ATOA01 3/17/06 9:53 AM Page i A Textbook of Neuroanatomy ·· ATOA01 3/17/06 9:53 AM Page ii Dedication To my father, Antonios, my mother, Garifalia, and my sister Oursikía for their contribution to my education MAP To my wife, Roseann, my daughter, Jen, and my mother, Mary LPG XXV The brain within its groove Runs evenly and true; But let a splinter swerve, ‘T were easier for you To put the water back When floods have slit the hills, And scooped a turnpike for themselves, And blotted out the mills! Emily Dickinson ·· ATOA01 3/17/06 9:53 AM Page iii A Textbook of Neuroanatomy Maria A. Patestas Associate Professor of Anatomy Department of Anatomy Des Moines University Des Moines, Iowa Leslie P. Gartner Professor of Anatomy Department of Biomedical Sciences Baltimore College of Dental Surgery Dental School University of Maryland at Baltimore Baltimore, Maryland ·· ATOA01 3/17/06 9:53 AM Page iv © 2006 by Blackwell Science Ltd a Blackwell Publishing company BLACKWELL PUBLISHING 350 Main Street, Malden, MA 02148-5020, USA 108 Cowley Road, Oxford OX4 1JF, UK 550 Swanston Street, Carlton, Victoria 3053, Australia The right of Maria A. Patestas and Leslie P. Gartner to be identified as the Authors of this Work has been asserted in accordance with the UK Copyright, Designs, and Patents Act 1988. 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, except as permitted by the UK Copyright, Designs, and Patents Act 1988, without the prior permission of the publisher. First published 2006 by Blackwell Publishing Ltd Library of Congress Cataloging-in-Publication Data Patestas, Maria A. A textbook of neuroanatomy / Maria A. Patestas and Leslie P. Gartner. p. cm. Includes bibliographical references and index. ISBN 1-4051-0340-X (paperback : alk. paper) 1. Neuroanatomy. I. Gartner, Leslie P. II. Title. QM451.G285 2004 611.8–dc22 2003026897 A catalogue record for this title is available from the British Library. Set in 9/11.5pt Palatino by Graphicraft Limited, Hong Kong Printed and bound in the United Kingdom by Replika, India The publisher’s policy is to use permanent paper from mills that operate a sustainable forestry policy, and which hasbeen manufactured from pulp processed using acid-free and elementary chlorine-free practices. Furthermore, the publisher ensures that the text paper and cover board used have met acceptable environmental accreditation standards. For further information on Blackwell Publishing, visit our website: http://www.blackwellpublishing.com ·· ATOA01 3/17/06 9:53 AM Page v Contents Preface viii Follow-up to clinical case 67 Questions to ponder 67 Part 1 General Principles of the Chapter 6 Gross Anatomy of the Brain 68 Nervous System 1 Clinical case 68 Cerebrum 69 Chapter 1 Introduction to the Nervous System 3 Diencephalon 77 Cells of the nervous system 3 Cerebellum 77 Central nervous system 5 Brainstem 80 Peripheral nervous system 7 Clinical considerations 82 Questions to ponder 9 Synonyms and eponyms 82 Follow-up to clinical case 82 Chapter 2 Development of the Nervous System 10 Questions to ponder 83 Clinical case 10 Chapter 7 Meninges and Cerebrospinal Fluid 84 Early development 11 Neurulation 12 Clinical case 84 Early development of the spinal cord and brain 18 Cranial meninges 85 Development of the spinal cord 19 Spinal meninges 92 Development of the brain 21 Venous sinuses of the cranial dura mater 93 Clinical considerations 26 Cerebrospinal fluid 95 Synonyms and eponyms 27 Ventricles of the brain 95 Follow-up to clinical case 27 Clinical considerations 97 Questions to ponder 27 Synonyms and eponyms 97 Follow-up to clinical case 97 Questions to ponder 98 Chapter 3 Histophysiology of the Nervous System 28 Clinical case 28 Chapter 8 Vascular Supply of the Central Neurons 29 Nervous System 99 Neuroglia 35 Clinical case 99 Generation and conduction of nerve impulses 38 Vascular supply of the spinal cord 100 Clinical considerations 42 Arterial supply of the brain 101 Synonyms and eponyms 43 