Human Nervous System An Anatomical Viewpoint Tenth Edition BARR’S The Human Nervous System An Anatomical Viewpoint Tenth Edition John A. Kiernan, MB, ChB, PhD, DSc Professor Emeritus Department of Anatomy and Cell Biology The University of Western Ontario London, Canada and Nagalingam Rajakumar, MB, BS, PhD Associate Professor Departments of Psychiatry and Anatomy and Cell Biology The University of Western Ontario London, Canada Acquisitions Editor: Crystal Taylor Product Manager: Jenn Verbiar Marketing Manager: Joy Fisher-Williams Vendor Manager: Bridgett Dougherty Manufacturing Coordinator: Margie Orzech Creative Director: Doug Smock Compositor: Integra Software Services Pvt. Ltd. Tenth Edition Copyright © 2014 Lippincott Williams & Wilkins, a Wolters Kluwer business 351 West Camden Street Two Commerce Square Baltimore, MD 21201 2001 Market Street Philadelphia, PA 19103 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 [email protected], or via Web site at lww.com (products and services). 9 8 7 6 5 4 3 2 1 Library of Congress Cataloging-in-Publication Data Kiernan, J. A. (John Alan) Barr’s the human nervous system: an anatomical viewpoint/John A. Kiernan, MB, ChB, PhD, DSc, professor, Department of Anatomy and Cell Biology, The University of Western Ontario, London, Canada, and Nagalingam Rajakumar, MB, BS, PhD, associate professor, Departments of Psychiatry and Anatomy and Cell Biology, The University of Western Ontario, London, Canada.—Tenth edition. pages cm Includes bibliographical references and index. ISBN 978-1-4511-7327-7 1. Neuroanatomy. I. Barr, Murray Llewellyn, 1908-1995. II. Rajakumar, Nagalingam, 1957- III. Title. IV. Title: Human nervous system. QM451.B27 2014 611'.8—dc23 2012041508 DISCLAIMER Care has been taken to confirm the accuracy of the information present and to describe generally accepted practices. The authors, editors, and publisher, however, 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 contents of the publication. 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P r e f a c e Murray Llewellyn Barr (1908 to 1995) obtained New to the Edition his medical degree in 1933 from the Univer- sity of Western Ontario in London, Canada, In this Tenth Edition, the improvement and and after a few years in practice, he entered the changing of illustrations has continued, and Department of Anatomy at the same institution. nearly all are now colored. The text and recom- He studied and taught neuroanatomy there until mended readings have also been updated. R eaders 1978. This period of service was interrupted of this edition have access to the publisher’s by the Second World War, when he served in Web site—www.lww.thepoint.com—which the Medical Branch of the Royal Canadian Air includes a variety of additional materials. These Force. In 1949, the direction of Barr’s research include labeled and unlabeled versions of all the changed abruptly from neurohistology to cyto- illustrations for instructors, sample exam questions genetics. With Ewart G. (“Mike”) Bertram, who and clinical cases, an extended glossary, and bio- was then a graduate student, he had observed graphical information about researchers and phy- an intranuclear inclusion in neurons of female sicians whose names are associated with anatomy, animals. This was the sex chromatin, now widely physiology, and diseases of the nervous system. known as the Barr body; its discovery was an early landmark in the science of human cytogenetics. Acknowledgments For this and his later work in the field, Murray Barr received more than 30 awards and honors, An important feature in the production of the including the Kennedy Foundation International Tenth Edition has been the publisher’s extensive Award in Mental Retardation, Fellowship of the use of external reviewers. The book is undoubtedly Royal Society of London, the Order of Canada, improved as a result of their recommendations and and 7 honorary doctorates. comments, for which we thank them. It is a plea- Although Barr’s research career was largely con- sure to acknowledge also the helpful advice given cerned with the cytological diagnosis of inherited by present and former colleagues during the writ- diseases, he continued to teach neuroanatomy. The ing of this and the five preceding editions: Drs. J. first edition of this book, published in 1972, was Ronald Doucette, Jonathan Hore, Kost Elisevich, one of the first medium-sized texts in its field. It Brian A. Flumerfelt, Elias B. Gammal, Alan W. was written to make life easier for those approach- Hrycyshyn, Arthur J. Hudson, Peeyush K. Lala, ing neuroscience for the first time, especially Peter Merrifield, D.G. Montemurro, David M. medical students and those in the allied health sci- Pelz, David Ramsay, A. Jon Stoessl, Shannon ences. This objective has not changed, although a Venance, Tutis Vilis, and Chris Watling. We greater variety of students now study the subject. appreciate also the numerous insightful comments Advances in the science necessitated much revi- of Professor Ronan O’Rahilly (Villars-sur-Glâne, sion over the years, and the book became larger Switzerland), especially on matters of embryology with successive editions. With the Eighth and and terminology. Ninth Editions, this trend was reversed, result- The artwork for the first seven editions of ing in a somewhat smaller book. The illustrations the book was prepared by local artists (Margaret were enhanced, however, with more extensive use Corrin, Louise Gadbois, Jeannie Ross, and of colors. Nancy Somerville) supervised by the authors. v vi Preface For this and the previous edition, all the illus- Tenth Edition, and Jennifer Verbiar and Bridgett trations were prepared in electronic form by the Dougherty, including Gordon Hammy Matchado publisher’s art department, Jennifer Clements from Integra Software Services presided over pro- and artist Kim Battista, working from older art- duction of the book. We thank all of them for work and from our sketches. Among other staff their contributions. at Lippincott Williams & Wilkins, Crystal Taylor J. A. KIERNAN arranged the reviewing and the contract for the N. RAJAKumAR London, Canada C o n t e n t s PREfACE v PART I: INTRoDuCTIoN AND NEuRohISTology 1 Development, Composition, and Evolution of the Nervous System 3 2 Cells of the Nervous System 13 3 Peripheral Nervous System 35 4 Imaging Techniques and Neuroanatomical Research methods 49 PART II: REgIoNAl ANAToMy of ThE CENTRAl NERvouS SySTEM 5 Spinal Cord 63 6 Brain Stem: External Anatomy 81 7 Brain Stem: Nuclei and Tracts 89 8 Cranial Nerves 115 9 Reticular formation 143 10 Cerebellum 159 11 Diencephalon 175 12 Corpus Striatum 201 13 Topography of the Cerebral Hemispheres 213 14 Histology of the Cerebral Cortex 221 15 functional Localization in the Cerebral Cortex 229 16 Cerebral White matter and Lateral ventricles 247 17 Olfactory System 261 18 Limbic System: The Hippocampus and the Amygdala 269 PART III: REvIEW of ThE MAJoR SySTEMS 19 General Sensory Systems 285 20 visual System 305 vii viii Contents 21 Auditory System 321 22 vestibular System 333 23 motor Systems 341 24 visceral Innervation 357 PART Iv: BlooD SuPPly AND ThE MENINgES 25 Blood Supply of the Central Nervous System 371 26 meninges and Cerebrospinal fluid 387 GLOSSARy Of NEuROANATOmICAL AND RELATED TERmS 399 INDEx 413