Anatomy SNC Nieuwenhuys

R.
Nieuwenhuys IVoogd
Chr. van Huijzen
The Human
Central Nervous System
A Synopsis and Atlas
Third Revised Edition
With 217 Figures
Springer-Verlag Berlin Heidelberg GmbH

RUDOLF NIEUWENHUYS, M.D., Ph.D.
Professor of Neuroanatomy, Department of Anatomy,
University of Nijmegen, The Netherlands
JAN VOOGD, M.D., Ph.D.

Professor of Anatomy, Department of Anatomy
Erasmus University, Rotterdam, The Netherlands
CHRISTIAAN VAN HUIJZEN, F.M.A.A.

Medical artist, Department of Anatomy,
University of Nijmegen, The Netherlands
1978 First English Edition

1980 German Edition
Italian Edition
1981 Second English Edition
1983 Spanish Edition
Greek Edition
Japanese Edition
ISBN 978-3-540-13441-1
Library of Congress Cataloging in Publication Data. Nieuwenhuys, R., 1927. The human central nervous
system. Bibliography: p. Includes index. 1. Central nervous system - Atlases. 2. Histology - Atlases. 3. Neuro-
anatomy - Atlases. 1. Voogd, J. (Johan), 1933. II. Huijzen, Chr. van. III. Title. [DNLM: 1. Central Nervous
System - anatomy and histology - atlases. WL 17 N682h) QM455.N48 1988 612'.82'0222 88-24868
ISBN 978-3-540-13441-1 ISBN 978-3-662-10343-2 (eBook)
DOI 10.1007/978-3-662-10343-2
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© Springer-Verlag Berlin Heidelberg 1978, 1981, 1988
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Preface to the Third Edition
The present edition of our The Human Central Nervous System: A Synopsis
and Atlas differs in several respects from its predecessor. An entirely new
section on the cerebrovascular system and the meninges has been added, in
accordance with the wishes of many colleagues. The text has been thoroughly
revised and extended in the light of new data and concepts. The functional
significance of the structures discussed and depicted has received more atten-
tion, and numerous correlations with neuropathology and clinical neurology
have been indicated.
The final section in the previous editions was devoted to the monoaminergic
neuron systems. It was our original plan to add sections on other important
transmitter-specified neuronal populations. However, the size of these sections
soon grew well beyond the limits set for the present work. Hence, it was
decided to produce a separate text on that subject, which has appeared in
the mean time (R.NIEUWENHUYS: Chemoarchitecture of the Brain, Springer-
Verlag 1985). The reader who is particularly interested in chemical neuroana-
tomy is referred to that work; numerous data on the nature of the neurotrans-
mitters present in the various centres and fibre systems of the neuraxis are
incorporated in the text of the present book, however.
Not only the text, but also the pictorial material has been extended and
brought into harmony with the present state of knowlegde. Eighty-two new
illustrations have been added and many others have been revised. As in the
previous editions, all of the figures, including the new ones, are original. The
bibliography has been extended from 272 references in the second edition
to 1553 in the present one. Due to all of these additions the book has grown
considerably; however, its aim - to provide a straightforward, clear and reliable
guide to the structural and functional organization of the human central ner-
vous system, for both apprentices and specialists in the neurological sciences
- has remained unchanged.
During the preparation of this and previous editions we received the advice
and help of many persons; hence our gratitude is manifold. The late Dr.
J.H.R. SCHOEN made available the Hiiggqvist material. The ample consulta-
tions we had with Dr. H.O.M. THIJSSEN, Professor of neuroradiology at the
University of Nijmegen, were of considerable help to us. We wish to thank
Mr. A. BINNENDIJK and Mr. C. CORNELISSEN for the preparation of the series
of brain and head slices, Mrs. C. DE VOCHT-POORT, Mrs. P. DRIESSEN-VERIJDT
and the late Mr. J. STINS for the preparation of the histological material,
and Dr. H. FEIRABEND and Mr. W. REIJCHARD for the injected specimens
in part two.
Preface to the Third Edition VI
Acknowledgement is made to the artists: to Mr. T. VAN GERWEN, who

made the half-tone illustrations, to Mr. A. GRUTER, who did the drawings
of the microscopical sections and the line drawings of part two, and to Mr.
W.P.J. MAAS, who prepared the line drawings of part five and also aided
in the labelling of the illustrations. Without their skill and patience this book
would not have been possible.
The invaluable secretarial assistance of Mrs. M. RIECK, E. KLINK, G. VAN
SON-VERSTRAETEN, H. FLIERVOET, J. VELZEBOER, M. VAN DE COEVERING and
W. DE HAAN is especially acknowledged.
Finally, we extend our most sincere thanks to the publishing house of
Springer-Verlag and their staff - especially Mr. B. LEWERICH, Mrs. Th. DEIG-
MOLLER, Mrs. D. GROSSHANS and Mrs. U. PFAFF - for their help during the
preparation of this book.
Summer 1988 R. NIEUWENHUYS

J. VOOGD
CHR. VAN HUIJZEN
Preface to the Second Edition
The particularly good reception enjoyed by our "The Human Central Nervous
System, a Synopsis and Atlas" has made a second edition necessary, hardly
more than two years after its first appearance. This new edition enabled us
to make a number of corrections, but it was judged premature to undertake
a thorough updating of the text. However, a major improvement - suggested
by some reviewers and many colleagues - is that in this new edition the abbre-
viations in the figures have been replaced by the full Latin terms.
We want to emphasize that the study of this book can facilitate and deepen
but never replace the study of the anatomical preparation. Acquaintance with
the basic cytology and histology of nervous tissue has been taken for granted.
This book is evidently often consulted with the aim of looking up a particu-
lar structure together with its name and its topographical relations. This is
certainly one of the purposes of the book. We are, however, of the opinion
that during a systematic study of the figures showing the functional systems
in part V, perusal of the accompanying text will be necessary. As a matter
of fact the spatial representations of the fibre systems are no more than a
visualization of the most salient features discussed in the text. The pictures
are so to speak a snapshot of the current knowledge of a particular functional
system within the central nervous system; no less, but no more either. The
mutual coherence between the pictures in the macroscopical, microscopical
and functional sections of the book, will be readily apparent during the use
of the work. The authors hope that for the readers this coherence will lead
to a better insight into the structure of the human central nervous system.
Summer 1981 R.NIEUWENRUYS

J. VOOGD
CRR. VAN HUIJZEN
Contents
Introduction
Purpose and Plan. . . . . . . . . . . . . . . . . . 1
Material, Techniques, and Preparation of the Illustrations 1
Annotations. . . . . . . 4
Terminology and Labelling. 4
Part I Gross Anatomy

Orientation . . . . . 6
External and Medial Views. 10
Internal Structures . . . . 26
Part II Vessels and Meninges

Arteries of the Brain . . 34
Veins of the Brain 45
Vessels of the Brain Stem 50
Meninges, Cisterns and Liquor System. 54
Vessels and Meninges of the Spinal Cord . 60
Part III Brain Slices

Coronal Sections. . 66
Sections Perpendicular to the Axis of the Brain Stem. 79
Sagittal Sections . . 84
Horizontal Sections. . . . . . 89
Part IV Microscopical Sections

Coronal Sections Through the Basal Part of the Prosencephalon . 96
Transverse Sections Through the Brain Stem and Spinal Cord . 104
Part V Functional Systems

Cranial Nerve Nuclei in the Brain Stem 144
General Sensory Systems and Taste . . 149
Introduction. . . . . . . . . . . 149
Primary Afferents and the Spinal Grey Matter 150
The Anterolateral System . . 151
The Medial Lemniscus System 154
The Somatosensory Cortex. . 154
x Contents
The Trigeminal System . . . . . . . . . . . . . . 155

Descending Connections to Somatosensory Relay Nuclei 160
The Visceral Afferent Systems 164
Special Sensory Systems. . . . . . 165
The Vestibular System. . . . . . 165
The Vestibular Nerve and Nuclei 165
Afferents to the Vestibular Nuclei . 168
Efferents from the Vestibular Nuclei . 169
Projections from the Vestibular Nuclei to the Cerebral Cortex 172
The Auditory System 172
Auditory Centres. . . . . . . . 172
Auditory Pathways . . . . . . . 173
The Ascending Auditory Projection 177
The Descending Auditory Projection 178
Auditory Reflex Pathways 179
The Visual System . . . . . 179
The Visual Pathway. . . . 179
The Accessory Optic System 184
Visual Reflexes. . . . 185
Eye Movements . . . 185
Ascending Reticular Systems 197
The Reticular Formation 197
The Raphe Nuclei 197
The Medial Reticular Formation 202
The Lateral Reticular Formation 202
Noradrenergic Cell Groups. . . 204
Adrenergic Cell Groups . . . . 205
Cholinergic Cell Groups; Locomotor and Relay Centres 205
Ascending Reticular Pathways . . . . . . . . . . . . 206
Ascending Projections from the Raphe Nuclei. . . . . 206
Ascending Projections from the Medial Reticular Zone . 208
Ascending Projections from the Lateral Reticular Zone . 211
Cerebellum . . . . . . . 221
Introduction. . . . . . 221
Cortex and Central Nuclei 222
Types of Afferent and Intrinsic Connections 222
Terminations of Mossy Fibre Systems . . . 224
Functional Zones: Organization of the Olivocerebellar Climbing Fibre
System and the Corticonuclear Projection 228
The Vestibulocerebellum. . . . . . . . . 232
Efferent Pathways . . . . . . . . . . . 232
Function and Malfunction of the Cerebellum 236
Thalamocortical and Corticothalamic Connections 237
The Thalamus: Its Parts and Peduncles . . . . 237
Specific and Non-specific Nuclei of the Thalamus 238
Thalamic and Thalamocortical Circuits . . 239
Connections of the Ventral Thalamic Nuclei . . 239
Contents XI
Connections of the Anterior Nucleus and the Lateral Dorsal Nucleus

of the Thalamus . . . . . . . . . . . . . . . . 240
Connections of the Medial Nucleus of the Thalamus . 240
Connections of the Medial Geniculate Body . . . . 241
Connections of the Posterior Nuclei of the Thalamus. 241
Connections of the Lateral Geniculate Body 241
Connections of the Lateral Thalamic Nuclei 244
Connections of the Intralaminar Nuclei 246
Motor Systems. . . . . . . 247
Long Corticofugal Pathways 247
Introduction. . . . . . 247
Medial and Lateral Motor Systems 253
Reticulospinal, Raphespinal and Coeruleospinal Systems 256
Direct and Indirect Cortico-motoneuronal Connections 257
The Pyramidal Tract Syndrome. . . . . . 258
The So-Called Extrapyramidal Motor System. 258
Introduction. . . 258
Structural Features . . . . . . . . . . . 259
Fibre Connections . . . . . . . . . . . 260
Aspects of the Organisation of the Striatum. 267
Subdivision of the Basal Ganglia into Dorsal and Ventral Sectors 269
Fibre Connections of the Ventral Striatum . . . . . . 270
The Magnocellular Basomedial Telencephalic Complex. 273
Descending Reticular Systems . . . . . . . . . . . . 279
Descending Projections from the Raphe Nuclei . . . . 279
Descending Projections from the Medial Reticular Zone 282
Descending Projections from the Lateral Reticular Zone 284
Olfactory and Limbic Systems . . . . . . . . . . . 293

Introduction. . . . . . . . . . . . . . . . . . 293
The Central Limbic Continuum: Sections and Centres 296
The Central Limbic Continuum: Circuitry and Major Conduction
Channels . . . . . . . . . . . . . . . . . . . . . . . . . 302
The Central Limbic Continuum: Functional Connections of Individual
Centres . . . . 309
The Amygdala. . . . . . . . . . . . 323
Introduction. . . . . . . . . . . . 323
Fibre Systems Related to the Amygdala 324
Afferents to the Amygdala. . . . . . 324
Efferents from the Amygdala. . . . . 328
Comments on the Connections and Functions of the Amygdala 329
Bed Nucleus of the Stria Terminalis 332
The Hippocampus . . . . . . . . . . . . 334
Introduction. . . . . . . . . . . . . . 334
Fibre Systems Related to the Hippocampus. 335
Intrinsic Connections of the Hippocampus 335
Afferents to the Hippocampus . 336
Efferents from the Hippocampus . . . . 340
XII Contents
The Circuit of Papez . . . . . . . . . . . . . . . 341

The Olfactory System . . . . . . . . . . . . . . . . 344
The Limbic System at Large: A Functional Commentary . 348
Long Association and Commissural Connections 365
General Features. . . . . 365
The Somatosensory Cortex. 370
The Visual Cortex . 371
The Auditory Cortex 372
The Frontal Lobe. . 373
The Paralimbic Areas 375
References. . 377
SUbject Index 419