Venous drainage of the brain 112 Follow-up to clinical case 43 Clinical considerations 116 Questions to ponder 43 Synonyms and eponyms 117 Follow-up to clinical case 117 Chapter 4 Neurotransmitter Substances 44 Questions to ponder 117 Clinical case 44 Classification of neurotransmitter substances 47 Chapter 9 Autonomic Nervous System 118 Clinical considerations 52 Clinical case 118 Follow-up to clinical case 53 Sympathetic nervous system 119 Questions to ponder 53 Parasympathetic nervous system 126 Enteric nervous system 129 Chapter 5 Spinal Cord 54 Neurotransmitters and receptors of the Clinical case 54 autonomic nervous system 130 Morphology of the spinal cord 55 Pelvic autonomic functions 131 Internal morphology of the spinal cord 60 Clinical considerations 132 Vascular supply of the spinal cord 63 Synonyms and eponyms 133 Clinical considerations 66 Follow-up to clinical case 133 Synonyms and eponyms 67 Questions to ponder 133 ·· ATOA01 3/17/06 9:53 AM Page vi vi === CONTENTS Part 2 Integrative Components of the Chapter 14 Reticular Formation 241 Nervous System 135 Clinical case 241 Morphology of the reticular formation 241 Zones of the reticular formation 242 Chapter 10 Ascending Sensory Pathways 137 Nuclei associated with the reticular formation 245 Clinical case 137 Input to and output from the reticular formation 245 Sensory receptors 138 Functions of the reticular formation 245 Anterolateral system 146 Clinical considerations 250 Tactile sensation and proprioception 154 Synonyms and eponyms 251 Sensory pathways to the cerebellum 157 Follow-up to clinical case 251 Clinical considerations 161 Questions to ponder 252 Modulation of nociception 166 Neuroplasticity 168 Chapter 15 Cranial Nerves 253 Synonyms and eponyms 169 Clinical case 253 Follow-up to clinical case 170 Olfactory nerve (CN I) 256 Questions to ponder 170 Optic nerve (CN II) 256 Oculomotor nerve (CN III) 258 Chapter 11 Motor Cortex and Descending Trochlear nerve (CN IV) 260 Motor Pathways 171 Trigeminal nerve (CN V) 261 Clinical case 171 Abducent nerve (CN VI) 266 Cortical areas controlling motor activity 172 Facial nerve (CN VII) 268 Descending motor pathways 174 Vestibulocochlear nerve (CN VIII) 272 Clinical considerations 187 Glossopharyngeal nerve (CN IX) 272 Synonyms and eponyms 188 Vagus nerve (CN X) 274 Follow-up to clinical case 189 Spinal accessory nerve (CN XI) 277 Questions to ponder 189 Hypoglossal nerve (CN XII) 277 Synonyms and eponyms 280 Chapter 12 Basal Ganglia 190 Follow-up to clinical case 280 Clinical case 190 Questions to ponder 281 Components of the basal ganglia 191 Nuclei associated with the basal ganglia 196 Chapter 16 Visual System 282 Input, intrinsic, and output nuclei of the basal ganglia 196 Clinical case 282 Connections of the basal ganglia 198 Eye 282 Circuits connecting the basal ganglia, thalamus, Central visual pathways 287 and cerebral cortex 205 Visual reflexes 291 Other circuits of the basal ganglia 207 Clinical considerations 298 Neurotransmitters of the basal ganglia 208 Synonyms and eponyms 302 “Direct” and “indirect” loops (pathways) of the Follow-up to clinical case 302 basal ganglia 209 Questions to ponder 303 Circuits that modulate activity of the basal ganglia 210 Clinical considerations 212 Chapter 17 Auditory System 304 Synonyms and eponyms 217 Clinical case 304 Follow-up to clinical case 218 Ear 304 Questions to ponder 218 Auditory transmission 308 Central auditory pathways 309 Chapter 13 Cerebellum 219 Clinical considerations 315 Clinical case 219 Synonyms and eponyms 316 Morphology of the cerebellum 220 Follow-up to clinical case 316 Cerebellar peduncles 227 Questions to ponder 317 Deep cerebellar nuclei 229 Afferents (input) to the cerebellum 231 Chapter 18 Vestibular System 318 Efferents (output) from the cerebellum 235 Clinical case 318 Functional organization of the cerebellum: Vestibular apparatus 319 intrinsic circuitry 236 Vestibular nerve (CN VIII) 325 Clinical considerations 