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Nieuwenhuys · Voogd · van Huijzen The Human Central Nervous System Fourth Edition (cid:2) (cid:3)(cid:4) R. Nieuwenhuys J. Voogd C. van Huijzen The Human Central Nervous System Fourth Edition With 391 Figures (cid:1) (cid:2) Rudolf Nieuwenhuys M.D., Ph.D. Professor emeritus of Neuroanatomy The Netherlands Institute (home) for Neuroscience Papehof 25 Meibergdreef 47 1391 BDAbcoude 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 LibraryofCongressControlNumber:2007926177 ISBN 978-3-540-34684-5 Springer-Verlag Berlin Heidelberg New York ISBN3-540-13441-7 3.AuflageSpringer-VerlagBerlinHeidelbergNewYork This work is subject to copyright.Allrights arereserved,whether thewholeor partofthematerialis con- cerned,specifically therightsoftranslation,reprinting,reuseofillustrations,recitation,broadcasting,repro- ductiononmicrofilmorinanyotherway,andstorageindatabanks.Duplicationofthispublicationorparts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its currentversion,andpermissionforusemustalwaysbeobtainedfromSpringer.Violationsareliableforpro- secutionundertheGermanCopyrightLaw. SpringerisapartofSpringerScience+BusinessMedia springer.com ©SpringerBerlinHeidelberg1978,1981,1988,2008 PrintedinGermany The use of generaldescriptivenames,registered names,trademarks,etc.inthispublicationdoes not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws andregulationsandthereforefreeforgeneraluse. ProductionManager:K.Schwind MedicalEditor:Dr.MariaMagdaleneNabbe Typesetting:K+VFotosatzGmbH,Beerfelden CoverDesign:WMXDesignGmbH,Heidelberg Printingandbinding:StürtzGmbH,Würzburg SPIN11768005 80/7231–5 4 3 2 1 0 Printedonacid-freepaper 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 Formation of the Brain Regions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Histogenesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Fundamental Morphological Pattern . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Spinal Cord . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 Coronal Sections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138 Sections Perpendicular to the Axis of the Brain Stem . . . . . . . . . . . . . . . 153 Sagittal Sections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158 Horizontal Sections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165 Section II Structure of Spinal Cord and Brain Parts . . . . . . . . 175 6 Topography of Spinal Cord, Brain Stem and Cerebellum . . . . . . . 177 Spinal Cord . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177 The Grey Matter of the Cord . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178 Primary Afferents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180 The White Matter of the Cord . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 Introductory Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253 Parcellation of the Thalamus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255 Thalamic and Thalamocortical Circuits . . . . . . . . . . . . . . . . . . . . . . . . . . 259 Ventral Nuclear Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262 Anterior Nuclear Group and Lateral Dorsal Nucleus . . . . . . . . . . . . . . . . 263 Mediodorsal Nucleus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 264 Medial Geniculate Body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 264 Lateral Geniculate Body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265 Lateral Nuclear Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265 Posterior Nuclear Complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266 Intralaminar and Midline Nuclei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266 Functional Aspects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 268 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271 Contents IX 9 Diencephalon: Ventral Thalamus or Subthalamus . . . . . . . . . . . . . . 281 Introductory Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281 Subthalamic Cell Masses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281 Subthalamic Fibre Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 284 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 286 10 Diencephalon: Hypothalamus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289 Introductory Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289 Topography and Subdivision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289 Major Conduction Channels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 290 Functional Aspects 1: Introductory Notes . . . . . . . . . . . . . . . . . . . . . . . . 293 Functional Aspects 2: Organization of Specific Hypothalamic Functional Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 298 Circadian Rhythms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 298 Wakefulness and Sleep . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301 Stress Response . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305 Thermoregulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307 Food Intake . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308 Thirst . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 311 Sexual Behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314 Defensive Behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 320 Functional Aspects 3: Summary and Concluding Remarks . . . . . . . . . . . 322 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323 11 Telencephalon: Introduction and Olfactory System . . . . . . . . . . . . . 337 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337 Olfactory System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341 Gross Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341 Olfactory Receptors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341 Olfactory Bulb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 343 Secondary Olfactory Projections and Their Sites of Termination . . . . . 347 Further Spread of Olfactory Information . . . . . . . . . . . . . . . . . . . . . . . 353 Accessory Olfactory System and Terminal Nerve . . . . . . . . . . . . . . . . . 355 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357 12 Telencephalon: Hippocampus and Related Structures . . . . . . . . . . 361 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361 Topography and Structure of Hippocampus . . . . . . . . . . . . . . . . . . . . . . . 366 Microcircuitry of Ammon’s Horn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372 Intrinsic Connections of the Hippocampus . . . . . . . . . . . . . . . . . . . . . . . 375 Major Fibre Systems Connecting the Hippocampal Formation and Its Dependencies with Other Parts of the Brain . . . . . . . . . . . . . . . . 378 Afferents to the Hippocampus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Efferents from the Hippocampus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 The Circuit of Papez . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 X Contents Synopsis of Anatomical Data: Functional and Clinical Aspect . . . . . . . . 