Introduction
Purpose and Plan between the basic neuroanatomy required by

medical students and the specialised litera-
This atlas has been designed with the object ture in the field comprising different sub-
of providing a comprehensive pictorial sur- disciplines. It may therefore be useful at
vey of the macroscopic and microscopic many levels of medical education and can
structure of the human central nervous sys- also serve as a quick pictorial review for prac-
tem and its surrounding structures. titioners in the various neurological sciences.
The pictorial material encompasses half-
tone illustrations and line drawings, all de-
rived from original macroscopic and micro-
Material, Techniques, and Preparation
scopic preparations. Considerable thought
has been spent during the preparation of
of the Illustrations
these drawings to achieve an optimal combi-
nation of clarity and exactness. Moreover, The gross anatomical part of this atlas is
great pains have been taken to achieve a max- based on eight brains and one spinal cord
imal coherence of thematically related fig- of adult individuals with no record of neu-
ures. The illustrations are arranged in five rological diseases. These specimens were
parts. The first part depicts the gross appear- fixed for at least two months in formalin.
ance and three-dimensional structure of the One specimen was used for the illustrations
brain and spinal cord. The second part shows showing the external morphology. This brain
three-dimensional illustrations of the adnexa was then serially sliced into 2-mm-thick sec-
of the central nervous system, i.e. the vessels, tions in the coronal plane. Three other brains
meninges and cisterns. The third part in- were sliced in the three other conventional
cludes drawings of a number of whole brain planes: sagittal, horizontal and perpendicu-
slices, sectioned in four different directions. lar to the axis of the brain stem. The serial
The fourth part consists of a carefully slices thus obtained were employed in the
selected series of 25 microscopic sections preparation of various graphical and three-
through the spinal cord, brain stem and basal dimensional reconstructions. The latter were
prosencephalon. In these sections the fibre made from 4-mm-thick styrofoam sheets.
systems are shown on the right and the cell The remaining four brains were dissected in
masses on the left. The final part integrates a number of ways. All of the figures in this
and amplifies the data presented in the pre- section showing features of the internal con-
ceding sections. The central nervous system figuration of the brain are based on recon-
is divided here into ten functional systems, structions or dissections or both. Therefore,
and the interrelationships of the centres and in these figures the size, shape and spatial
fibre paths belonging to each of these systems relationships of the structures revealed are
are depicted and briefly described. Emphasis reproduced with great exactitude.
has been laid on aspects of clinical signifi-
cance. This atlas is intended to fill the gap
2 Introduction
The part of the book on vessels and meninges is the same as in the half-tone Figs. 43-45,
presented a specific difficulty. Due to the for which the specimen was basic; the speci-
high degree of variability within the normal men was also used in composing the half-
scope of vascular patterns, it was not easy tone Fig. 41.
to find "standard" specimens to serve as the
basis for the illustrations. Therefore, apart The cisterns were studied by using a sliced
from depicting our own material, we made head. A complete head of an aged person
two compromises. Firstly, if a vascular detail was frozen-sectioned at intervals of 4 mm,
was apparently uncommon, we replaced it by leaving slices of 3-mm thickness, the remain-
the most common configuration. Secondly, ing 1 mm being lost due to the thickness of
we added figures, mostly taking the form of the band saw. The direction of sectioning was
diagrams and based on different sources, to coronal, perpendicular to the horizontal
illustrate the more general configuration. To plane of Frankfurt. Both the frontal and the
know what is common or general in this occipital sides of each slice, were drawn in
respect we studied not only the anatomical detail. These drawings then served as the ba-
literature on the subject matter, but also - sis for two types of reconstruction: a spatial
because of the clinical relevance - some reconstruction in foam cardboard (Depafit),
radiological publications. and a number of graphic reconstructions.
For the preparation of the illustrations in
this section four specimens were studied. Two Of the literature consulted during the prepa-
brains were used to study the arteries; in the ration of our illustrations on vessels and
third the sinuses and veins were the special meninges, several anatomical and radiologi-
objects of investigation; and the fourth was cal publications should be mentioned. Sys-
employed for reconstruction purposes, which tematic descriptions of the cisterns and of the
helped in studying the topography of the vascular patterns, their variations and their
brain's surrounding and internal structures. occurrence can be found in the works of
Lang [756] and Salamon and Huang [1183].
The arteries of two formalin-fixed specimens Waddington [1465] presents many variations
were filled and coloured by injecting coloured in vascular patterns together with abnormal
latex into both internal carotids and into one studies. General or typical patterns can be
or both of the vertebral arteries. The speci- found in Lang [756] and Salamon and Huang
mens were then dissected carefully; at each [1183] and in the publications by
step of dissection both photographs and Lazorthes [769] and Szikla et al. [1348].
drawings were made, mostly in standard For special items we also referred to other
views. After having completed the documen- publications. The central arteries are well
tation of the four halves, one half was se- illustrated in the book by Schlesinger [1216].
lected for having the most common configu- Vascular details as they may appear in radio-
ration. This half served as the base for the graphs are amply illustrated in Salamon's
half-tone Figs. 31-35,40 and 46. atlas [1182], while topographical details can
be found in Seeger's comprehensive atlas
The sinuses and veins were filled and coloured [1226] and in the publication by Hovelacque
by injecting coloured latex into both internal [559]. Thijssen's thesis [1371] served us as a
jugular veins of a fresh, i.e. non-fixed, com- guide for obtaining a better understanding
plete head of a young adult. The head was of the spatial relationships of vessels, brain
then submerged in formalin for several weeks and skull and of their consequences for (ra-
and after that dissected step by step. The dif- diologic) projection.
ferent steps were documented by photo- The configuration of the cisterns has been
graphs, mostly in standard views, and by studied by Lilieqvist [789] and Lang [755] by
drawings in a fixed oblique view. This view using corrosion preparations. Duvernoy pro-
Introduction 3
duced a standard work on the brainstem ves- apparatus. These samples were employed for
sels [315], while the spinal vascularisation depicting the cell masses in the left halves
and topography is well presented in the pub- of the sections. Thus, it should be appreciat-
lications of Rickenbacher [1126], Djindjian ed that the cells are represented at a magnifi-
[307], Theron and Moret [1370] and Renard cation ten times that of the section as a
et al. [1116]. whole.
The sections through the brain stem and
The brain slices that constitute the third part the spinal cord are based on Nissl and
of this atlas were selected from the four series Haggqvist material. The latter technique was
of slices already mentioned. The brains were selected for the analysis of the white matter,
embedded in gelatin and sliced on a rotary- because it shows both the axon and the mye-
blade commercial meat slicer. Due to the lin sheath of the individual nerve fibres in
elasticity of the large gelatin blocks we expe- contrasting colours. Neuronal somata can
rienced initially considerable difficulties in also be observed in Haggqvist material,
maintaining the intended plane of sectioning though with less distinctness than in Nissl
throughout a series. However, this problem sections. The procedure followed in the prep-
was ingeniously solved by Mr. A. Binnendijk, aration of the drawings involved the follow-
our laboratory assistant, by embedding the ing sequence of steps:
brains in boxes, prepared from styrofoam
1. Haggqvist sections of the levels to be
plates, and by subsequently slicing the brains
depicted were selected.
while still encased in the surrounding box.
2. With the aid of a photographic enlarger,
This procedure yielded perfect, well-oriented,
negative photographic prints were made of
continuous series of slices.
these sections at a magnification of seven
diameters.
The microscopic sections that comprise the
3. Under microscopic control the various
fourth part were all drawn from original
areas of grey and white matter were delineat-
preparations. The sections through the basal
ed on the photographs. In this way outline
prosencephalon are based on one of the ex-
drawings for the final figures were obtained.
cellent Weiger-Pal series which the late Pro-
4. The fibre composition and the fibre pat-
fessor G. Jelgersma employed for his Atlas
tern of the various tracts and more diffuse
Anatomicum Cerebri Humani [618] and on a
areas of white matter were analysed; the
series of KlUver-Barrera preparations. The
fibres were graded into three groups: coarse,
procedure was as follows. Seven sections
medium or thin. By using dots and lines of
were selected from the Weigert-Pal series.
corresponding diameters the results of the
From these sections outline drawings were
analysis were represented semi-diagrammati-
prepared at a magnification of four diameters
cally in the drawings.
in which the position of the fibre systems and
5. From corresponding Nissl preparations
cell masses was indicated. Since the series
samples of the various nuclei and other cellu-
only includes one half of the brain, the other
lar areas were drawn at a magnification of
half was added in the drawings as a mirror
70 diameters. These samples were employed
image. In order to obtain bilateral symmetry
for depicting the cell masses in the left halves
slight corrections appeared to be necessary.
of the drawings. Thus, in this series of draw-
In the right half of the drawings the fibre
ings, as in those of the sections through the
pattern was drawn in semi-diagrammatically.
forebrain, the neuronal somata are repre-
The cell masses were studied in correspond-
sented at a magnification ten times that of
ing sections of the Kliiver-Barrera series.
the sections as a whole.
From each griseum one or several character-
istic samples were drawn at a magnification The pictures constituting the fifth and final
of 40 diameters with the aid of a projection part fall into two categories: (1) plates show-
4 Introduction
ing the topographic relationships of the of these structures various works have been
structures belonging to the various functional consulted, among which the atlases of Riley
systems and (2) diagrams illustrating the neu- [1127], Schaltenbrand and Bailey [1209], and
ronal interrelationships within these systems. Singer and Yakovlev [1256] should be
The plates are largely based on reconstruc- especially mentioned. Our principal sources
tions prepared from our own macroscopic for interpretation and terminology of struc-
and microscopic material. The diagrams are tures in particular brain regions were as fol-
based on data compiled from the literature. lows: cell masses of the brain stem: Olszews-
ki and Baxter [1029]; cerebellum: Angevine,
et al. [46]; thalamus: Dewulf [298], Van
Buren and Borke [1407] and Jones [629];
Annotations
hypothalamus: Nauta and Haymaker [986];
allocortical and adjacent structures: Stephan
An effort has been made to enhance the use- [1291]; amygdala: Crosby and Humphrey
fulness of this atlas by including brief de- [265]. It was not feasible to label all of the
scriptions of the functional systems depicted. recognisable structures on every plate. In the
These annotations include the most impor- series of slices (Part III) and microscopic sec-
tant findings of modern experimental neuro- tions (Part IV), structures that appear repeat-
anatomical research. edly have been labelled on alternative plates
or, in a number of instances, even less
frequently.
Terminology and Labelling
Since the Latin terminology has the obvious

advantage of accepted international usage, it
was decided to employ this terminology
wherever possible. However, to facilitate use
of the atlas we have included the English or
Anglicized equivalents in the index wherever
they seemed important. As regards gross
anatomy, the Paris Nomina Anatomica,
which were adopted in 1955 by the Interna-
tional Nomenclature Committee and revised
in 1960, 1965 and 1975, has been used ac-
cording to the last edition. in this fourth edi-
tion the list of cerebral arteries was amplified
at the insistence of radiologists and others
(page A64). Nevertheless the list is still in-
complete, as has been stated by Lang [756]
and others. We filled gaps by Latinizing the
Anglicized terms in relevant radiological
works (Salamon and Huang [1183], and
Duvernoy [315]) or by adapting Latin terms
from anatomical works (Lang [756]) to the
already existing terms in the NA. Unfortu-
nately an internationally approved nomen-
clature for the microscopic structures of the
brain does not exist. For the nomenclature
Part I Gross Anatomy
Orientation
External and Medial Views
Internal Structures
6 Gross Anatomy
1 Cerebrum
2 Cerebellum
3 Truncus cerebri
4 Medulla spinalis
Fig. 1. The central nervous system in situ (1 /6 x)

Orientation 7
Q
1 Telencephalon 0
(Cerebrum) :; ~
..<:i
0- "
:;
....
2 Telencephalon impar "Q
u
"E
'""....0
0)
u
3 Diencephalon ~
a
;:l
Q
0 '"0
:; >....
4 Mesencephalon ..<:i 0)
0- Q
8Q to
}
"C
5 Pons Q .!> S
Meten- 0
....
0) ~
~>-.
6 Cere- :;
cephalon ..c: 0)
u
bellum 0- ~
"uQ ~
u
Q
'"
"R
7 Myelencephalon "a
.!> ;:l
E!::
to
....
;:l
0)
(Medulla 0
..<:i Z
oblongata) P:::
8 Medulla spinalis
Fig. 2. Medial surface of the right half of the brain in the bisected head. The position of its major
subdivisions is indicated (2/ 5 x)
8 Gross Anatomy
I Ventriculus lateralis
2 Foramen interventriculare
3 Ventriculus tertius
4 Aqueductus cerebri
5 Ventriculus quartus
6 Canalis centralis
Fig. 3. The ventricular system of the brain. The arrow passes through the foramen interventriculare
from the third ventricle to the lateral ventricle (3/5 x )
Orientation 9
A B
1 Polus occipitalis 5 Operculum fronto-parietale 9 Lobus frontalis

2 Lobus occipitalis 6 Operculum temporale 10 Polus frontalis
3 Lobus parietalis 7 Operculum frontale 11 Lobus limbicus
4 Lobus temporalis 8 Polus temporalis
10
c 4
Fig. 4 A-D. Subdivision of the right cerebral hemisphere into lobes. A lateral view; B medial view;
C superior view; D interior view (1/2 x)
10 Gross Anatomy
Fig. SA, B. Subdivision of the cortex of the right cerebral hemisphere into cytoarchitectonic fields
according to Brodmann (3/4 x ). A lateral view ; B medial view
External and Medial Views 11
I Fissura longitudinalis cerebri

2 Gyrus frontalis superior
3 Sulcus frontalis superior
4 Gyrus frontalis medius
5 Sulcus frontalis inferior
6 Gyrus frontalis inferior
7 Sulcus precentralis
8 Gyrus precentralis
9 Sulcus centralis
10 Lobulus paracentralis
II Gyrus postcentralis
12 Sulcus postcentralis
13 Sulcus intraparietalis
14 Lobulus parietal is superior
15 Lobulus parietalis inferior
16 Sulcus parielo-occipitalis
17 Gyri occipitales
18 Sulci occipitales
Fig. 6. The brain seen from above (1/1 x )

12 Gross Anatomy
18 Sulcus precentralis
19 Sulcus frontalis superior
20 Gyrus frontalis superior
21 Gyrus frontalis medius
22 Sulcus frontalis inferior
23 Pars opercularis }
24 Pars triangularis Gyrus frontalis inferior
1 Sulcus ceotralis 25 Pars orbitalis
2 Gyrus postcentralis 26 Sulcus lateralis, ramus ascendens
3 Sulcus postcentralis 27 Sulcus lateralis, ramus anterior
4 Lobulus parietalis superior 28 Sulci orbitales
5 Sulcus parieto-occipitalis 29 Gyri orbitales
6 Lobulus parietalis inferior 30 Bulbus olfactorius
7 Sulcus iotraparietalis 31 Tractus olfactorius
8 Gyrus angularis 32 Sulcus lateralis
9 Gyrus supramarginaJis 33 Gyrus temporalis superior
10 Sulcus lateralis, ramus posterior 34 Sulcus temporalis superior
11 Gyri occipitales 35 Gyrus temporalis medius
12 Sulcus luoatus 36 Sulcus temporalis inferior
13 Sulcus occipitalis anterior 37 Gyrus temporalis inferior
14 Sulci occipitales 38 Pons
15 lncisura preoccipitalis 39 Flocculus
16 Hemispherium cerebeUi 40 Medulla oblongata
Fig. 7. Lateral view of the brain (1 /1 x )

