A Textbook of Neuroanatomy 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 A Textbook of Neuroanatomy Second Edition Maria A. Patestas Professor of Anatomy Department of Anatomy Des Moines University Des Moines, Iowa Leslie P. Gartner Professor of Anatomy (retired) Department of Biomedical Sciences Baltimore College of Dental Surgery Dental School University of Maryland at Baltimore Baltimore, Maryland Copyright © 2016 by John Wiley & Sons, Inc. All rights reserved Published by John Wiley & Sons, Inc., Hoboken, New Jersey Published simultaneously in Canada 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, scanning, or otherwise, except as permitted under Section 107 or 108 of the 1976 United States Copyright Act, without either the prior written permission of the Publisher, or authorization through payment of the appropriate per-copy fee to the Copyright Clearance Center, Inc., 222 Rosewood Drive, Danvers, MA 01923, (978) 750-8400, fax (978) 750 4470, or on the web at www.copyright.com. Requests to the Publisher for permission should be addressed to the Permissions Department, John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, (201) 748-6011, fax (201) 748-6008, or online at http://www.wiley.com/go/permission. Limit of Liability/Disclaimer of Warranty: While the publisher and author have used their best efforts in preparing this book, they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose. No warranty may be created or extended by sales representatives or written sales materials. The advice and strategies contained herein may not be suitable for your situation. You should consult with a professional where appropriate. Neither the publisher nor author shall be liable for any loss of profit or any other commercial damages, including but not limited to special, incidental, consequential, or other damages. For general information on our other products and services or for technical support, please contact our Customer Care Department within the United States at (800) 762-2974, outside the United States at (317) 572 3993 or fax (317) 572-4002. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic formats. For more information about Wiley products, visit our web site at www.wiley.com. Library of Congress Cataloging-in-Publication Data Names: Patestas, Maria Antoniou, author. | Gartner, Leslie P., 1943- , author. Title: A textbook of neuroanatomy / Maria A. Patestas, Leslie P. Gartner. Description: Second edition. | Hoboken, New Jersey : John Wiley & Sons Inc., [2016] | Includes index. Identifiers: LCCN 2015039616 | ISBN 9781118677469 (pbk.) Subjects: | MESH: Nervous System–anatomy & histology. Classification: LCC QM451 | NLM WL 101 | DDC 611.8–dc23 LC record available at http://lccn.loc.gov/2015039616 10 9 8 7 6 5 4 3 2 1 Contents Preface viii Chapter 7 Brainstem 95 About the Companion Website x Internal Organization of the Brainstem 95 Medulla 99 Part I General principles of the nervous system 1 Pons 105 Midbrain 110 Chapter 1 Introduction to the Nervous System 3 Synonyms and Eponyms of the Brainstem 119 Cells of the Central Nervous System 5 Questions to Ponder 119 Central Nervous System 5 Peripheral Nervous System 8 Chapter 8 Meninges and Cerebrospinal Fluid 120 Questions to Ponder 9 Cranial Meninges 121 Spinal Meninges 128 Chapter 2 Development of the Nervous System 10 Venous Sinuses of the Cranial Dura Mater 129 Early Development 11 Cerebrospinal Fluid 133 Neurulation 13 Ventricles of the Brain 133 Early Development of the Spinal Cord and Brain 18 Synonyms and eponyms of the Nervous System 135 Development of the Spinal Cord 20 Questions to Ponder 135 Development of the Brain 22 Synonyms and Eponyms of the Nervous System 29 Chapter 9 Vascular Supply of the Central Questions to Ponder 29 Nervous System 136 Vascular Supply of the Spinal Cord 137 Chapter 3 Histophysiology of the Nervous System 30 Arterial Supply of the Brain 138 Neurons 31 Venous Drainage of the Brain 151 Neuroglia 38 Synonyms and Eponyms of the Vascular Supply of the Generation and conduction of Nerve Impulses 42 Central Nervous System 155 Synonyms and Eponyms of Nervous System Questions to Ponder 155 Histopathology 46 Questions to Ponder 46 Chapter 10 Autonomic Nervous System 156 Sympathetic Nervous System 159 Chapter 4 Neurotransmitter substances 47 Parasympathetic Nervous System 166 Classification of Neurotransmitter Substances 50 Enteric Nervous System 169 Questions to Ponder 57 Neurotransmitters and Receptors of the Autonomic Nervous System 170 Chapter 5 Spinal cord 58 Pelvic Autonomic Functions 171 Morphology of the Spinal Cord 59 Synonyms and Eponyms of the Autonomic Internal Morphology of the Spinal Cord 65 Nervous System 174 Questions to Ponder 174 Vascular Supply of the Spinal Cord 68 Synonyms and Eponyms of the Spinal Cord 72 Chapter 11 Spinal Reflexes 175 Questions to Ponder 72 Components of Reflexes 