THE HUMAN BRAIN An Introduction to Its Functional Anatomy A B C Reproduction of a woodcut in De Homine, 1662, by René Descartes, who thought that the pineal gland was the seat of the soul, monitoring the movement of “animal spirits” in sensory nerves and controlling the movement of animal spirits through motor nerves. Cover image: An axial slice of a whole human brain, cut by the author’s brother Jack about 30 years ago at the University of Colorado Medical School and stained by Pam Eller, using a phenol/copper sulfate technique. The brain, and the brain alone, is the source of our pleasures, joys, laughter, and amusement, as well as our sorrow, pain, grief, and tears. It is especially the organ we use to think and learn, see and hear, to distinguish the ugly from the beautiful, the bad from the good, and the pleasant from the unpleasant. The brain is also the seat of madness and delirium, of the fears and terrors that assail us by night or by day, of sleeplessness, awkward mistakes, and thoughts that will not come, of pointless anxieties, forgetfulness, and eccentricities. Hippocrates, ca. 400 BC The human mind can be described as a slow-clockrate modifi ed-digital machine with multiple distinguishable parallel processing, all working in salt water. Philip Morrison: The mind of the machine, Technology Review75:17, 1973 One of the diffi culties in understanding the brain is that it is like nothing so much as a lump of porridge. R.L. Gregory: Eye and brain: the psychology of seeing,New York, 1966, McGraw-Hill Were I to await perfection, My book would never be fi nished. Tai T’ung, 13th century Chinese scholar: The six scripts: principles of Chinese writing. Quoted in Edmunds LN Jr:Cellular and molecular bases of biological clocks, Sixth Edition, New York, 1988, Springer-Verlag THE HUMAN BRAIN An Introduction to Its Functional Anatomy Sixth Edition JOHN NOLTE, PhD Professor of Cell Biology and Anatomy The University of Arizona College of Medicine Tucson, Arizona THREE-DIMENSIONAL BRAIN RECONSTRUCTIONS by JOHN SUNDSTEN, PhD University of Washington School of Medicine Seattle, Washington 1600 John F. Kennedy Blvd. Ste 1800 Philadelphia, PA 19103-2899 THE HUMAN BRAIN: AN INTRODUCTION TO ISBN: 978-0-323-04131-7 ITS FUNCTIONAL ANATOMY Copyright © 2009, 2002, 1999, 1993, 1988, 1981 by Mosby, Inc., an affi liate of Elsevier Inc. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permissions may be sought directly from Elsevier’s Rights Department: phone: (+1) 215 239 3804 (US) or (+44) 1865 843830 (UK); fax: (+44) 1865 853333; e-mail: [email protected]. You may also complete your request on-line via the Elsevier website at http://www.elsevier.com/permissions. Notice Neither the Publisher nor the Author assumes any responsibility for any loss or injury and/or damage to persons or property arising out of or related to any use of the material contained in this book. It is the responsibility of the treating practitioner, relying on independent expertise and knowledge of the patient, to determine the best treatment and method of application for the patient. The Publisher Library of Congress Cataloging-in-Publication Data Nolte, John. The human brain : an introduction to its functional anatomy / John Nolte.—6th ed. p. ; cm. Includes bibliographical references and index. ISBN 978–0–323–04131–7 (alk. paper) 1. Neuroanatomy. 2. Brain—Anatomy. I. Title. [DNLM: 1. Central Nervous System—physiology. 2. Brain—physiology. WL 300 N798h 2009] QM451.N64 2009 611.8—dc22 2007048134 Acquisitions Editor: Madelene Hyde Developmental Editor: Katie DeFrancesco Publishing Services Manager: Linda Van Pelt Design Direction: Gene Harris Cover Designer: Gene Harris Working together to grow libraries in developing countries Printed in China www.elsevier.com | www.bookaid.org | www.sabre.org Last digit is the print number: 9 8 7 6 5 4 3 2 1 Preface The few years since the publication of the fi fth edition Readings have been updated, but the same philosophy of this book have seen continued rapid growth in our was retained: the intent in choosing references was not knowledge of the structure and function of the brain, only to document old and new fi ndings, but also to and increasingly elegant techniques for studying it. Basic provide access through those references to the vast lit- neuroscience has never been more relevant to clinical erature of neuroscience. For these reasons, the emphasis practice. The normal function of neural structures is is on recent reviews and research papers. Many classic becoming understood in more and more detail, as and scientifi cally more important papers have been are molecular-level details of neurological disorders. omitted, although some older or peripherally related Imaging studies now allow observation of regional articles that make interesting reading are listed; where changes in blood fl ow and metabolism in the brains of the content is not obvious from the title of a reference, subjects as they perform various mental tasks, and annotation was added. reconstruction of white matter pathways in live subjects. Beginning with this edition, an electronic collection This, together with dozens of things that were left undone of images and movies is also available. It includes nearly the last time around and dozens of new things I have all of the images from this book, from the accompanying learned, provided the incentive for a new edition. Essentials of the Human Brain, and from the atlas I tried to follow the same principles used since the designed to complement the book (The Human Brain inception of this book. Although the book has grown a in Photographs and Diagrams, third edition). It also little more in length, a minimum of detail has been includes a series of animations of three-dimensional added. In writing or updating various sections, I tried brain reconstructions produced by Drs. John Sundsten not to include facts simply for their own sake. Rather, and Kathleen Mulligan of the University of Washington. new observations and concepts