1 Sulcus centralis
2 Sulcus lateralis, ramus posterior
3 Sulcus lateralis, ramus ascendens
4 Operculum fronto-parietale
5 Sulcus circularis insulae
6 Sulcus lateralis, ramus anterior
7 Gyrus longus insulae
8 Sulcus centralis insulae
9 Gyri breves insulae
10 Operculum frontale
11 Operculum temporale
12 Limen insulae
13 Polus insulae
Fig. 8. Dissection of the right cerebral hemisphere to display the insula (1 /1 x)

14 Gross Anatomy
8 Bulbus olfactorius 18 Gyrus longus insulae

9 Tractus olfactorius 19 Pedunculus cerebri
10 Polus insulae 20 Pons
1 Fissura longitudinalis cerebri 11 Stria olfactoria medialis 21 Pyramis
2 Chiasma opticum 12 Trigonum olfactorium 22 Oliva
3 Tractus opticus 13 Substantia perforata anterior 23 Flocculus
4 Infundibulum 14 Stria olfactoria lateralis 24 Plexus choroideus ventriculi
5 Tuber cinereum 15 Gyrus diagonalis quarti
6 Corpus mamillare 16 Limen insulae 25 Hemispherium cere belli
7 Fossa interpeduncularis 17 Gyri breves insulae 26 Vermis cere belli
Fig. 9. Basal view of the brain. The frontal portion of the left temporal lobe has been removed to
expose the underlying structures (1 /1 x)
12 Gyrus rectus
13 Sulcus olfactorius
14 Area subcallosa
1 Sulci orbitales 15 Gyrus paraterminalis
2 Gyri orbitales 16 Gyrus diagonalis
3 Gyrus temporalis superior 17 Sulcus rhinalis
4 Sulcus temporalis superior 18 Gyrus ambiens
5 Sulcus temporalis inferior 19 Sulcus collateralis
6 Gyrus temporalis medius 20 Gyrus parahippocampalis
7 Sulcus occipitotemporalis 21 Pulvinar thalami
8 Gyrus temporalis inferior 22 Splenium corporis callosi
9 Gyrus occipitotemporalis 23 Isthmus gyri cinguli
lateralis 24 Sulcus calcarinus
10 Incisura preoccipitalis 25 Gyrus occipitotemporalis
11 Gyri occipitales medialis
Fig. 10. Basal view of the right cerebral hemisphere. The olfactory tract has been sectioned (1 /1 x)
16 Gross Anatomy
19 Sulcus centralis
1 Gyrus frontalis superior 20 Lobulus paracentralis
2 Sulcus cinguli 21 Sulcus cinguli, pars marginalis
3 Gyrus cinguli 22 Lobulus parietalis superior
4 Sulcus corporis callosi 23 Precuneus
5 Gyrus paraterminalis 24 Sulcus subparietalis
6 Sulcus parolfactorius posterior 25 Indusium griscum
7 Area subcallosa 26 Sulcus parieto-occipitalis
8 Sulcus parolfactorius anterior 27 Cuneus
9 Gyrus rectus 28 Sulcus calcarinus
10 Gyrus intralimbiCUS} 29 Gyrus fasciolaris
11 Limbus Giacomini Uncus 30 Taenia thalami
12 Gyrus uncinatus 31 Isthmus gyri cinguli
13 Gyrus semilunaris 32 Gyrus dentatus
14 Gyrus am biens 33 Gyrus occipitotemporalis medialis
15 Incisura unci 34 Sulcus collateralis
16 Gyrus parahippocampalis 35 Gyrus occipitotemporalis lateralis
17 Sulcus rhinalis 36 Sulcus occipitotemporalis
18 Gyrus temporalis superior 37 Gyrus temporalis inferior
Fig. 11. Medial aspect of the right cerebral hemisphere (1 /1 x)

1 Truncus corporis callosi

2 Septum pellucidum
3 Fornix 17 Splenium corporis callosi
4 Genu corporis callosi 18 Thalamus
5 Rostrum corporis callosi 19 Tela choroidea ventriculi tertii
6 Foramen interventriculare 20 Corpus pineale
7 Commissura anterior 21 Lamina quadrigemina
8 Lamina terminalis 22 Aqueductus cerebri
9 Hypothalamus 23 Velum medullare superius
10 Chiasma opticum 24 Ventriculus quartus
11 Nervus opticus 25 Velum medullare inferius
12 Bulbus olfactorius 26 Vermis cere belli
13 Nervus oculomotorius 27 Hemispherium cere belli
14 Pons 28 Tela choroidea ventriculi quarti
15 Medulla oblongata 29 Apertura mediana ventriculi quarti
16 Medulla spinalis 30 Canalis centralis
Fig. 12. Medial view of the right half of the bisected brain (1 /1 x)
18 Gross Anatomy
1 Corpus fornicis
2 Crus fomicis 24 Lingula cerebelli
3 Recessus suprapinealis 25 Fissura precentralis
4 Habenula 26 Lobulus centralis
5 Commissura habenularum 27 Fissura preculminata
6 Corpus pineale 28 Culmen
7 Recessus pinealis 29 Fissura prima
8 Commissura posterior 30 Declive
9 Plexus cboroideus ventriculi tertii 31 Fissura superior posterior
10 Adbesio intertbalamica 32 Folium vermis
11 Co=issura anterior 33 Fissura horizontalis
12 Columna fomicis 34 1\iber vermis
13 Sulcus bypothaJamicus 35 Fissura prepyramidalis
14 Lanrinaterallnalis 36 Pyramis vermis
15 Corpus mamillare 37 Fissura secunda
16 Fossa interpeduncularis 38 Uvula vermis
17 Recessus opticus 39 Fissura posterolateralis
18 Chiasma opticum 40 Nodulus
19 Recessus infundibuli 41 Velum medullare inferius
20 Infundibulum 42 Fastigium
21 Colliculus superior 43 Lobulus semilunaris inferior
22 Colliculus inferior 44 Lobulus biventer
23 Velum medulJare superius 45 Tonsilla cerebelli
Fig. 13. Medial view of the bisected brain stem and cerebellum (3/2 x)
--------/
19 Thberculum anterius thalami

20 Tractus opticus
21 Pedunculus cerebri
1 Thalamus 22 Nervus opticus
2 Pulvinar thalami 23 Nervus oculomotorius
3 Colliculus superior 24 Nervus trigeminus, radix motoria
4 Corpus geniculatum laterale 25 Nervus trigeminus, radix sensoria
5 Colliculus inferior 26 Pons
6 Trigonum lemnisci 27 Nervus facialis
7 Nervus trochlearis 28 Nervus intermedius
8 Pedunculus cerebellaris superior 29 Nervus vestibulocochlearis
9 Pedunculus cerebellaris medius 30 Nervus abducens
10 Pedunculus cerebellaris inferior 31 Nervus glossopharyngeus
11 Recessus lateralis ventriculi quarti 32 Pyramis
12 Plexus choroideus ventriculi q uarti 33 Nervus vagus
13 Tela choroidea ventriculi quarti 34 Nervus accessorius
14 Thberculum nuclei cuneati 35 Nervus hypoglossus
15 Obex 36 Oliva
16 Thberculum nuclei gracilis 37 Radices craniaJes nervi accessorii
17 Sulcus lateralis posterior 38 Radices spinales nervi accessorii
18 Radix dorsalis nervi spinalis 39 Radices ventrales nervi spinalis
Fig. 14. Lateral view of the brain stem and the diencephalon after removal of the structures surrounding
the thalamus (3/2 x )
20 Gross Anatomy
1 Ventriculus lateralis 25 Taenia choroidea

2 Ventriculus tertius 26 Lamina affixa
3 Corpus pineale 27 Stria terminalis
4 Brachium coUiculi superioris 28 Stria medullaris thalami
5 Colliculus superior 29 Taenia thalami
6 Brachium colliculi inferioris 30 Trigonum habenulae
7 Colliculus inferior 31 Pulvinar thalami
8 Pedunculus cerebri 32 Corpus geniculatum media Ie
33 Corpus genicu la tum laterale
9 Velum medu.llare superius

10 Eminentia medialis
11 Sulcus medianus (ventriculi quarti) 34 Nervus trochlearis
12 CoUiculus facialis 35 Lingula cerebelli
13 Area vestibularis 36 Nervus trigeminus
14 Trigonum nervi hypoglossi 37 Pedunculus cerebellaris superior
.1 S Trigonum nervi vagi 38 Pedunculus cerebellaris medius
.16 Taenia ventriculi quarti 39 Pedunculus cerebellaris inferior
.11 Tuberculum nuclei cuneati 40 Recessus lateralis ventriculi quarti
18 Tuberculum nuclei gracilis 41 Apertura lateralis ventriculi quarti
19 Funiculus lateralis 42 Plexus choroideus ventriculi quarti
20 Sulcus lateralis posterior 43 Tela choroidea ventriculi quarti
21 Fasciculus cuneatus 44 Nervus accessorius
22 Sulcus intermedius posterior 4S Apertura mediana ventriculi quarti
23 Fasciculus gracilis 46 Obex
24 Sulcus medianu posterior 47 Radix dorsalis nervi spinalis
Fig. 15. Dorsal view of the brain stem and the diencephalon after removal of the structures surrounding
the thalamus. The contour of the cerebellum is indicated (3/2 x)
16
1 Culmen 7 Lobulus quadrangularis 13 Lobulus semilunaris inferior

2 Declive 8 Fissura prima 14 Fissura ansoparamediana
3 Folium vermis 9 Lobulus simplex 15 Lobulus gracilis
4 Tuber vermis 10 Fissura superior posterior 16 Fissura prebiventeris
5 Pyramis vermis 11 Lobulus semilunaris superior 17 Lobulus biventer
6 Uvula vermis 12 Fissura horizontalis 18 Fissura secunda
19 Tonsilla cerebelli
., hemispherium cerebelli
E
.,> pars
inler-
I
med ia
fissura pr ima
"
fissura secunda
Fig. 16. Dorsal view of the cerebellum (6/5 x; diagram: 1/1 x)

22 Gross Anatomy
1 Taenia choroidea 24 Ventriculus lateralis

2 Lamina aiflXa 25 Ventriculus tertius
3 Taenia thalami 26 Infundibulum
4 Tuberculum anterius thalami 27 Corpus mamillare
5 Thalamus 28 Pedunculus cerebri
6 Adhesio interthalamica 29 Substantia perforata posterior
7 Chiasma opticum 30 Fossa interpeduncularis
8 Nervus opticus 31 Pons
9 Tractus optic us 32 Sulcu basilaris pontis
10 Corpus geniculatum laterale
11 Nervus oculomotorius
12 Nervus trochlearis
,/
,/
(
J
13 Nervus trigeminus, radix motoria 33 Pedunculus cerebeUaris medius
14 Nervus trigeminus, radix sensoria 34 Plexus choroideus ventriculi
15 Nervus abducens quarti
16 Nervus facialis 35 Oliva
17 Nervus intermedius 36 Pyramis
18 Nervus vestibulocochlearis 37 Decussatio pyramidum
19 Nervus glossopharyngeus 38 Funiculus lateralis
20 Nervus vagus 39 Sulcus lateralis anterior
21 Nervus accessorius 40 Funiculus anterior
22 Nervus hypoglossus 41 Fissura mediana anterior
23 Radices ventrales nervi spinalis
Fig. 17. Ventral view of the brain stem and the diencephalon. The structures surrounding the thalamus
have been removed (3/2 x)
24
1 Velum medullare superius 12 Culmen 24 Fissura horizontalis

2 Pedunculus cerebella ris superior 13 Fissura preculminata 25 Lobulus semilunaris inferior
3 Fastigium 14 Lobulus quadrangularis 26 Fissura ansoparamediana
4 Velum medullare inferius 15 Lobulus centralis 27 Lobulus gracilis
5 Pedunculus cerebellaris inferior 16 Ala lobuli centralis 28 Fissura prebiventeris
6 Pedunculus cerebellaris medius 17 Lingula cerebelli 29 Lobulus biventer
7 Nervus intermedius 18 Fissura prima 30 Fissura intrabiventeris
8 Nervus vestibulocochlearis 19 Lobulus simplex 31 Fissura secunda
9 Recessus lateralis ventriculi quarti 20 Fissura superior posterior 32 Tonsilla cerebelli
10 Tela choroidea ventriculi quarti 21 Lobulus semilunaris superior 33 Nodulus
11 Plexus choroideus ventriculi quarti 22 Pedunculus flocculi 34 Fissura posterolateralis
23 Flocculus 35 Uvula vermis
hemis pher i um cere b elli
~
<II
pars
inler-
I
> me d ia
, issura secunda
Fig. 18. Ventral view of the cerebellum (6/5 x; diagram : 1/1 x)

Th12
Th1 0
L 1
C1
C2 L2
C3 L3
Th 11
L4
C4 L5
S 1
C5
S 2 Th12
C6
C7
C8 L1
Th 1
8
Th 2
L2
Th 3 Cauda equina
Th 4
Th 5
Th 6
-L 5
- - -Th5
Th 7 - - Fila radicularia
Th 8 S1
~--s2
Th 9
''---- S 3
Thl0
""--- S 4
Th 11
~--- S5
"---- - -Co
Fig. 19. Dorsal view of the spinal cord showing attached dorsal root filaments and spinal ganglia.
The Cervical (C), Thoracic (Th), Lumbar (L), Sacral (S) and Coccygeal (Co) spinal nerves have been
transected at their site of exit from the intervertebral foramina . The position of the spinal segments
is indicated left to the cord (2/3 x )
Medulla oblongata
Medul la spinalis,
Pars lu mbalis
_ Intumescentia lumballs
Medul la spinalis,
Pars cervicalis
_ Intumescenlia cervlcalis
- - - - Conus medullaris
Fissura mediana anterior
-~ Funiculus anterior
- _. Fila radicularla
- - . Funiculus lateralls
- - - Fi lum terminale
Medulla spina li s,
Pars thoracalis
Fig. 20. Ventral view of the spinal cord; the ventral root filaments have been transected (2/3 x)
26 Gross Anatomy
16
1 Ventriculus lateralis, pars centralis 9 Recessus suprapinealis