175 Chapter 6 Gross Anatomy of the Brain 73 Lower Motoneurons 176 Skeletal Muscle Innervation 177 Cerebrum 74 Skeletal Muscle Receptors 177 Diencephalon 85 Muscle Stretch Reflex 179 Cerebellum 86 Reciprocal Inhibition 179 Brainstem 89 Autogenic Inhibition (Inverse Myotatic Reflex) 180 Synonyms and Eponyms of the Brain 94 Flexor Reflex (Withdrawal Reflex, Nociceptive Reflex) 180 Questions to Ponder 94 vi ● ● ● CONTENTS Crossed Extension Reflex 181 Nuclei Associated with the Reticular Formation 307 Maintenance of Muscle Tone via the Gamma Loop 181 Input to and Output from the Reticular Formation 307 Alpha-Gamma Coactivation 182 Functions of the Reticular Formation 308 Synonyms and Eponyms of the Spinal Reflexes 184 Synonyms and Eponyms of the Reticular Formation 314 Questions to Ponder 184 Questions to Ponder 314 Part II Integrative Components of the Nervous Chapter 17 Cranial Nerves 315 System 185 Olfactory Nerve (CN I) 320 Optic Nerve (CN II) 320 Oculomotor Nerve (CN III) 321 Chapter 12 Ascending sensory Pathways 187 Trochlear Nerve (CN IV) 323 Sensory Receptors 188 Trigeminal Nerve (CN V) 325 Anterolateral System 197 Abducent Nerve (CN VI) 332 Tactile Sensation and Proprioception 207 Facial Nerve (CN VII) 336 Sensory Pathways to the Cerebellum 212 Vestibulocochlear Nerve (CN VIII) 338 Modulation of Nociception 224 Glossopharyngeal Nerve (CN IX) 339 Neuroplasticity 226 Vagus Nerve (CN X) 342 Synonyms and Eponyms of the Ascending Sensory Spinal Accessory Nerve (CN XI) 345 Pathways 226 Hypoglossal Nerve (CN XII) 346 Questions to Ponder 227 Synonyms and Eponyms of the Cranial Nerves 348 Questions to Ponder 349 Chapter 13 Motor Cortex and Descending Motor Pathways 228 Chapter 18 Visual System 350 Cortical Areas Controlling Motor Activity 229 Eye 350 Descending Motor Pathways 231 Central Visual Pathways 355 Synonyms and Eponyms of the Motor Cortex and Visual Reflexes 365 Descending Motor Pathways 248 Synonyms and Eponyms of the Visual System 375 Questions to Ponder 249 Questions to Ponder 375 Chapter 14 Basal Nuclei 250 Chapter 19 Auditory System 376 Components of the Basal Nuclei 251 Ear 376 Nuclei Associated with the Basal Nuclei 255 Auditory Transmission 381 Input, Intrinsic, and Output Nuclei of the Basal Nuclei 257 Central Auditory Pathways 383 Connections of the Basal Nuclei 259 Synonyms and eponyms of the Auditory System 391 Circuits Connecting the Basal Nuclei, Thalamus, and Questions to Ponder 391 Cerebral Cortex 265 Other Circuits of the Basal Nuclei 267 Chapter 20 Vestibular System 392 Neurotransmitters of the Basal Nuclei 268 Vestibular Apparatus 393 “Direct” and “Indirect” Loops (Pathways) of the Basal Vestibular Nerve (CN VIII) 399 Nuclei 270 Central Pathways of the Vestibular System 400 Circuits that Modulate Activity of the Basal Nuclei 271 Control of Ocular Movements 403 Synonyms and Eponyms of the Basal Nuclei 277 Vestibular Nystagmus 406 Questions to Ponder 278 Caloric Nystagmus 408 Synonyms and Eponyms of the Vestibular System 409 Chapter 15 Cerebellum 279 Questions to Ponder 410 Morphology of the Cerebellum 281 Cerebellar Peduncles 289 Chapter 21 Olfactory System 411 Deep Cerebellar Nuclei 291 Olfactory Receptor Cells 412 Afferents (Input) to the Cerebellum 292 Olfactory Transduction 412 Efferents (Output) from the Cerebellum 296 Olfactory Nerve (CN I) 414 Functional Organization of the Cerebellum: Intrinsic Central Connections of the Olfactory System 415 Circuitry 298 Synonyms and Eponyms of the Olfactory System 418 Synonyms and Eponyms of the Cerebellum 302 Questions to Ponder 419 Questions to Ponder 302 Chapter 22 Limbic System 420 Chapter 16 Reticular Formation 303 Limbic Lobe 421 Morphology of the Reticular Formation 304 Brainstem Centers Associated with Limbic Zones of the Reticular Formation 305 System Function 432 CONTENTS ● ● ● vii Pathways of the Limbic System 432 Thalamic Nuclei 469 Limbic Association Cortex 435 Synonyms and Eponyms of the Thalamus 477 Limbic System Input to the Endocrine, Autonomic, Questions to Ponder 478 and Somatic Motor Systems 435 Synonyms and Eponyms of the Limbic System 437 Chapter 25 Cerebral Cortex 479 Questions to Ponder 438 Cells of the Cerebral Cortex 481 Types of Cortex 483 Chapter 23 Hypothalamus 439 Cell layers of the Neocortex 484 Borders 440 Vertical Columnar Organization of the Cerebral Cortex 485 Hypothalamic Zones and Component Nuclei 441 Afferents (Input) to the Cerebral Cortex 485 Hypothalamic Regions (areas) and Component Nuclei 446 Efferents (Output) from the Cerebral Cortex 486 Connections of the Hypothalamus 448 Internal Capsule and Corona Radiata 489 Pathways of the Hypothalamus 449 Lobes of the Cerebral Cortex 489 Functions of the Hypothalamus 452 Functional Organization of the Cerebral Cortex 490 Hypothalamohypophyseal Connections 455 Cerebral Dominance 498 Synonyms and Eponyms of the Hypothalamus 462 Synonyms and Eponyms