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R. Nieuwenhuys J. Voogd C. van Huijzen The Human Central Nervous System R. Nieuwenhuys J. Voogd C. van Huijzen The Human Central Nervous System Fourth Edition With 391 Figures 12 Rudolf Nieuwenhuys M.D., Ph.D. Professor emeritus of Neuroanatomy The Netherlands Institute (home) for Neuroscience Papehof 25 Meibergdreef 47 1391 BD Abcoude 1105 BA Amsterdam The Netherlands The Netherlands [email protected] Jan Voogd M.D., Ph.D. Professor emeritus of Anatomy Department of Neuroscience (home) Erasmus University Rotterdam Rhijngeesterstraatweg 1 P.O. Box 2040 2342 AN Oegstgeest 3000 CA Rotterdam The Netherlands The Netherlands [email protected] Christiaan van Huijzen F.M.A.A. Medical Artist (retired) (home) Willem Degenstraat 2 6525 BW Nijmegen The Netherlands Library of Congress Control Number: 2007926177 ISBN 978-3-540-34684-5 Springer-Verlag Berlin Heidelberg New York ISBN 3-540-13441-7 3. Auflage Springer-Verlag Berlin Heidelberg New York This work is subject to copyright. All rights are reserved, whether the whole or part of the material is con- cerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, repro- duction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer. Violations are liable for pro- secution under the German Copyright Law. Springer is a part of Springer Science+Business Media springer.com © Springer Berlin Heidelberg 1978, 1981, 1988, 2008 Printed in Germany The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Production Manager: K. Schwind Medical Editor: Dr. Maria Magdalene Nabbe Typesetting: K+V Fotosatz GmbH, Beerfelden Cover Design: WMX Design GmbH, Heidelberg Printing and binding: Stürtz GmbH, Würzburg SPIN 11768005 80/7231 – 5 4 3210 Printedonacid-free paper Preface to the Fourth Edition The present edition of The Human Central Nervous System differs considerably from its predecessors. In previous editions, the text was essentially confined to a section dealing with the various functional systems of the brain. This section, which has been rewritten and updated, is now preceded by 15 newly written chapters, which introduce the pictorial material of the gross anatomy, the blood vessels and meninges and the microstructure of its various parts and deal with the development, topography and functional anatomy of the spinal cord, the brain stem and the cerebellum, the diencephalon and the telencephalon. Great pains have been taken to cover the most recent concepts and data. As suggested by the front cover, there is a focus on the evolutionary development of the human brain. Throughout the text numerous correlations with neuropathology and clinical neu- rology have been made. After much thought, we decided to replace the full Latin terminology, cherished in all previous editions, with English and Anglicized Latin terms. It has been an emotional farewell from beautiful terms such as decussatio hipposideriformis Wer- nekinkii and pontes grisei caudatolenticulares. Not only the text, but also the pic- torial material has been extended and brought into harmony with the present state of knowledge. More than 230 new illustrations have been added and many others have been revised. The number of macroscopical sections through the brain has been extended considerably. Together, these illustrations now comprise a complete and convenient atlas for interpreting neuroimaging studies. The bibliography increased from 272 references in the first and second edi- tions, via 1553 references in the third edition, to almost 4000 references in the present one. Therefore, the book has grown again considerably; however, its aim – to provide a straightforward, clear and reliable guide to the structural and func- tional organization of the human central nervous system for both apprentices and specialists in the neurological sciences – has not changed. During the preparation of this and previous editions we received advice and help from many persons; hence, our gratitude is manifold. Professor Luis Puelles and Dr. Iain H.M. Smart were so kind as to critically read the chapter on the de- velopment of the central nervous system. Professor Jean Büttner-Ennever read and offered valuable advice for changes in the chapter on the visual system. The many consultations we had with Professor Harry B.M. Uylings on the structure and subdivision of the cerebral cortex were also of considerable help to us. Pro- fessor Jaak Duysens gave valuable suggestions for the section on Locomotion of Chapter 21, Motor systems. VI Preface to the Fourth Edition We would also like to acknowledge the artists: Mr. Wil P.M. Maas, who pre- pared numerous line drawings, and Mr. Gerben van der Meulen, who gave many figures their final shape and also aided in labelling them. Mrs. Jenneke Kruis- brink, librarian of the Netherlands Institute of Neuroscience assisted in retrieving the literature. The secretarial assistance afforded by Mrs. Inge Eijkhout and the reference management of Mrs. Irene S. Benne are especially acknowledged. In concluding this preamble, the invaluable moral and practical support of Mrs. Suzanne Bakker M.Sc. in all phases of this project is gratefully acknowledged and, finally, we extend our most sincere thanks to our publisher, Springer-Verlag, and their staff – especially Dr. Maria Magdalene Nabbe, Mrs. Sherryl Sundell, and Mr. Klemens Schwind, for their kind help during the preparation of this book. October 2007 Rudolf Nieuwenhuys Jan Voogd Chris van Huijzen Contents Section I Orientation, Development, Gross Anatomy, Blood Supply and Meninges ...................... 1 1 Orientation ............................................... 3 References ................................................. 3 2 Development ............................................. 7 Early Morphogenesis ........................................ 7 FormationoftheBrainRegions................................ 9 Histogenesis ............................................... 16 Fundamental Morphological Pattern ............................ 24 SpinalCord................................................ 26 Hindbrain ................................................. 29 Cerebellum................................................ 32 Midbrain .................................................. 39 Forebrain ................................................. 41 Diencephalon .............................................. 44 Telencephalon .............................................. 48 New Insights into the Development of Forebrain . ................. 53 References ................................................. 59 3 Gross Anatomy ........................................... 67 References ................................................. 71 4 Blood Supply, Meninges and Cerebrospinal Fluid Circulation ... 95 Introduction............................................... 95 Arteries of the Brain ......................................... 95 Meninges, Cisterns and Cerebrospinal Fluid Circulation . ........... 97 Circumventricular Organs .................................... 99 Veins of the Brain ........................................... 99 Vessels and Meninges of the Spinal Cord . ....................... 100 References ................................................. 101 VIII Contents 5 Brain Slices .............................................. 137 Introduction .............................................. 137 CoronalSections........................................... 138 Sections Perpendicular to the Axis of the Brain Stem ............... 153 Sagittal Sections ............................................ 158 HorizontalSections......................................... 165 Section II Structure of Spinal Cord and Brain Parts ........ 175 6 Topography of Spinal Cord, Brain Stem and Cerebellum ....... 177 SpinalCord............................................... 177 Introduction ............................................ 177 The Grey Matter of the Cord ................................ 178 Primary Afferents ........................................ 180 TheWhiteMatteroftheCord............................... 182 Topography of the Brain Stem and the Cerebellum ................. 190 Introduction ............................................ 190 Cranial Nerve Nuclei in the Brain Stem ....................... 190 Medulla Oblongata ........................................ 191 Metencephalon and the Cerebellum ........................... 198 Mesencephalon .......................................... 204 Addendum: Diencephalon and the Corpus Striatum ................ 209 References ................................................ 210 7 Diencephalon: Introduction and Epithalamus ................. 247 Introduction .............................................. 247 Epithalamus ............................................... 247 References ................................................ 250 8 Diencephalon: Dorsal Thalamus ............................ 253 IntroductoryNotes ......................................... 253 Parcellation of the Thalamus .................................. 255 Thalamic and Thalamocortical Circuits .........................