2 Ventriculus lateralis, cornu anterius 10 Recessus pinealis
3 Adhesio interthalamica 11 Ventriculus lateralis, cornu posterius
4 Foramen interventriculare 12 Aqueductus cerebri
5 Ventriculus tertius 13 Fastigium
6 Recessus opticus 14 Ventriculus quartus
7 Recessus infundibuli 15 Recessus lateralis ventriculi quarti
8 Ventriculus lateralis, cornu inferius 16 Canalis centralis
Fig. 21. The ventricles of the brain; oblique view from behind and above (6/5 x)
Internal Structures 27
1 Lamina affixa 9 Corpus fornicis

2 Stria terminalis 10 Commi ura fornicis
3 Taenia thalami 11 Corpus pineale
4 Taenia choroidea 12 Crus fornici
5 Taenia fornicis 13 Taenia fimbriae
6 Thalamus 14 Fimbria hippocampi
7 Columna fornicis
8 Corpus mamillare
...........-:....:.:.:...:....... .
Fig. 22. Topography of diencephalic and telencephalic taeniae; oblique view from behind and above.
In the complementary diagram the choroid walls of the lateral and third ventricles are shown. Dense
stippling: ventricular surface; light stippling: meningeal surface; double arrow: interventricular foramen
(2/1 x ; diagram: 5/3 x)
28 Gross Anatomy
1 Stria longitudinalis medialis 14 Gyrus cinguli

2 Stria longitudinalis lateralis 15 Sulcus corporis caUosi
3 Corpus callosum 16 Stria longitudinalis medialiS} Hippocampus
4 Fornix 17 Indusium griseum supra-
5 Taenia fornicis 18 Stria longitudinalis lateralis commissuralis
6 Gyrus fasciolaris 19 Gyrus fasciolaris
7 Gyri Andreae Retzii 20 Fasciola cinerea
8 Hippocampus precommissuralis 21 Gyri Andreae Retzii
9 Gyrus dentatus 22 Cornu ammOniS} HO
10 Alveus hippocampi 23 Subiculum Ippocampu °
11 Fimbria hippocampi 24 Gyrus dentatus retrocommJ urahs

12 Cornu ammonis 25 Sulcus hippocampi
13 Digitationes hippocampi 26 Sulcus fimbriodentatus
27 Fimbria hippocampi
28 Gyrus inlralimbicus
29 Limbus Giacomini
30 Sulcus hippocampi
31 Gyrus uncinatu
Fig. 23. Dissection showing the hippocampus and some related structures in oblique view from behind
and above (2/1 x)
9
10
12
13
9 Nucleus ventralis lateralis

1 Nucleus anterior thalami 10 Nucleus anterior thalami
2 Nucleus ventralis lateralis 11 Nucleus medialis thalami
3 Nucleus lateralis dorsalis 12 Nucleus lateralis dorsalis
4 Nucleus lateralis posterior 13 Nucleus lateralis posterior
5 Nucleus ventralis anterior 14 Nuclei habenulae
6 Nucleus ventralis posterolateral is 15 Nuclei pulvinares
7 Nuclei pulvinares 16 Corpus genicula tum laterale
8 Corpus geniculaturn latera Ie 17 Corpus geniculatum mediale
Fig. 24. A model of both thalami; oblique view from behind and above. The nucleus reticularis and
the so-called midline nuclei have been omitted (2/1 x)
I Nucleus anterior thalami

2 Nucleu ventralis lateralis
3 Nucleus medialis thalami
4 Fasciculus mamillothalamicus
5 Nucleus ventralis anterior
6 Nucleus lateralis dorsalis
8 Lamina medullaris interna
9 Nucleus ventralis la teralis
10 Nucleus centromedianus 14 Nucleus lateral.is posterior
11 Nucleus parafascicuIaris 15 Nuclei pulvinares
12 Nucleus ventralis posterolateral is 16 Corpus geniculatum mediale
13 Nucleus ventralis posteromedialis 17 Corpus geniculatum latera Ie
Fig. 25. Left half of the same model as shown in Figure 24. The position of its major nuclei is indicated
on three frontal sections (2/1 x)
30 Gross Anatomy
1 Nucleus caudatus }
2 Nucleus lentiformi Corpus striatum
3 Thalamus sinister
4 Thalamus dexter
5 Stria medullaris thalami
Fig. 26. The corpus striatum and the thalamus of both sides in oblique view from behind and above
(6/5 x)
1 Corona radiata (some isolated fibres)

2 Cauda nuclei caudati
3 Pontes grisei caudatolenticulares
4 Corpus nuclei caudati
5 Putamen
6 Outline of thalamus
7 Capsula interna
(one isolated fibre indicated)
8 Globus pallid us, pars lateralis
9 Globus pallid us, pars medialis
10 Commissura anterior
11 Caput nuclei caudati
12 Nucleus accumbens
13 Pedunculus nuclei lentifonnis
14 Junction of cauda to 13
Fig. 27. The basal ganglia in medial view (6/5 x)

8 Capsula interna, pars sublentifonnis

9 Capsula interna, pars retrolentifonnis
10 Capsula interna, crus posterius
12 Mesencephalon
1 Capsula interna, crus anterius

2 Ventriculus lateralis, par centralis
3 Ventriculus lateralis, cornu anterius 13 Thalamus
4 Caput nuclei caudati 14 Pontes gri ei caudatolenticulares
5 Corpus nuclei caudati 15 Ventriculus lateralis, cornu posterius
6 Globus pallid us 16 Cauda nuclei caudati
7 Putamen 17 Ventriculus latera lis, cornu inferius
Fig. 28. The corpus striatum of both sides, viewed from above. The thalamus and the extent of the
lateral ventricle can be seen on the left side. The internal capsule and its convergence upon the peduncu-
lus cerebri are indicated on the right side (6/5 x)
7 Ventriculus lateralis, pars centralis

1 Corpus nuclei caudati '" 8 Pontes grisei caudatoienticulares
2 Ventriculus lateralis, cornu anterius 9 Cauda nuclei caudati
3 Caput nuclei caudati 10 Thalamus
4 Putamen 11 Ventriculus iateraiis, cornu posterius
5 Nucleus accumbens 12 Pedunculus nuclei lentiformis
6 Ventriculus tertius 13 Ventriculus iateralis, cornu inferius
Fig. 29. The corpus striatum and the thalamus in lateral view. The contours of some parts of the
ventricular system are indicated (6/5 x)
Part II Vessels and Meninges
Arteries of the Brain
Veins of the Brain
Vessels of the Brain Stem
Meninges, Cisterns and Liquor System
Vessels and Meninges of the Spinal Cord

34 Vessels and Meninges
1 Cerebromeningeal anastomosis *
2 Calvaria (outer and inner surface)
3 Cerebrum (outer surface)
4 A. Callosomarginalis
5 A. pericallosa
6 Corpus callosum
7 A. cerebri anterior
8 A. supratrochlearis
9 A. dorsalis nasi •••
to Incisura frontalis
(Foramen frontale)
49
--_._-----
~--------
11 A. meningea anterior 34 A. cerebri media, pars insularis
12 A. lacrimalis 35 A. communican posterior
13 Foramen ethmoidale anterius 36 A. basilari
14 A. ethmoidalis anterior· .. • 37 A. cerebri posterior
15 Foramen ethmoidale posterius 38 Foramen parietalc"
16 A. ethmoidalis posterior 39 A. occipitalis
17 A. ophlhalmica 40 A. occipitalis, R. mastoideus··
18 Fissura orbitalis superior 41 Foramen mastoideum
19 A. meningea media, R. anastomoticus**** 42 A. meningea posterior
20 A. meningea media, R. frontalis 43 Junction of vertebral arteries
21 Canalis opticus 44 Foramen jugulare
22 A. conchae superioris (Anastomosis)*** 45 A. temporalis superficialis
23 A. sphenopalatina 46 A. pharyngea ascendens
24 A. infraorbitalis 47 A. carotis extern a
25 Canalis infraorbitalis 48 A. carotis interna
26 Foramen infraorbitale*** 49 A. carotis communis
27 A. angularis 50 A. vertebralis
28 A. facialis
29 A. maxillaris Anastomoses
30 A. meningea media 1 Cerebromeningeal *
31 Foramen spinosum 38+40 Extracraniomeningeal **
32 A. carotis interna, pars petrosa 9+22+26 Extracranio-orbital ***
33 A. meningea media, R. parietalis 14+19 Orbitomeningeal ****
Fig. 30. Collateral circulation in the arterial system of the head; semidiagrammatic lateral view (2/3 x).
Black: a. carotis externa with extracranial branches; Black hatched: a. vertebralis system (main trunk);
Solid red: meningeal arteries; Red hatched: a. carotis interna with orbital and lateral cortical branches;
Open red: medial cortical branches of a. carotis interna
Arteries of the Brain 35
17
1 Sulcus centralis 20 A. cerebri posterior, pars precommunicalis

2 Ramus marginalis sulci cinguli 21 Aa. centrales posteromediales
3 Precuneus 22 R. choroideus posterior medialis
4 A. precunealis 23 A. cerebri posterior, pars postcommunicalis
5 A. pericallosa, R. Posterior 24 Rr. thalamici (posteriores)
(anastomosing with 28) 25 A. occipitalis medialis
6 A . paracentralis 26 A. cingulothalamica
7 Sulcus cinguli 27 R. thalamicus (superior)
8 R. frontalis posteromedialis 28 R. corporalis callosi dorsalis
9 R. frontalis intermediomedialis (anastomosing with 5)
10 R . frontalis anteromedialis 29 R . parietalis
11 A. callosomarginalis 30 Sulcus parieto-occipitalis
12 A. pericallosa 31 R. parieto-occipitalis
13 A . mediana corporis callosi 32 R. calcarinus (in the depth
14 A. cerebri anterior, pars postcommunicalis of the calcarine sulcus)
15 A. communicans anterior 33 Rr. temporales posteriores
16 A . frontobasalis medialis 34 R. temporalis intermedius medialis
17 A. temporopolaris 35 A. occipitalis lateralis
18 A. carotis intern a 36 Sulcus collateralis
19 A. communicans posterior 37 Rr. temporales anteriores
Fig. 31. The arteries of the medial cerebral cortex; the anterior and posterior cerebral arteries (1 /1 x).
Of the posterior. cerebral artery some of its central branches are also shown. End branches of the
anterior cerebral artery, that reach the lateral side of the superior frontal gyrus can be seen in the
next figure. The Figs. 31-35 are all derived from the same specimen
26
25
1 Sulcus centralis
2 R . frontalis posteromedialis 15 Sulcus occipitalis transversus
3 R . frontalis intermediomedialis 16 A. temporo-occipitalis
4 R. frontalis anteromedialis 17 Sulcus temporalis superior
5 A. frontobasalis medialis 18 A. temporalis posterior
6 A. frontobasalis lateralis 19 A. temporalis intermedia
7 A. prefrontalis 20 Cist. fossae lateral is cerebri
8 Sulcus frontalis inferior 21 A. temporalis anterior
9 A. sulci precentralis 22 Cist. pontis
10 A. sulci centralis 23 N. abducens
11 A. sulci postcentralis 24 Cist. pontocerebellaris
(A. parietalis anterior) 25 Cist. medullaris
12 A. parietalis posterior 26 A. vertebralis
13 A. gyri angularis 27 Cist. cerebellomedullaris
14 Sulcus intraparietalis 28 A. cerebelli inferior posterior, R. lateralis
Fig. 32. The arteries of the lateral cerebral cortex: the middle cerebral artery (1 /1 x). In this figure
the lateral and medullar cisterns are left intact. On the lateral surface of the cerebellum one inferior
and two superior cerebellar branches are illustrated (cf. Fig. 40). On the superior frontal gyrus some
end branches of the anterior cerebellar artery can be seen
25
17 Sulcus lemporaJis superior

18 A. temporo-occipitalis
19 Sulcus lateralis, ramus posterior
20 A. sulci postcentralis
1 Sulcus centralis (a. parietalis anterior)
2 A. sulci centralis (branches) 21 A. parietalis posterior
3 Gyrus postcentralis 22 A. gyri angularis
4 Gyrus precentralis 23 Gyrus angularis
5 A. sulci precentralis 24 Sulcus intraparietalis
6 Sulcus frontalis inferior 25 Sulcus parieto-occipitalis
7 Gyrus frontalis inferior, 26 Sulcus lunatus
pars triangularis 27 Sulcus occipitalis anterior
8 A. prefrontalis (A. candelabra) *
9 A. frontobasalis lateralis Alternative subdivision
(in this case two separate branches)
11 + 15 A. temporalis anterior
10 Truncus anterior Ae. cerebri mediae
16 A. temporalis intermedia
(A. frontalis ascendens)
18 A. temporalis posterior
11 A. temporalis anterior (branches)
20 A. parietalis
12 A. temporopolaris
21 + 22 A. gyri angularis
13 Truncus intermedius Ae. cerebri mediae
14 Truncus posterior Ae. cerebri mediae *All branches of the truncus anterior
15 A. temporalis intermedia may be candelabra-like
16 A. temporalis posterior (the so-called candelabra group)
Fig. 33. The branches of the middle cerebral artery seen at their full extent; lateral view (1/1 x).
In this specimen, as in most cases, a trifurcation can be seen of the artery. The branches of the
anterior (frontal) trunk are illustrated in black and red; the branches of the middle (parietal) trunk
are in black only, the branches of the posterior (temporal) trunk are red coloured. The candelabralike
branching of especially the anterior trunk is a common phenomenon
1 Cis!. laminae terminalis