238 Central pathways of the vestibular system 326 Synonyms and eponyms 240 Control of ocular movements 329 Follow-up to clinical case 240 Vestibular nystagmus 332 Questions to ponder 240 Caloric nystagmus 334 ·· ATOA01 3/17/06 9:53 AM Page vii CONTENTS === vii Synonyms and eponyms 334 Clinical considerations 382 Follow-up to clinical case 334 Synonyms and eponyms 383 Questions to ponder 335 Follow-up to clinical case 383 Questions to ponder 384 Chapter 19 Olfactory System 336 Clinical case 336 Chapter 22 Thalamus 385 Olfactory receptor cells 336 Clinical case 385 Olfactory transduction 338 Borders 385 Olfactory nerve (CN I) 338 Anatomy 385 Central connections of the olfactory system 339 Internal and external medullary laminae 388 Clinical considerations 342 Thalamic nuclei 389 Synonyms and eponyms 342 Clinical considerations 395 Follow-up to clinical case 342 Synonyms and eponyms 396 Questions to ponder 343 Follow-up to clinical case 396 Questions to ponder 397 Chapter 20 Limbic System 344 Chapter 23 Cerebral Cortex 398 Clinical case 344 Clinical case 398 Limbic lobe 345 Cells of the cerebral cortex 400 Brainstem centers associated with limbic Types of cortex 402 system function 355 Cell layers of the neocortex 402 Pathways of the limbic system 356 Vertical columnar organization of the cerebral cortex 404 Clinical considerations 358 Afferents (input) to the cerebral cortex 404 Synonyms and eponyms 359 Efferents (output) from the cerebral cortex 404 Follow-up to clinical case 359 Internal capsule and corona radiata 406 Questions to ponder 360 Lobes of the cerebral cortex 408 Functional areas of the cerebral cortex 408 Chapter 21 Hypothalamus 361 Cerebral dominance 417 Clinical case 361 Clinical considerations 417 Borders 362 Synonyms and eponyms 421 Hypothalamic zones and component nuclei 362 Follow-up to clinical case 422 Hypothalamic regions (areas) and component nuclei 368 Questions to ponder 423 Connections of the hypothalamus 370 Pathways of the hypothalamus 370 Questions to ponder: answers to odd questions 425 Functions of the hypothalamus 374 Hypothalamohypophyseal connections 376 Index 435 ·· ATOA01 3/17/06 9:53 AM Page viii Preface This Textbook of Neuroanatomywas written with the student • clinical cases open each chapter, setting the stage for the in mind in full knowledge of the apprehension with which relevance of that chapter’s context; he or she faces the prospect of learning a subject matter that • key points in the chapter are highlighted in the text; frequently intimidates the student population. The material • clinical case questions indicate the relevance of the chap- appears daunting, not only due to its very nature of being ter opening case at key points in the chapter—the answers associated with polysyllabic words, but also because there appear on the related website; are so many eponyms and synonyms for the myriad of terms • summary tables within each chapter function as study that are necessary for the proper description of the nervous guides to assist students in learning and memorizing; system. The good news is that neuroanatomy is simply being • clinical considerations sections indicate the medical con- maligned by its nefarious reputation, for it is a logical, relat- ditions relevant to the chapter topics; ively benign subject, whose comprehension does not require • synonym/eponym tables help organize the many possi- Einsteinian brilliance, but merely a willingness to learn the ble terms for each vocabulary word; meaning of a large fistful of new words and the ability to read • a follow-up to each clinical case is given at the end of a map and follow it from one place to another. In order to each chapter and discusses the opening case, helping tie help the student easily achieve the goal of learning neuro- together the text and its medical application; anatomy we made this textbook complete, concise, yet easy • questions to ponder at the end of each chapter reinforce