of the Cerebral Cortex 504 Questions to Ponder 463 Questions to Ponder 506 Chapter 24 Thalamus 465 Questions to Ponder: Answers to Odd Questions 507 Borders 465 Anatomy 467 Index 517 Internal and External Medullary Laminae 468 Preface to the Second Edition It is always gratifying to publish the second edition of a text it is the possession of a rich vocabulary that permits one to book because it is an indication that our efforts were well communicate with speakers of that tongue. The second part received, that our colleagues endorsed our work, and that of this textbook, Chapters 12–25, utilizes the vocabulary of students have been well served by it. This new edition of the first part to detail information concerning the various the Textbook of Neuroanatomy, as indeed was the first, was pathways and discrete systems that act in concert to written with the student in mind in full knowledge of the perform the myriad of functions of the human nervous trepidation with which he or she faces the prospect of learn system. ing a subject matter that appears to be daunting and intimi We retained in this edition the well-liked special features, dating. The material appears overwhelming because of the which include: many eponyms that have been retained and add to the in herent complexity of the nervous system. Happily, for the • chapter opening outlines that provide a quick overview of student the reports of being an overwhelming subject is an the chapter content and organizational logic; unjust reputation, for neuroanatomy is a fascinating topic, • clinical cases in the beginning of each chapter, setting the whose comprehension does not require amazing brilliance, stage for the relevance of that chapter’s context; but merely a willingness to learn new words, the aptitude • key points in the chapter that are highlighted in the text; for reading maps, and following pathways from one place to • clinical case questions that emphasize the relevance of the another. In order to help the student more easily achieve chapter opening case at key points in the chapter – the the goal of learning neuroanatomy we made this textbook answers appear on the related website; complete, concise, yet easy to read, and well-illustrated • summary tables within each chapter that function as study with many schematic diagrams to expound upon the con guides to assist students in learning and memorizing; cepts being discussed. Presenting the material in a relevant • clinical considerations that underscore the medical condi context will make the learning experience simpler, more tions relevant to the topics; enjoyable, and more memorable. • synonym/eponym tables that help organize the many Every chapter of this edition was revised to include possible terms for each vocabulary word; current material and to make it even easier to read than the • a follow-up to each clinical case at the end of each chapter first edition. Two new chapters were added, one concerning that discusses the opening case, helping to correlate the the Brainstem (Chapter 7) and the other Spinal Reflexes text and its medical application; (Chapter 11), in order to stress the importance of those • questions to ponder at the end of each chapter to rein two topics and to enhance the student’s understanding of force the relevance of the material, with the answers to their significance in the functioning of the central nervous even-numbered questions given on the website and the system. answers to odd-numbered questions appearing at the end We continued to highlight the interrelationships between of the book; systems, structures, and the rest of the body as we describe the • an accompanying website that includes all the illustrations, various regions of the brain. We think a broad understanding the even answers for questions to ponder, and all the an of neuroanatomy, supported with a basic understanding of swers for clinical case questions. It also features animations its physiology, is critical so that students instead of merely of key processes and links to useful sources. The site can be memorizing structures learn the principles thus establishing found at www.wiley.com/go/patestas/neuroanatomy2e. the foundation for future studies in the health sciences and for the logical forces that help drive it. As with the first edition, this textbook is divided into ACKNOWLEDGMENTS two sections. The first 11 chapters provide an overview of neuroanatomy that introduces terms and should be We would like to thank the many individuals who helped viewed as the vocabulary lessons that are necessary evils us bring this project to fruition, including our editors at in the mastering of a new language. For no matter how John Wiley & Sons Publishing: Justin Jeffryes, our Editorial well one understands the grammar of a foreign language, Director, who signed and oversaw the publishing process;