2 . opticus (N. II)
3 Cis!. valleculae cerebri 19 A. frontobasalis lateralis
4 Cis!. chiasma tis (branch of the middle cerebral artery)
5 A. carotis interna (pars cerebralis) 20 Gyrus frontalis inferior, pars orbitalis
6 N. oculomotorius (N. III) 21 A. temporopolaris
7 Hypophysis 22 A. temporalis anterior
8 Cist. interpeduncularis 23 Sulcus temporalis inferior
9 N. abducens (N. VI) 24 Sulcus collateralis; A. occipitalis lateralis
10 N. trochlearis (N. IV) 25 Rr. temporales anteriores
11 Cis!. pontis 26 Sulcus occipitotemporalis
12 Cis!. trigemini 27 A. vertebralis
13 Cis!. meatus acustici (interni); 28 A. cerebelli inferior posterior; Rr. laterales
Nn. VII + VIII 29 A. cerebelli inferior posterior, R. medialis
14 Nn.IX,X+XI 30 Fissura horizontalis cerebelli
15 Cis!. pontocerebellaris 31 R . temporalis intermedius medialis
16 Cist medullaris 32 Rr. temporales posteriores
17 Cis!. cerebellomedullaris 33 Sulcus occipitotemporalis
18 A. frontobasalis medialis 34 Sulcus collateralis
(branch of the anterior cerebral artery) 35 A. occipitalis lateralis
Fig. 34. The arteries of the brain viewed from the basal side (1 /1 x ). In this figure the basal, cerebellar
and medullary cisterns are left intact
1 A. tcmpo ropolari
2 Rr. temporales anleriorcs
3 A. temporalis anterior
4 A. temporalis intermedia
21 A. communicans posterior
5 A. temporalis posterior
22 R. hypothalamicus
6 R. temporalis intermedius medialis 23 R. thalamicus (anteroinferior)
7 A. occipitalis medialis
24 A. cerebri posterior, pars precommunicalis
8 A. occipitalis lateralis
25 Aa. centrales posteromediales
9 Rr. temporales posteriores
26 A. cerebri posterior, pars postcommunicalis
10 R. calcarinus (Ae. occipitalis medialis)
27 R. choroideus posterior medialis
11 A. frontobasalis medialis
28 A. choroidea anterior
12 A. frontobasalis lateralis
29 Rr. choroidei Ae. choroideae anterioris
13 A. cerebri media, pars insularis
30 R. choroideus posterior lateralis
14 Limen insulae
15 Aa. centrales anterolaterales, Rr. laterales 32 R. thalamicus (inferior)
16 Aa. centrales anterolaterales, Rr. mediales
33 R. thalamicus (posterior)
17 A. cerebri media, pars sphenoidalis
34 R . thalamicus (superior)
18 Aa. centrales anteromediales 35 R. corporis callosi dorsalis
19 A. communicans anterior
20 A. cerebri anterior, pars precommunicalis 19 + 20 + 21 + 24 Circulus arteriosus (left half)
Fig. 35. The cerebral arteries viewed from the basal side (1/1 x). Part of the left temporal lobe has
been removed to show the sphenoid part of the middle cerebral artery and the arterial supply of
the choroid plexus of the lateral ventricle. The lateral occipital artery has been interrupted to gain
a free sight on the diencephalic, mesencephalic and retrosplenial branches of the posterior cerebral
artery
Fig. 36. Cortical territories of the three cerebral arteries; semidiagrammatic lateral and medial views
of a left cerebral hemisphere (2/3 x). The territories correspond to the vascularisation pattern illustrated
in the Figs. 31-33. Stippled areas: sites of possible cerebrocerebral arterial anastomoses, mostly accord-
ing to Gillilan [402]
1 Nucleus caudatus 20 Nucleus dorsomedialis thalami

2 Putamen 21 Nucleus medialis thalami
3 Globus pallid us, pars lateralis 22 Nucleus anterior thalami
4 Globus pallid us, pars medialis 23 Globus pallidus, pars medialis
5 Thalamus 24 Cauda nuclei caudati
6 Substantia perforata anterior 25 A. choroidea anterior
7 Aa. centrales anterolaterales, Rr. laterales 26 Subthalamic area;
8 Aa. centrales anterolaterales, Rr. mediales Aa. centrales posteromediales
9 A. centralis longa (Heubneri) 27 Hypothalamic area;
10 Aa. centrales anteromediales R . hypothalamicus
11 A. cerebri anterior 28 Corpus amygdaloideum
12 Substantia perforata posterior 29 A. cerebri posterior
13 A. cerebri media, pars sphenoidalis 30 A. communicans posterior
14 A. hypophysialis superior 31 A. basilaris
15 A. hypophysialis inferior 32 A. vertebralis
16 A. carotis interna, pars cerebralis
17 A. carotis interna, pars cavernosa (carotissiphon)
18 A. carotis interna, pars petrosa
19 A. carotis interna, pars cervicalis
Fig. 37. The central arteries from the carotis and the vertebralis system in a frontal view (1 /1 x).
Substrate based on a reconstruction. The frontal section is perpendicular to the horizontal plane
of Frankfurt, passing through the center of the insula. The central arteries themselves have been
derived from different sources
OH OH
FH FH
1 Calvaria (inner oulline)

2 A. occipitalis medialis,
R. paricto-occipitalis
4 Ventriculus lateral is
5 Insula
6 A. occipitaJi media lis
7 A. cerebelli superior, R. mcdialis
8 A. occipitalis latcralis
9 Margo liber alae minoris ossis sphenoidalis
10 Intraosseous part of 11 (inconstant)
11 A. meningea media, R . frontalis 31 A. paracentralis
12 A. meningea media, R. parietalis 32 A. pericallosa
13 Margo superior partis petrosae ossis temporalis 33 A. callosomarginalis
14 A. cerebelli superior, R. lateralis 34 A. cerebri media, pars terminalis
15 A. cerebri posterior 35 A. cerebri media, pars insularis
16 A. cerebelli superior 36 A . cerebri anterior, pars postcommunalis
17 A. basilaris 37 A. communicans anterior
18 A. cerebelli inferior anterior 38 A. cerebri anterior, pars precommunicalis
19 A. cerebelli inferior posterior, R. medialis 39 A. cerebri media, pars sphenoidalis
20 A. cerebelli inferior posterior, R. lateralis 40 A. carotis interna, pars cavernosa (carotissiphon)
21 A. cerebelli inferior posterior 41 A. carotis interna, pars petrosa
22 A. vertebralis, pars intracranialis 42 A. carotis interna, pars cervicalis
23 A. maxillaris, pars pterygoidea 43 A. carotis communis
24 A. meningea media
25 A. temporalis superficialis OH: Upper horizontal plane (Kronlein):
26 A. maxillaris, pars mandibularis tangential to supraorbital margin
27 A. vertebralis, pars atlantis FH: Horizontal plane of Frankfurt (Reid):
28 A. carotis externa tangential to infraorbital margin
29 A. facialis Double arrow: Sulcus lateralis
30 A. vertebralis, pars transversaria Single arrow: Foramen magnum
Fig. 38. Orthogonal frontal projection of the cerebral and cerebellar arteries in situ together with
some bony landmarks and the lateral ventricles (2/3 x). The projection was made parallel to the
horizontal plane of Frankfurt by using a graphical reconstruction from the frontal slices of one speci-
men, and by cross-reference with Fig. 39. In this figure and the next an ample use have been made
of indications by Thijssen [cf. 1371]. Most vessels are illustrated only in one half of the skull; the
vertebral artery is shown bilaterally
OH
Be
FH FH
36 A. vertebrali • pars intracranialis

37 A. vertebralis, pars atlantis
38 A. carotis interna, pars cervicalis
39 A. maxillaris
1 Sulcus centralis 20 A. carotis interna, pars cerebralis 40 A. meningea media
2 A. pericallosa 21 A. carotis interna, pars cavernosa 41 A. carotis extern a
3 A. callosomarginalis 22 Siphon point 42 A. vertebralis, pars transversaria
4 Corpus callosum 23 A. cerebri media, pars sphenoidalis 43 A. carotis communis
5 Outline of ventricles 24 Ektokanthion (Canthus externus) 44 Medulla spinalis
6 Outline of insula 25 Glabella 45 Inion (Protuberantia occipitalis
7 A. cerebri anterior 26 Orbitale (on infraorbital margin) externa)
8 A. cerebri media, frontal trunk 27 A. carotis interna, pars petrosa
9 Commissura anterior 28 A. basilaris BC Bicommissuralline (Talairach)
10 A. cerebri media, parietal trunk 29 Margo superior partis petrosae ossis CM Canthus-Meatus line
11 A. cerebri media, temporal trunk temporalis FR Rorizontalline or plane of
12 Commissura posterior 30 A. cere belli inferior anterior Frankfurt (Reid)
13 A. occipitalis medialis 31 Porion (on suprameatal margin) GI Glabella - Inion line
14 A. occipitalis lateralis 32 Ventriculus quart us OR Upper horizontal line or plane
15 A. cerebelli superior, R. medialis 33 A. cerebelli inferior posterior, (Kriinlein)
16 A. cere belli superior, R. lateralis R. medialis VCA Vertical tangential to anterior
17 A. cere belli superior 34 A. cerebelli inferior posterior, commissure
18 A. cerebri posterior R.lateralis VCP Vertical tangential to posterior
19 A. communicans posterior 35 A. cerebelli inferior posterior commissure
Fig. 39. Orthogonal lateral projection of the cerebral and cerebellar arteries, together with external
and bony landmarks, in a schematized composition of data from different specimens and publications
(2/3 x) . Some neural structures are also illustrated in their outlines: the left hemisphere, cerebellum,
left insula, corpus callosum and ventricular system. Within the outlines of the orbita the bulbus oculi
and the optic nerve are indicated. On the outer side of the figure a number of reference lines are
added. In the center two lines tangential to the anterior (AC) and posterior (PC) commissures can
be seen: the one passing above the AC and beneath the PC is part of the bicommissural line of
Talairach (BC); the other tangent is part of the upper horizontal line of Kronlein (0H)
17 R. capsulae internae +>-
+>-
(R. thalamicus lateroinferior)
18 R. choroideus posterior medialis
19 A. cerebri posterior,
pars postcommunicalis
20 R. choroideus posterior lateralis
21 Rr. thalamici (posteroinferiores)
22 R. thalamicus (posterior)
23 A. occipitalis medialis
24 A. cingulothalamica
25 R. thalamicus (superior)
26 R. corporis callosi dorsalis
(anastomosing with 27)
27 A. pericallosa, R . posterior
28 A. vermis superior
29 A. cerebelli superior, R. medialis
30 A. cerebelli superior, R. lateralis
31 Colliculus inferior
32 R. mesencephalicus
34 A. basilaris
35 Aa. pontis mediales
36 Aa. pontis laterales
37 Nervus trigeminus
38 A. cerebelli inferior anterior
39 Nervus vestibulocochlearis
40 A. labyrinthi
1 A. pericallosa 41 Nervus facialis
2 Nucleus caudatus 42 Rr. medullares
3 Capsula interna 43 A. vertebralis
4 Thalamus 44 Radix spinalis nervi accessorii
5 Putamen 45 A. cere belli inferior posterior
6 A. cerebri anterior 46 A. cerebelli inferior posterior,
7 Aa . centrales anterolaterales, Rr. laterales 47
R. lateralis
8 Aa . cenlrales anterolatera les, Rr. mediales 47 A. cerebelli inferior posterior,
9 A. cerebri media, pars sphenoidalis R. medialis
10 Nervus opLieus
11 A. carotis intema, pars cerebralis
12 A. communicans posterior
~
en
13 R . hypothalamic us 11
en
14 A. choroidea anterior ~
15 R. thalamicus (anteroinferior) ::l
0-
16 Aa . centrales posteromediales
s:::
~
Fig. 40. The arteries of cerebellum, brain stem, thalamus and striate body in a lateral view (3/2 x). Some arteries are slightly simplified in order =:!.
to show their course and relations more clearly. The three arrow points indicate the choroid branches of the three choroidal arteries. The same ::l
specimen as in Figs. 31-35, with some slight simplifications ~

en
11 Nervus oculomotorius 24 Ganglion lrigeminale ~
12 Margin of incisura lenlorii 25 Cavum trigeminale S·
13 Sinus sphenoparielalis 26 Plexus venosus foraminis ovalis '"o-,
14 Nervus trochlearis 27 A. carotis inlema, pars petrosa
15 A.ophthalmica 28 A. carolis interna. pars cervicalis
16 V. ophlhalmica superior 29 Bulbus venaejugularis superior
~
t:l:I
17 Nervus oplicus 30 Sinus petrosus inferior ....
I>l
18 V. ophthalmica inferior (belween Nn. IX and X) S·
19 Ala major ossis sphenoidalis 31 V. sulci pontomedullaris
20 V. cerebri media supcrlicialis 32 Plexus basilaris
21 Nervus ophthalmicus 33 Vv. ponlis
22 Sinus cavemosus 34 Sinus petrosus superior
23 Nervus abducens 35 V. petrosa
36 Vv. ponlis Iransversae
(superior + inferior)
37 V. pontis laleralis
38 V. pedunculi cerebellaris superioris
39 Vv. hemispherii superiorcs
14 40 V. precentralis cere belli
41 V. vermis superior
16
42 Sinus reclus
11 43 V. vermis inferior
44 Con(Juens sinuum
45 Sinus lenlorii
(collecting infralenlorial veins)
18 46 Vv. hemispherii inferiores
47 Sinus lransversus
48 Sinus tentorii
(collecting supratenloria.1 veins)
49 Sinus sigmoideus
50 Inferior petrosal vein (inconslant)
51 Vv. spinalcs
1 Vv. insulares (anterior, lateralis, poslerior)
2 Vv. cerebri anleriores 52 Plexus venosus vertebralis
3 V. cerebri media profunda internus anterior
4 V. interpcduncularis 28 53 V. emissaria mastoidea
5 V. basalis 54 V. emissaria condylaris
6 V. cercbri interoa
7 V. cerebri magna
8 Vv. mesencephalicae
9 V. mesencephalica laleraJjs
10 V. ponlomesencephalica anterior
Fig. 41. Sinuses and veins of the diencephalon, brain stem and cerebellum in a lateral view (3/2 x). Composite drawing from two specimens with
additions from other sources. The cortical origins of the basal vein have been added: i.e. the insular veins, the deep middle cerebral vein and
the anterior cerebral veins. The tentorium bas been made fu1Jy transparent and the cavernous sinus has been deprived of its lateral dural wall. VI
The inner, lateral wall of the cavum trigeminale has also been removed. The orbita has been opened by a sagittal cut through its center
"""
I Vv. diploicae
2 Sinus sagittalis superior
3 Vv. cerebri superiores 26 Sinus sigmoideus
4 V. emissaria parietalis 27 Sinus pelrosus superior
5 Vv. temporales superficiales (parietal branch) 28 Sinus petrosus inferior
6 V. anastomotica superior (Trolard) 29 Plexu basilari
7 Sinus sagiltalis inferior 30 Vv. meningeae mediae
8 V. lhalamostriata superior 31 Sinu cavernosus
9 V. choroidea superior 32 Plexus pterygoideu
JOY cerebri interna 33 Y ophthalmica uperior
J 1 Y cerebri media superficialis 34 V. angulari
12 V. cerebri media profunda 35 V. ophthalmica inferior
13 Y choroidea inferior 36 Foramen infraorbitale
14 V. basal is 37 Y infraorbitalis
15 V. mesencephalica lateralis+ V. petrosa 38 Y faciei profunda
J 6 V. anastomotica inferior (Labbe) 39 Y facialis
17 V. cerebri magna 40 Y palatina
18 Sinus rectus 41 Vv. maxillares
19 Vv. cerebri inferiores 42 Vv. temporales superficiales (cf. no. 5)
20 Confluens sinuum 43 V. jugularis intema
21 V. emissaria occipitalis 44 V. retromandibulari
22 Sinus transversus 45 V. jugularis externa
23 Sinus occipitalis 46 Y cervicalis profunda
24 V. emissaria mastoidea 47 Plexu venosi vertebrale inlerni
25 V. emissaria condylaris 48 Y occipitali
Fig. 42. Collateral circulation in the venous system of the head; semidiagrammatic lateral view (2/3 x ).
Unpaired sinuses in the median plane are drawn without outlines; the extracranial veins draining
into the internal and external jugular veins are in black; between the intra vertebral venous plexuses
a fragment of the spinal medulla can be seen. The arrows indicate the continuity of the superficial
temporal veins
1 A. meniogea anterior 31 Conflueos sinuum

