The Human Central Nervous System

967 Pages·2008·3.02 MB·English

by  Rudolf Nieuwenhuys M.D.

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
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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