to read, and well illustrated with many schematic diagrams the relevance of the material, with the answers to even to expound upon the concepts being discussed. Presenting questions given on the website and the answers to odd the material in a relevant context will make the learning questions appearing at the end of the book; experience simpler, more enjoyable, and more memorable. • an accompanying website includes all the illustrations, We envisioned writing a book that was accessible and the even answers for questions to ponder, and all the clear—one that students would not have to wade through in answers for clinical case questions. It also features anima- order to discern key information of human neuroanatomy. tions of key processes and links to useful sources. The site We have also highlighted the interrelationships between sys- can be found at www.blackwellpublishing.com/patestas. tems, structures, and the rest of the body as we go through An instructor’s CD-ROM version is also available. the various regions of the brain. We think a broad under- standing of neuroanatomy, supported with a basic under- standing of its physiology, is critical so that students do not ACKNOWLEDGMENTS get bogged down memorizing structures but instead learn the principles. This conceptual basis of the “big picture” We would like to thank the many individuals who helped helps establish the foundation for future studies in the health us bring this project to fruition, including our editors sciences and for the logic that helps drive it. at Blackwell Publishing: Nancy Whilton, our publisher, who The text is divided into two sections. The first nine chap- signed and oversaw the publishing process; Elizabeth Frank, ters provide an overview of neuroanatomy that introduces our assistant editor, who managed the flow and review of terms, and should be viewed as the vocabulary lessons which multiple drafts; our desk editor, Jane Andrew, who scrut- are necessary evils in the mastering of a new language. For no inized the fine detail; and our production editors Rosie matter how well one understands the grammar of a foreign Hayden, Sarah Edwards, and Cee Pike, who ably managed language, it is the possession of a rich vocabulary that the complex physical construction of this project. Our artist, permits one to communicate with speakers of that tongue. Todd Smith, worked closely with us to ensure a first-rate art The second part of this textbook, Chapters 10–23, utilizes the program, and Dr Scott Thompson wrote the clinical cases that vocabulary of the first part to detail information concerning begin and close each chapter. the various pathways and discrete systems that act in concert Additionally, we would like to thank the many reviewers to perform the myriad of functions of the human nervous who commented on the manuscript in its development. Dr system. Robert Sikes, in particular, was an invaluable reviewer, who Special featuresinclude: provided specific and thorough comments on many draft chapters, and the entire art program. We also would like to • chapter opening outlines that provide a quick overview acknowledge the many helpful comments made by the of the chapter content and organizational logic; reviewers: Stephen Lahr, Curt Anderson, Melody Harrison, ·· ATOA01 3/17/06 9:53 AM Page ix PREFACE === ix Lyn Turkstra, Angela Ciccia, Denise Johnson, Min Yu, John omissions, and shortcomings is ours. In view of that fact, Stein, Mohammad Jamali, Robert Berry, and Judy Schotland. we welcome criticisms and suggestions for improvement of Their help was instrumental in the crafting of this book. this text. And finally, we would like to thank our respective families for all the time we took away from them in the writing of this Maria A. Patestas ([email protected]) textbook. Leslie P. Gartner ([email protected]) While we appreciate all of the assistance that we received from our editors and colleagues, the responsibility for errors, ··