2 Sinus spheooparietalis (orifices of occipital sinus)
3 V. ophthalmica superior 32 Aa . ethomoidales anterio r
4 A. communicans anterior + posterior
5 A. cerebri anterior 33 A. lacrimalis
6 A. carotis iotema, pars cerebral is 34 V. diploica temporalis anterior,
7 Recessu opticus 19 ervus trigeminus (sensory and dtaining into sphenoparietal sinus
(on the optic chiasm) motor roots) 35 A. ophthalmica
8 Dor urn sellae 20 Meatus acusticus intemus ; 36 Cav um trigeminale
9 Sinus intercavemosus nervi VII + VUJ 37 A. basilaris + Plexus basilaris
(posterio r part) 21 Sinus petros us superior 38 V. pontomesencephalica anterior
10 A. meoingea media, 22 Sinu petrosus inferior 39 Aa. po ntis, Rr. latera le
R. anastomoticus 23 Inferior petrosal vein (inconstant) 40 A. ccrcbclli inferior an terior
II A. cerebri media 24 R. meningeus po terior 41 I ervus hypoglossu
12 ervus oculomotorius Ae. vertebralis + Canalis hypoglossal is
13 A. communicans posterior 25 Sinus sigmoideus 42 A. spinalis anterior
14 A. cerebri posterior 26 V. emissaria rna toidea 43 Sinus marginalis
15 A. meningea media, R . fron talis + R . meningeus Ae. occipitalis 44 A. cerebelli inferior posterior
16 Vv. meningeae mediae 27 Sinus transversus 45 Nervus acce sorius, R. spinalis
17 A. meningca media, R. parietalis 28 A. meningea posterior
18 Nervus abducens (piercing the 29 Falx cerebelli 4 + 5 + 11 + 13 + 14 Circulus arteriosus
dura mater) 30 Sinus occipitalis (left halJ)
Fig. 43. The skull base in an oblique view from the posterior right side (1 /1 x) . The dura has been
left in place with the exception of the roof of the greater part of the sinuses and the inner, lateral
sheet of the wall of the cavum trigeminale on the right The arterial circle of Willis is left intact;
the basilar artery has been interrupted to show the underlying subdural venous plexus of the clivus.
1 Sinus sagittalis superior 12 Sulcus parieto-occipital is 22 Nervus oculomotorius

2 Vv. prefrontales}Vv. cerebri (behind falx) 23 V. interpeduncularis
3 Vv. frontales superiores 13 Vv. cerebri anteriores 24 V. petrosa
4 Sulcus centralis 14 V. gyri olfactorii 25 Sinus petrosus superior
5 Lacunae laterales 15 Vv. gyrorum orbitalium (attachment of tentorium)
6 Vv. parietales } Vv. cerebri 16 Sinus sphenoparietalis 26 Vv. cerebri inferiores
7 Vv . occipitales superiores 17 Vv. thalamostriatae inferiores 27 Incisura tentorii
8 Sulcus cinguli (behind falx) 18 V. cerebri media superficialis 28 Tentorium cere belli
9 Sinus sagittalis inferior 19 Plica petroC\inoidea anterior 29 Sinus tentorii (collecting
10 V. corporis callosi dorsalis 20 Plica petroC\inoidea posterior supratentorial veins)
11 Falx cerebri 21 V. basalis 30 Sinus transversus
Fig. 44. The veins of the cerebral cortex; same view and same specimen as Fig. 43 (1/1 x). The falx
cerebri has been made transparent. In the drawing the most common drainage of the superficial
medial cerebral vein (into the cavernous sinus) and the basal vein (into the great cerebral vein of
Galen) has been illustrated, contrary to the actual situation in the specimen: drainage respectively
into the transverse sinus and via the petrosal vein only into the superior petrosal sinus. Variability
of venous patterns doesn't occur only interindividually, but also bilaterally: on the left seven individual
superior cerebral veins can be seen, of which one coming only from the medial side and one frontopolar
vein draining into the anterior tip of the superior sagittal sinus. On the right all superior veins drain
into the sinus via three stems altogether. For insular veins see next figure
1 Falx cerebri
2 Gyrus cinguli
(sinister)
3 Genu corporis
callosi
4 Nucleus caudatus 28 Sinus petrosus superior
(caput) 29 V. petrosa
5 Vv. septi pellucidi 17 Vv. insulares (anterior, 30 V. mesencephalica lateralis
anteriores centralis, posterior) 31 V. pedunculi cerebellaris
6 Vv. nuclei caudati 18 V. cerebri media profunda superioris
7 V. thalarnostriata superior 19 V. ba salis 32 V. pontis lateralis
8 V. choroidea superior 20 Vv. hippocampi anteriores 33 V. recessus lateralis ventriculi
9 Connecting vein between 8 and 10 21 Hippocampus quarti
10 V. thalami superior 22 V. ventricularis inferior 34 Pedunculus cerebellaris medius
11 Vv. cerebri intemae 23 V. thalamostriata superior 35 Pedunculus cerebellaris superior
(sinistra + dextral (inferior part) 36 V. vermis superior
12 V. atrii medialis 24 V. choroidea inferior 37 V. vermis inferior
13 V. basalis 25 Vv. temporales mediales 38 Sinus tentorii
14 V. cerebri magna 26 V. thalamostriata superior 39 Vv. hemispherii inferiores
15 Sinus rectus (superior part) 40 Vv. hemispherii superiores
16 Sinus transversus (medial extremity) 27 V. atrii lateral is 41 Sinus transversus (lateral extremity)
(atrial part of 23)
Fig. 45. The deep cerebral and the cerebellar veins; same view and same specimen as Figs. 43 and
44 (1 /1 x). The falx cerebri has been made transparent in order to show the left periventricular veins.
The veins of the medial and posterior thalamus can be seen on both sides; the inferior cerebellar
veins are shining through the cerebellum. The insula has been added to present the insular origins
of the deep middle cerebral vein. In the inferior part of the specimen the choroid plexus ends by
a cut, at the site where the hippocampus appears, the posterosuperior part of it being cut away
-
./, 2
4 2
6
21
24 Fissura horizontalis
36 25 V. vermis inferior
26 V. hemispherii inferior (medial branch)
27 A. vennis inferior
Nucleus caudatus (ventricular surface) 28 A. cerebelli inferior posterior, R. medialis
2 Vv. nuclei caudati 29 Retrotonsillar veins
3 V. septi pellucidi anterior 30 A. cerebelli inferior posterior, Rr. laterales
4 V. thalamostriata superior 31 Tonsilla cerebelli
5 Lamina affixa 32 R. tonsillae (of 35)
6 Plexus choroideus ventriculi lateralis 33 Rr. choroidei ventriculi quarti (of 35)
7 Septum pellucidum . 34 R . recessus latera lis ventriculi quarti (of 35)
8 Fornix 35 A. cerebelli inferior posterior
9 Foramen interventriculare 36 A. vertebralis
10 Plexus choroideus ventriculi tertii 37 A. cerebelli inferior anterior
11 V. cerebri interna 38 Aa. pontis, Rr. mediales
12 V. basalis 39 A. basilaris
13 A. vermis superior 40 A. cere belli superior, R. lateralis
14 V. cerebri magna 41 A. cerebelli superior, R. medialis
15 Commissura fornicis 42 Aa centrales posteromediales
16 Sinus sagittalis inferior 43 A. cerebri posterior, pars precommunicalis
17 V. precentralis cere belli 44 Nervus oculomotorius
18 V. vermis superior 45 A. communicans posterior
19 Sinus rectus 46 A. carotis interna
20 Falx cerebri 47 A mediana corporis callosi
21 Sinus sagittalis superior 48 A. cerebri anterior, pars precommunicalis
22 Confluens sinuum 49 A. communicans anterior
23 Sinus occipitalis 50 A. frontobasalis medialis
Fig. 46. Medial arteries and veins of the cerebellum; medial view of the left half of a bisected brain
(3/2 x). Most of the medial perforating arteries to the pons and the mesencephalon are also illustrated.
Superiorly the major subependymal veins on the caudate body and the internal cerebral vein can
be seen
1 A. quadrigemina (see 19) 22 V. precentralis cere belli

2 R. choroideus posterior medialis (see 18) 23 Vv. mesencephalicae (superior median collicular vein)
3 Colliculus superior 24 V. basalis (posterior segment)
4 Griseum centrale mesencephali 25 V. mesencephalica lateralis
5 Corpus geniculatum mediale 26 V. pontomesencephalica anterior (peripeduncular branch)
6 Corpus geniculatum laterale 27 Vv. pedunculares
7 Pedunculus cerebri 28 V. communicans posterior
8 Substantia nigra 29 V. interpeduncularis
9 Nucleus TUber 30 V. pontomesencephalica anterior (median branch)
10 Nervus oculomotorius 31 V. basalis (anterior segment)
11 Aa. centrales posteromediales
12 A. basilaris
13 A. cerebri posterior, pars precommunicalis
14 A. communicans posterior (with R . mesencephalicus)
15 A. cerebri posterior, pars postcommunicalis
16 A. cerebelli superior, R. lateralis
17 A. cerebelli superior, R. medialis;
arrow.' R . mesencephalicus, far more distally
18 R . choroideus posterior medialis
19 A. quadrigemina
20 Rr. pedunculares
21 A. choroidea anterior
Fig. 47. Vascularisation of the mesencephalon; transverse section in a superior view, with the vessels
of about that level (4/1 x). The plane of section is that of Fig. 110. Territories according to Duvernoy
[315], arteries and arterial territories in the right half, veins and their territories in the left half
1 Branch to inferior cerebellar peduncle 15 Ventriculus quartus

2 Nuclei vestibulares 16 Posterior tegmental veinlets, ascending to the lateral
3 Tegmentum pontis mesencephalic vein
4 Continuation of 11 to horizontal fissure 17 Vv. pontis transversae
5 Branch to middle cerebellar peduncle 18 V. petrosa
6 Anastomosis around trigeminal nerve (inconstant) 19 V. pontis lateral is (anterior cerebellar vein)
7 Radix sensoria nervi trigemini 20 Vv. pontis
8 Radix motoria nervi trigemini 21 V. pontomesencephalica anterior
9 Aa. pontis, R. lateralis (superolateral pontine artery)
10 Aa. pontis, R. lateralis (inferolateral pontine artery)
11 A. cerebelli inferior anterior
12 Pons (Nuclei pontis)
13 Aa. pontis, R. medialis
14 A. basilaris
Fig. 48. Vascularisation of the metencephalon; transverse section in a superior view, with the vessels
of about that level (4/1 x). The plane of sectioning is that of Fig. 112. Source and execution as
in Fig. 47
1 Nucleus cuneatus medialis 16 Veinlet descending to (posteromedian) medullary vein

2 Pedunculus cerebellaris inferior 17 V. medullae oblongatae (lateralis) . } ascendin to the lexus of
3 Nervus vagus 18 V. medullae oblongatae (anterolaterahs) g d' p. .
4 Rr. medullares (posteriores) 19 V. medullae oblongatae (median a) corrrespon mg pontme vems
5 Rr. medullares (laterales)
6 Nucleus nervi hypoglossi
7 Lemniscus medialis
8 Nucleus olivaris inferior
9 A cerebelli inferior posterior
10 Rr. medullares (mediales)
11 Rr. medullares (anterolaterales)
12 Tractus pyramidalis
13 A. spinalis anterior
14 Rr. medullares (anteromediales)
15 A. vertebralis
Fig. 49. Vascularisation of the myelencephalon; transverse section in a superior view, with the vessels
of about that level (4/1 x). The plane of section is that of Fig. 114. Source and execution as in
Fig. 47
D Nervous Ii sue
D Intraneural liquor pace

(ventricles)
r:
L.. _ _ _ J
Extraneural liquor space
(cisterns)
Dura mater (pachymeninx)
AraChnOid} (leptomenmges)
. 1 Sinus agittalis superior + Lacunae laterales
Pia mater 2 Granulationes arachnoideales
3 Cavitas subarachnoideaLi
Arteries 4 Vv. cerebri (cortical + deep cerebral veins)
Veins 5 Aa. cerebri (cortical + deep cerebral arterie )
6 Tela choroidea (of all ventricles)
Venous sinuses 7 Aa. choroideae (anterior + posterior
Lymphe vessels choroid arteries)
-
Blood 8 Vv. choroideae (superior + inferior
choroid veins)
Liquor cerebro pinalis 9 Connective ti ue
---~ Lymphe 10 Sinu rectus
A Liquor- blood (valve-like) passage 11 Sinus transver u + sinus igrnoideu
12 Aperturae (mediana + lateralis) ventriculi quarti
6 Blood-brain barrier of non-fenestrated endothelial wall +
13 A. carotis interna
arachnoidal barrier layer 14 V. jugulari interna
c Blood-liquor barrier of non-fenestrated epithelial wall + 15 Ductus thoracicus
arachnoidal barrier layer 16 ervi spinales
o Liquor-blood passage via lymphatic system 17 Filarnentum tenninaJe
Fig. 50. Diagram of the blood and liquor circulation in the brain. For reasons of simplification only
one channel represents the arterial supply instead of the two systems that exist in reality (Cf. next
figure)
Meninges, Cisterns and Liquor System 55