were added where they My hope is that faculty will fi nd these images and anima- help to illuminate function. Because much of our new tions useful in teaching. knowledge has been made possible by new techniques, The challenge of writing a clear, accurate, up-to-date illustrations of these techniques have been maintained, survey of human neurobiology becomes more formida- updated, and to some extent distributed throughout the ble (though more fascinating) by the year, and it is inevi- book: anatomical labeling and tract-tracing methods, table that I have committed errors of hyperbole, structural and functional magnetic resonance imaging, obfuscation, and ignorance. I welcome the comments and electrophysiological techniques are presented in and suggestions of students and colleagues who use this abundance. I went back over all of the illustrations, tried book. They will improve my knowledge and my teaching, to make them a little more clear and attractive where and make the next edition that much better. possible, and added a few new ones. I also added more structural and clinical images, such as diffusion tensor John Nolte images showing normal white matter anatomy and Tucson, Arizona images illustrating germane cases. A new chapter on April 2008 neural plasticity and repair was added. The Suggested v This page intentionally left blank Acknowledgments As in the case of the fi rst fi ve editions, this revision was visual illusions in Chapter 17; Jim Fujimoto, for the far from a solitary accomplishment, and I am indebted optical coherence tomography images; Steve Ringel, for to many. Friends and colleagues from around the world the muscle biopsies; Wally Welker and www.brain- kindly provided illustrations that are either new in this museum.org, for the comparative neuroanatomy; and edition or carried over from previous ones. The sources Susumu Mori, for most of the wonderful new diffusion are acknowledged, with my gratitude, in the fi gure tensor images (and for helping me understand them). legends, but I especially thank Jay Angevine for giving Thanks to Tom Finger, Ed French, Ted Glattke, Erwin me access to the wonderful Weigert-stained brain sec- Montgomery, Sarah Orjada, Dan Stamer, and Todd Van- tions and for being a mainstay for me here in Arizona in derah for their helpful and valued comments on various discussing things neuroanatomical; John Sundsten, for parts of the manuscript. Thanks also to Katie DeFran- the beautiful three-dimensional reconstructions that cesco and Linda Van Pelt of Elsevier/Mosby for their help appear throughout the book and for the fun we had and their patience. A big thanks to my teaching col- making them; Pam Eller, for preparing the brainstem leagues in Arizona for their enthusiasm and good humor sections used in Chapters 11 through 15 and elsewhere and for new ways of looking at things; and to the stu- and for being my buddy all these years; Dave Asher, for dents of The University of Arizona College of Medicine, the temporal bone sections and for being there for me; who use the book and offer many helpful comments and Ray Carmody, for his unstinting help with clinical images suggestions, and who by their curiosity and caring con- over the last 17 years; Nate McMullen and Allen Bell, for tinue to make teaching fun. Finally, my love and special the light micrographs in Chapter 1 and elsewhere; Drs. thanks to Kathy—it was right, and now you’re back Ennio Pannese and Alan Peters, for the beautiful elec- again.* tron micrographs; Grant Dahmer and Norm Koelling, for many of the brain dissections; Ken Catania, for the mole John Nolte stories and pictures; Pagie Beeson, for the aphasia cases; Tucson, Arizona Beau Lotto and Dale Purves, for some of the stunning April, 2008 *Paraphrased from Full Circle, by Gene Clark. vii This page intentionally left blank Contents 1 Introduction to the Nervous System 1 The nervous system has central and peripheral parts, 1 The principal cellular elements of the nervous system are neurons and glial cells, 2 2 Development of the Nervous System 37 The neural tube and neural crest give rise to the central and peripheral nervous systems, 37 Adverse events during development can cause congenital malformations of the nervous system, 49 3 Gross Anatomy and General Organization of the Central Nervous System 53 The long axis of the CNS bends at the cephalic fl exure, 54 Hemisecting a brain reveals parts of the diencephalon, brainstem, and ventricular system, 54 Humans, relative to other animals, have large brains, 55 Named sulci and gyri cover the cerebral surface, 55 The diencephalon includes the thalamus and hypothalamus, 64 Most cranial nerves are attached to the brainstem, 65 The cerebellum includes a vermis and two hemispheres, 67 Sections of the cerebrum reveal the basal ganglia and limbic structures, 67 Parts of the nervous system are interconnected in systematic ways, 68 4 Meningeal Coverings of the Brain and Spinal Cord 80 There are three meningeal layers: the dura mater, arachnoid, and pia mater, 80 The dura mater provides mechanical strength, 82 The dura mater has an arachnoid lining, 86 Pia mater covers the surface of the CNS, 90 The vertebral canal contains spinal epidural space, 91 Bleeding can open up potential meningeal spaces, 92 Parts of the CNS can herniate from one intracranial compartment into another, 93 5 Ventricles and Cerebrospinal Fluid 99 The brain contains four ventricles, 99 Choroid plexus is the source of most CSF, 103 Imaging techniques allow both the CNS and CSF to be visualized, 110 Disruption of CSF circulation can cause hydrocephalus, 117 6 Blood Supply of the Brain 122 The internal carotid arteries and vertebral arteries supply the brain, 122 Imaging techniques allow arteries and veins to be visualized, 132 Blood fl ow to the CNS is closely controlled, 132 A system of barriers partially separates the nervous system from the rest of the body, 140 Superfi cial and deep veins drain the brain, 141 ix
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