386 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 391 13 Telencephalon: Amygdala and Claustrum . . . . . . . . . . . . . . . . . . . . 401 Amygdala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 401 Topography, Subdivision and Structure . . . . . . . . . . . . . . . . . . . . . . . . 401 Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403 Fibre Systems Related to the Amygdala . . . . . . . . . . . . . . . . . . . . . . . . 404 Afferents to the Amygdala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 404 Intra-Amygdaloid Connections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409 Efferents from the Amygdala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409 The Bed Nucleus of the Stria Terminalis and the “Extended Amygdala” Concept . . . . . . . . . . . . . . . . . . . . . . . . 414 Fibre Connections of the BST . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 415 Functional and Clinical Aspects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 415 Claustrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 417 Topography, Subdivision and Structure . . . . . . . . . . . . . . . . . . . . . . . . 417 Fibre Connections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 417 Circuitry and Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 419 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 420 14 Telencephalon: Basal Ganglia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427 Cell Masses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 429 Fibre connections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 439 Introductory Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 439 The Direct Circuit: Cerebral Cortex – Striatum – Globus Pallidus – Thalamus – Cerebral Cortex; or: Cerebral Cortex – Striatum – Substantia Nigra, pars reticulata – Thalamus – Cortex . . . . . . . . . . . . 439 Subdivision of Direct Striatal Circuit . . . . . . . . . . . . . . . . . . . . . . . . . 444 A Brief Excursion to the Cerebellum . . . . . . . . . . . . . . . . . . . . . . . . . . 447 Thalamostriate Projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 448 The Indirect Circuit: Cerebral Cortex – Striatum – External Segment of Globus Pallidus – Subthalamic Nucleus – Internal Segment of Globus Pallidus and Substantia Nigra, pars reticulata – Thalamus – Cerebral Cortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 449 The Striato-Nigro-Striatal Circuit . . . . . . . . . . . . . . . . . . . . . . . . . . . . 451 Additional Striatal Connections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 454 Input Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 454 Output Channels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 454 Connections of the Ventral Striatum . . . . . . . . . . . . . . . . . . . . . . . . . . 456 Connections of the Substantia Innominata, Basal Nucleus of Meynert and Related Cell Groups . . . . . . . . . . . . . . . . . . . . . . . . . . 460 Clinical Aspects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 468 Introductory Note . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 468 Résumé of the Circuitry of the Basal Ganglia . . . . . . . . . . . . . . . . . . . 468 Disorders of the Basal Ganglia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 471 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474 Contents XI 15 Telencephalon: Neocortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 491 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 491 Sulcal Pattern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 498 Structural and Functional Subdivision of Neocortex . . . . . . . . . . . . . . . . 498 Structural Subdivision 1: Cytoarchitecture . . . . . . . . . . . . . . . . . . . . . . 498 Structural Subdivision 2: Myeloarchitecture . . . . . . . . . . . . . . . . . . . . . 506 Structural Subdivision 3: Myelogenesis . . . . . . . . . . . . . . . . . . . . . . . . 510 Structural Subdivision 4: Connectivity . . . . . . . . . . . . . . . . . . . . . . . . . 510 Functional Subdivision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 516 Structural and Functional Subdivision: Overview . . . . . . . . . . . . . . . . 528 Structural and Functional Localization in the Neocortex: Current Research and Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . 530 Neocortical Afferents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 536 Neocortical Neurons and Their Synaptic Relationships . . . . . . . . . . . . . . 544 Introductory Note . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 544 Typical Pyramidal Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 544 Atypical Pyramidal Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 559 Local Circuit Neurons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 560 Microcircuitry of Neocortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 569 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 569 Networks of Pyramidal Neuron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 570 Interneuronal Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 571 Neocortical Columns and Modules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 575 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 575 The Investigations of Lorente de Nó: Elementary Units and Glomérulos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 576 The Columnar Organization of the Somatosensory Cortex . . . . . . . . . 576 The Columnar Organization of the Visual Cortex . . . . . . . . . . . . . . . . 578 The Auditory Cortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 579 The Motor Cortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 579 Columnar Patterns Shown by the Cells of Origin and the Terminal Ramifications of Cortico-cortical Connections . . . . . . . . . . . . . . . . . . . 579 Minicolumns and the Radial Unit Hypothesis of Cortical Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 581 Dendritic Clusters, Axonal Bundles and Radial Cell Cords As (Possible) Constituents of Neocortical Minicolumns . . . . . . . . . . . . 582 Microcircuitry of Neocortical Columns . . . . . . . . . . . . . . . . . . . . . . . . 586 Neocortical Columns and Modules: A Critical Commentary . . . . . . . . 586 Comparative Aspects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 591 Synopsis of Main Neocortical Regions . . . . . . . . . . . . . . . . . . . . . . . . . . . 592 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 592 Association and Commissural Connections . . . . . . . . . . . . . . . . . . . . . 592 Functional and Structural Asymmetry of the Two Hemispheres . . . . . 599 Occipital Lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 600 Parietal Lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 605 Temporal Lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 611 Limbic Lobe and Paralimbic Belt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 617

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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
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