2 Sinus sagittalis inferior
3 Corpus callosum
4 Plexus choroideus ventriculi lateralis 21 A. carotis interna, pars cavernosa
5 V. cerebri interna 22 A. carotis interna, pars petrosa
6 Plexus choroideus ventriculi tertii 23 A. cere belli superior
7 Foramen interventriculare 24 A. basilaris
8 A. cerebri anterior 25 A. cerebelli inferior anterior
9 A. cerebri media 26 Plexus choroideus ventriculi quarti
10 V. basalis 27 R. choroideus of 28
11 V. cerebri magna 28 A. cerebelli inferior posterior
12 Sinus rectus 29 A. cere belli inferior posterior, R. medialis
13 Tentorium cerebelli 30 A. cere belli inferior posterior, R. lateralis
14 Confluens sinuum 31 V. vermis inferior
15 Sinus transversus 32 Canalis caroticus (inferior opening)
16 A. choroidea anterior 33 Foramen jugulare
17 Rr. choroidei posteriores mediales 34 Foramen magnum
18 Rr. choroidei posteriores laterales 35 A. carotis interna, pars cervicalis
19 A. cerebri posterior (here displaced inferiorly) 36 V. jugularis interna
20 A. communicans posterior 37 A. vertebralis
Fig. 51. Vascular supply of the choroid plexuses of the ventricles, as seen in a semidiagrammatic
lateral view of the left side (1/1 x). All vessels are paired, with the exeption of the basilar artery
(24), the superior and inferior sagittal sinuses (1, 2), the great cerebral vein of Galen (11), the straight
sinus (12) and the inferior vermian vein (31)
1 Lacunae latera les

2 GranuJationes arachnoideales
4 Vv. cerebri superiores
5 Arachnoidea (mater)
6 Dura mater encephali
7 Falx cerebri
8 Margo liber faJcis
9 Cist. pericallosa + A. pericallosa 23 Sinus sagittal is inferior
10 Pia mater (surface and cut edge) 24 Cist. fissurae transversae + V. cerebri interna
11 Cist. laminae terminalis + A. cerebri anterior 25 Cist. laminae tecti + V. basalis
12 Arachnoidea (surface and cut edge) 26 Cist. venae cerebri magnae + V. cerebri magna
13 Crista galli 27 Cist. cere belli superior
14 Lamina cribrosa 28 Sinus rectus + Tentorium cerebelli (cut)
15 Connection with cist. gyri olfactorii 29 Confluens sinuum
16 Sinus intercavernosus (anterior) }. d· h II 30 Protuberantia occipitalis interna
17 Sinus intercavernosus (posterior) In lap ragma se ae 31 Protuberantia occipitalis externa
18 Cist. chiasmatis 32 Falx cere belli
19 Cist. interpeduncularis 33 Sinus marginalis
20 Cist. pontis + A. basilaris 34 Tonsilla cerebelli (pial surface)
21 Plexus basilaris 35 Apertura median a ventriculi quarti
22 Cist. medullaris 36 Cist. cerebellomedullaris
Fig. 52. Meninges and cisterns from the medial side (1 /1 x). The sagittal section passes just left of
the falx cerebri and of the sinuses situated in the median plane, thus cutting through the lateral
lacunae. The pachymeninx (dura mater) is black outlined, the leptomeninges (arachnoidea and pia
mater) are in red. The cisterns are indicated with a screen in red
/ \
,
J \\
I
/
,I
l
/
"
1"'/
\
\
\I
!
I
I
\
1 Pericranium (periosteum cranii) 13 Venae diploicae

2 Lamina extern a } 14 Foveola granularis (Pacchioni)
3 Diploe Calvaria 15 Lacuna lateralis
4 Lamina interna 16 Endosteal part of dura mater
5 Dura mater encephali 17 Granulationes arachnoideales
6 Spatium subdurale (Pacchioni)
7 Arachnoidea (mater) 18 Endothelium
8 Cavitas subarachnoidealis 19 Sinus sagittalis superior
9 Pia mater 20 Venae cerebri superiores
10 Cortex cerebri 21 Falx cerebri
11 Venula 22 Villi arachnoideales
12 Vena emissaria 23 Vena meningea
Fig. 53. Diagrammatic frontal section through the superior sagittal sinus and surrounding structures,
showing the different compartments of blood and liquor and the layers involved (about 3/1 x)
Cisternae subarachnoideales Impressiones (structures in contact with cisterns)

a Granulationes arachnoideales 1 Gyrus precentralis
b Cistern of the central sulcus 2 Insula
c Cist. pericallosa 3 Genu corporis callosi
d Cist. laminae terminalis 4 Gyrus temporalis superior
e Cist. fossae lateralis cerebri 5 Gyrus parahippocampalis (temporal tip)
f Cist. valleculae cerebri 6 Tractus opticus
g Cist. chiasma tis 7 Fornix
h Cist. interpeduncularis 8 Splenium corporis callosi
Cist. fissura transversae 9 Vena cerebri magna
j Cist. venae cerebri magnae 10 Pulvinar thalami
k Cist. cere belli superior 11 Corpus genicula tum laterale
I Cist. ambiens 12 Subiculum
m Cist. pontis 13 Incisura tentorii (arrow)
n Cist. trigemini 14 Sulcus basilaris
o Cist. meatus acustici (interni) 15 Pedunculus cerebellaris medius
p Cist. pontocerebellaris 16 Nervi IX, X, XI
q Cist. cerebellomedullaris 17 Fissura horizontalis
r Cist. medullaris 18 Lobulus biventer
19 Vallecula cere belli
20 Foramen magnum
Fig. 54. Model of the cisterns after a spatial reconstruction in an oblique view from behind (1/1 x).
In the insert the site within the brain is indicated. The places of contact with different neural structures
can be recognized and are labeled
Cisternae subarachnoideales impressiones (structures in contact with cisterns)

a Cist. pericallosa Genu corporis callosi
b Cist. laminae terrninalis 2 Gyri orbitales
c Cist. valleculae cerebri 3 Tuber cinereum
d Cist. interpeduncularis 4 Corpus mamillare
e Cist. pon tis 5 Pedunculus cerebri
f Cist. crural is . . } Cist. ambiens 6 Fossa interpeduncularis
g Ala cisternae amblenlis 7 Pyramis (medullae oblongatae)
h Cist. trigemini 8 Corpus genicula tum laterale
Cist. meatus acustici (interni) 9 Corpus geniculatum mediale
j Cist. pontocerebellaris 10 Pulvinar thalami
k Cist. medullaris 11 Recessus suprapinealis
I Cist. cerebellomedullaris 12 Columnae fornicis
m Cist. cerebelli superior 13 Corpus fornicis
n Cist. pericallosa (posterior) 14 Crus fornicis
o Cist. venae cerebri magnae 15 Splenium corporis callosi
p Cist. laminae tecti 16 Vena cerebri magna
q Cist. fissurae transversale 17 Sulcus precentralis
r Cist. fossae lateralis cerebri 18 Operculum frontoparietale
19 Sulcus circularis insulae
20 Insula
Fig. 55. The same model as in the former figure, in an oblique view from above (1/1 x). To gain
a free sight on the central and lower cisterns, the cortical and sylvian cisterns of the left side and
part of the cortical cisterns of the right side are removed . Further procedure as in Fig. 54.
... 1 A. vertebralis, pars intracranialis

3 A. spinalis posterior
4 Rr. spinales
5 A. vertebralis, pars transversaria
6 A. radicularis posterior
7 A. radicularis anterior
8 A. cervicalis ascendens
9 A. cervicalis profunda (may also be source of
radicular arteries)
10 Truncus costocervicalis
11 A. subclavia
12 A. intercostalis suprema
13 Aorta
14 Hairpin junction of radicular artery to spinal artery
15 A. radicularis magna (Adamkiewiczi)
16 Ansa anastomotica
17 A. subcostalis
18 A lumbalis I
I, II etc. Aa. intercostales posteriores I, II etc.
C Cervical segments of the spinal cord
Co Coccygeal segment of the spinal cord
L Lumbar segment of the spinal cord
S Sacral segment of the spinal cord
T Thoracic segment of the spinal cord
1 A. spinalis posterior (dextra)

2 V. spinalis (posterior)
v 3 A. spinalis posterior (sinistra)
4 A. radicularis posterior (sinistra, C5)
5 V. radicularis anterior (sinistra, C5)
6 V. radicularis anterior (dextra, C5)
7 A. radicularis anterior (dextra, C5)
8 A. radicularis anterior (dextra, C6)
9 V. spinalis (anterior)
T 11 Radix ventralis nervi spinalis C6
Fig. 56A. Arterial supply of the spinal cord in a semi diagrammatic ventral view (slightly less than
1/2 x). Spinal and spinal root arteries are in solid red; dorsal arteries behind the medulla in broken
line. Arteries of origin and aorta red outlined. Of the spinal nerve roots only those are illustrated,
that belong to the same segment as the illustrated spinal root arteries. The levels of these arteries
are somewhat arbitrary, but about the most frequent according to different, through not fully concor-
dant sources (Rickenbacher [1126] and Djindjian [307])
Fig. 56B. Vascularisation areas in the spinal cord, from different sources. Left: arterial supply; right:
venous drainage. Upper section cf. Fig. 117
Vessels and Meninges of the Spinal Cord
1 V. spinalis (anterior)
2 V. spinalis (posterior)
3 V. vertebralis
4 V. cervicalis profunda
5 V. radicularis posterior
6 V. radicularis anterior
7 R. spinalis Ve. intercostalis posterioris
8 V. intercostalis suprema
9 V. subclavia
10 Bulbus venae jugularis inferior
11 V. brachiocephalica dextra
12 V. brachiocephalica sinistra
13 V. cava superior
14 V. intercostalis superior dextra
15 V. intercostalis superior sinistra
16 V. hemiazygos accessoria
17 V. azygos
18 V. hemiazygos
19 V. subcostalis
20 V. lumbalis ascendens
21 V. lumbalis I
J, II etc. Vv. intercostales posteriores J, II etc.
C Cervical segment of the spinal cord
Co Coccygeal segment of the spinal cord
L Lumbar segment of the spinal cord
S Sacral segment of the spinal cord
T Thoracic segment of the spinal cord
Fig. 57. Venous drainage of the spinal cord in a semidiagrammatic ventral view (slightly less than
1/2 x). Ventral spinal root veins are in solid black, dorsal ones with open outlines. Collecting veins
are half-hatched; the final path (the main branches of the superior vena cava) is characterized by
stippling. The lowest arrows indicate the drainage via the ascending lumbar veins into the common
iliac veins. Levels of the spinal root veins mostly according to Djindjian [307]
1 Costa VIII dex tra

2 Facies articularis capitis costae
3 Pediculus arcus vertebrae
4 Processus transversus
5 V. intercostalis posterior
6 A. intercostalis posterior
7 Nervus intercostalis (R. ventralis
nervi thoracici)
8 R. spinalis (Ve. intercostalis)
9 R. dorsalis nervi thoracici (VIII)
10 Foramen costotransversarium
11 Rr. communicantes
12 Truncus nervi thoracici (VIII)
13 A. spinalis posterior (dextra)
14 V spinalis posterior
15 Plexus venosus vertebralis internus
(posterior)
16 Medulla spinalis
17 Pia mater
18 Arachnoidea + Cavitas subarachnoidealis
19 Spatium subdurale
20 Dura mater spinalis
21 Cavitas epiduralis
22 Periosteum
23 Ligamentum denticulatum
24 Fibrous closure of the intervertebral
foramen
25 Dural sac of the spinal ganglion
and roots
26 Vv. spinales anteriores
27 Radix ventralis nervi thoracici (IX)
28 A. radicularis magna
30 V. radicularis posterior
31 R. spinalis
32 R. dorsalis
33 R. dorsalis (may drain into
intercostal vein)
34 V. hemiazygos
35 R. spinalis
36 V. radicularis anterior
37 Pia mater
38 Arachnoidea
40 Cavitas epiduralis
41 Ligamentum longitudinale posterius
42 R. meningeus nervi thoracici (X)
43 V. intervertebralis (superior)
44 R. communicans ad ramum
meningeum
45 V. intervertebralis (inferior)
46 Plexus venosus vertebralis internus
(anterior)
47 Vv. basivertebrales
48 Plexus venosus vertebralis externus
(anterior)
49 Corpus vertebrae thoracalis XI
50 V. azygos
Fig. 58. Topography of the contents of the vertebral canal at the level of the 8th to 11th rib; ventral
view, slightly from above (3/2 x). Semidiagrammatic representation. Vascular configurations are de-
rived from Figs. 56, 57 and 59. Pachymeninx in black outlines; leptomeninges in red. The dural sac
around the spinal ganglion and the ventral root has been cut in different ways. At the left side
the spinal roots are left out with the exception of part of the ventral root of Tl 0
Vessels and Meninges of the Spinal Cord 63
I Dura mater spinali 5

2 Conus medullaris
3 Medial epi.ddural vein} Plexus venosus vertebralis internus (anterior)
4 Lateral ept ura l vem
5 V. azygos
6 Ligamentum arcuatum mediale
7 V. lumbalis ascend ens
8 Pediculus arcus vertebrae lumbali
9 Plexus venosus vertebralis internus (posterior)
10 Spinal roots within dural sheath (between epidural veins)
11 V. intervertebralis (superior)
12 V. intervertebralis (inferior)
13 Segmental communicating vein
14 R. dorsalis (Ve. lumbalis ascendentis)
15 R. dorsalis (Ve. intervertebralis)
16 Plexus venosus vertebralis externus (posterior)
17 Ligamentum longitudinale posterius
18 Retrocorporeal venous anastomosis
19 V. basi vertebra lis
20 Plexus venosus vertebralis internus (anterior)
21 R. dorsalis Ve. sacralis lateralis (ascending sacral vein)
22 V. sacralis lateralis
23 V. iliaca communis
24 V. iliaca extema
25 V. iliaca intema
26 Pediculus arcus vertebrae sacral is III
27 Foramina sacralia pelvina
28 Foramina sacralia dorsalia
Fig. 59. The epidural veins of -the lumbar and sacral vertebral canal in a dorsal view (2/3 x). At
the dorsal and right sides the wall of the bony canal has been removed, unto the middle of the
pedic1es of the vertebral arch. At right the lateral outline of the sacrum is indicated in a broken
line. At the level of the 5th lumbar vertebra the transverse vein has been interrupted to show the
sagittal vein piercing the body of the vertebra (v. basivertebralis). This figure is mainly based on
Theron and Moret [1370] and Renard c.s. [1116]
64 Vessels and Meninges of the Spinal Cord
1 Cavitas cpiduralis
2 Cavitas subarachnoidalis
3 Sixth cervical spinal segment
5 Arachnoidea
6 Pia mater
7 Eighth thoracic spinal segment
8 Second to third lumbar spinal segment
9 Cauda equina (fila radicularia)
A
Fig. 60. A Semidiagrammatic presentation of the spinal cord and the meninges within the vertebral
canal at four different levels (1/1 x). Outlines of the vertebrae as in a superior view; the position
of the sections through the spinal cord has been indicated in the additional figure (8) of the median
sectioned vertebral column (1 /5 x)
Part III Brain Slices
Coronal Sections
Sections Perpendicular to the Axis

of the Brain Stem
Sagittal Sections
Horizontal Sections
66 Brain Slices
72 71 70 69 68 67 66 65 64 63 62
A
Coronal Sections 67
Fig. 61 A, B. Key diagrams showing the level and plane of the coronal sections in Figures 62-72 (2/3 x)
68 Brain Slices
15
1 Fissura longitudinalis cerebri 10 Sulcus cinguli

2 Gyrus frontalis superior 11 Gyrus cinguli
3 Sulcus frontalis superior 12 Genu corporis callosi
4 Gyrus frontalis medius 13 Radiatio corporis callosi
5 Sulcus frontalis inferior 14 Gyrus rectus
6 Gyrus frontalis inferior 15 Tractus olfactorius
7 Sulci orbitales
8 Gyri orbitales
9 Sulcus olfactorius
Fig. 62. Section through the anterior part of the cerebral hemispheres (6/5 x)
Coronal Sections 69
1 Sulcus corporis callosi 9 Stria longitudinalis medialis

2 Gyrus frontalis inferior 10 Stria longitudinalis lateralis
3 Sulcus circularis insulae 11 Radiatio corporis callosi
4 Sulcus lateralis 12 Truncus corporis callosi
5 Gyri breves insulae 13 Septum pellucidum
6 Gyrus temporalis superior 14 Ventriculus lateralis, cornu anterius
7 Sulcus temporalis superior 15 Rostrum corporis callosi
8 Gyrus temporalis medius 16 Caput nuclei caudati
18 Putamen
19 Capsula externa
20 Claustrum
21 Capsula extrema
22 Tractus olfactorius
Fig. 63. Section through the head of the caudate nucleus and the putamen (6/5 x)
70 Brain Slices
1 Gyrus frontalis superior 20 Indusium griseum

2 Sulcus frontalis superior 21 Stratum sUbependymale
3 Gyrus frontalis medius 22 Corona radiata
4 Sulcus precentralis 23 Caput nuclei caudati
5 Gyrus precentralis 24 Pontes grisei caudatolenticulares
6 Sulcus lateralis, ramus posterior 25 Capsula interna, crus anterius
7 Sulcus circularis insulae 26 Putamen
8 Gyrus frontalis inferior, pars opercularis 27 Lamina medullaris lateral is
9 Gyri breves insulae 28 Globus pallidus
10 Sulcus centralis insulae 29 Columna fornicis
11 Gyrus longus insulae 30 Commissura anterior
12 Gyrus temporalis medius 31 Lamina terminalis
13 Sulcus temporalis inferior 32 Substantia innominata
14 Gyrus occipitotemporalis lateralis 33 Arteria cerebri media, rami striati
15 Sulcus collateralis 34 Stria olfactoria lateralis
16 Sulcus rhinalis 35 Recessus opticus
17 Gyrus parahippocampalis 36 Chiasma opticum
18 Substantia perforata anterior 37 Infundibulum
19 Gyrus diagonalis
Fig. 64. Section through the anterior commissure and the optic chiasm (6/5 x)
Coronal Sections 71
1 Sulcus circularis insulae 16 Ventriculus lateral is, pars centralis

2 Gyri breves insulae 17 Corpus nuclei caudati
3 Sulcus centralis insulae 18 Vena thalamostriata
4 Gyrus longus insulae 19 Plexus choroideus ventriculi lateralis
5 Gyrus temporalis superior 20 Corpus fomicis
6 Sulcus temporalis superior 21 Foramen interventriculare
7 Gyrus temporalis medius 22 Nucleus anterior thalami
8 Sulcus temporal is inferior 23 Capsula intema, genu
9 Gyrus temporalis inferior 24 Putamen
10 Gyrus occipitotemporalis lateralis 25 Lamina medullaris lateralis
11 Sulcus rhinalis 26 Globus pallid us, pars lateralis
12 Gyrus para hippocampal is 27 Lamina medullaris medialis
13 Gyrus ambiens 28 Globus pallid us, pars medialis
14 Gyrus semilunaris 29 Pedunculus thalami inferior
15 Arteria cerebri media, rami striati 30 Ventriculus tertius
31 Columna fomicis
33 Tractus opticus
34 Hypothalamus
35 Infundibulum
36 Corpus amygdaloideum
Fig. 65. Section through the interventricular foramen, the infundibulum and the amygdaloid body
(6/ 5 x )
72 Brain Slices
1 Gyrus frontalis superior 18 Radiatio corporis callosi 26 Nucleus ventralis lateralis

2 Sulcus precentralis 19 Truncus corporis callosi 27 Nucleus medialis thalami
3 Gyrus precentralis 20 Corona radiata 28 Nucleus reticularis thalami
4 Sulcus cinguli 21 Septum pellucidum 29 Capsula interna, crus posterius
5 Gyrus cinguli 22 Corpus nuclei caudati 30 Ventriculus tertius
6 Sulcus corporis callosi 23 Stria terminalis 31 Zona incerta
7 Indusium griseum 24 Nucleus anterior 32 Tractus mamillothalamicus
8 Sulcus centralis thalami 33 Nucleus subthalamicus
9 Gyrus postccntralis 25 Plexus choriodeus 34 Fasciculus mamillaris prmceps
10 Sulcus lateralis, ramus posterior ventriculi tertii 35 Substantia nigra
11 Gyrus uncinatus 36 Corpus mamillare
12 Gyrus ambiens 37 Pedunculus cerebri
13 Gyrus para hippocampal is 38 Corpus amygdaloideum
14 Sulcus collateralis 39 Stria terminalis
15 Gyrus occipitotemporalis lateralis 40 Cauda nuclei caudati
16 Sulcus occipitotemporalis 41 Ventriculus lateralis, cornu inferius
17 Gyrus temporalis inferior 42 Pes hippocampi
43 Pons
Fig. 66. Section through the anterior end of the hippocampus, the mamillary body and the mamillotha-
lamic tract (6/5 x)
Coronal Sections 73
1 Sulcus lateralis, ramus posterior 12 Fissura longitudinalis cerebri 18 Nucleus reticularis thalami
2 Sulcus circularis insulae 13 Ventriculus lateralis, 19 Capsula intern a, crus posterius
3 Gyrus longus insulae pars centralis 20 Capsula extrema
4 Gyrus intralimbicus 14 Corpus fornicis 2t Claustrum
5 Sulcus hippocampi t 5 Nucleus lateralis dorsalis 22 Capsula externa
6 Gyrus dentatus 16 Nucleus medialis thalami 23 Putamen
7 Gyrus parahippocampalis 17 Nucleus ventralis lateralis 24 Globus pallidus
8 Sulcus collateralis 25 Nucleus ventralis posterolateralis
9 Gyrus occipitotemporalis lateralis 26 Nucleus centromedianus
10 Sulcus occipitotemporalis 27 Ventriculus tertius
11 Gyrus temporalis inferior 28 Nucleus ruber
29 Nucleus subthalamicus
30 Tractus opticus
31 Capsula interna, pars sublentiformis
32 Plexus choroideus ventriculi lateralis
34 Substantia nigra
35 Fossa interpeduncularis
36 Pons
Fig. 67. Section through the thalamus, the cerebral peduncle and the pons (6/5 x)
74 Brain Slices
4
5
17 Septum pellucidum
19 Plexus choroideus ventriculi lateralis
20 Corpus fornicis
21 Crus fornicis
22 Plexus choroideus ventriculi tertii
23 ucleus medialis thalami
1 Lobulus paracentralis 24 uclei pulvinares
2 Gyrus precentralis 25 ucleus lateralis posterior
3 ulcus centrali 26 Ponte grisei caudatolenticulares
4 Gyrus postcentrali 27 Capsula interna, pars retrolentiformi
5 Sulcus postcentral is 28 uclei habenulae
6 Lobulus parietalis inferior 29 Commissura posterior
7 Sulcus lateralis, ramus posterior 30 Aqueductus cerebri
8 Pla num temporale + gyri temporales transversi (He chi) 31 Corpus geniculatum mediale
9 Gyru tempora.1i uperior 32 Corpus geniculatum laterale
10 Sulcus tem poralis uperior 33 Cauda nuclei caudati
11 Gyrus temporalis mediu 34 Ventriculus la tera lis, cornu inferius
12 Sulcus tem poralis inferior 35 Hippocam pus
13 Gyrus temporalis inferior 36 Pedunculus cerebri
14 Sulcus occipitotemporalis 37 Decu sa tio peduncu lo rum cerebellarium
15 Gyrus occipitotemporalis lateralis superiorum
16 Gyrus parahippocampalis 38 Pons
Fig. 68. Section through the posterior part of the thalamus (6/5 x)
Coronal Sections 75
1 Sulcus centralis
2 Lobulus paracentralis 15 Sulcus corporis callosi 27 Colliculus inferior
3 Sulcus cinguli, pars marginal is 16 Indusium griseum 28 Aqueductus cerebri
4 Gyrus postcentralis 17 Truncus corporis callosi 29 Pedunculus cerebellaris superior
5 Sulcus postcentralis 18 Ventriculus lateralis 30 Hemispherium cere belli
6 Lobulus parietalis inferior 19 Stratum subependymale 31 Pedunculus cerebellaris medius
7 Sulcus lateralis, ramus posterior 20 Splenium corporis callosi 32 Flocculus
8 Gyrus fasciolaris 21 Crus fornicis 33 Nervus glossopharyngeus
9 Fasciola cinerea 22 Plexus choroideus ventriculi 34 Nervus vagus
10 Gyrus dentatus lateralis 35 Nervus accessorius
11 Gyrus parahippocampalis 23 Cauda nuclei caudati 36 Plexus choroideus ventriculi quarti
12 Sulcus collateralis 24 Fimbria hippocampi 37 Oliva
13 Gyrus occipitotemporalis lateralis 25 Alveus hippocampi 38 Pyramis
14 Sulcus occipitotemporalis 26 Hippocampus 39 Nervus hypoglossus
Fig. 69. Section through the inferior colliculus and the inferior olive. The thickness of this slice is
three times the standard of 2 mm (6/5 x)
76 Brain Slices
21 Cauda nuclei cauda ti

22 Plexu choroideus ventriculi laterali
23 Radiatio oplica
24 Vermi cere belli
I LobuJus parielalis superior 25 Hemi pherium cerebelli
2 Sulcus inlraparielalis 26 Velum medullare superius
3 Lobulus parielali inferior 27 Pedunculus cerebellaris superior
4 Gyrus lemporalis superior 13 Lobulus paracentrali 28 Ventriculus quartus
5 Gyru lemporalis medius 14 ulcu cinguli, 29 Plexus choroideus ventriculi quarti
6 Gyrus temporali inrerior pars marginali 30 Pedunculus cerebellaris medius
7 Gyrus occipilotemporaJis lateralis 15 Precuneus 31 Pedunculus cerebellaris inferior
8 Sulcus collaleralis 16 Sulcus subparietalis 32 Flocculus
9 Gyrus occipi totemporalis medialis 17 Gyru cinguli 33 Recessus lateralis ventriculi quarti
10 Sulcus calcarinus 18 Splenium corporis callosi 34 Plexus choroideus ventricul i quarti
11 Isthmus gyri cinguli 19 Ventriculus latera li 35 Oliva
12 Gyrus rasciolaris 20 Fimbria hippocampi 36 Pyramis
Fig. 70. Section through the splenium of the corpus callosum and the fourth ventricle (6/5 x)
Coronal Sections 77
t 2 Precuneus
t 3 Sulcus subparietali
14 Gyru cinguli
15 Radiatio corporis callo i
~27 16 Ventriculus lateralis,
cornu posteriu
17 Radiatio optica
1 Lobulus parietali superior 18 Verrni cerebelli
2 Sulcus intraparietalis t 9 Hemispherium cerebelli
3 Lobulus parietali inferior 20 ucleu emboliforrnis
4 Gyrus angularis 21 ucleu fa tigii
5 Gyrus temporal is uperior 22 ucleus globo u
6 Gyrus temporalis mediu 23 ucleu dentatu
7 Gyrus temporal is inferior 24 Corpus medullare cerebclli
8 Gyrus occipitotemporali laterali 25 Laminae albae cerebell i
9 Sulcus collateralis 26 Medulla oblongata
10 Gyrus occipitotemporalis mediali 27 Canalis centralis
11 Sulcus calcarinus 28 Medulla spinalis
Fig. 71. Section through the posterior horns of the lateral ventricles and the central cerebellar nuclei
(6/5 x )
78 Brain Slices
1 Fissura longitudinalis cerebri 14 Stria Gennari

2 Lobulus parietalis superior 15 Area striata
3 Sulcus intraparietalis 16 Radiatio corporis callosi
4 Lobulus parietalis inferior 17 Ventriculus lateralis, cornu posterius
5 Sulcus parieto-occipitalis 18 Radiatio optica
6 Precuneus 19 Vermis cere belli
7 Cuneus 20 Hemispherium cerebelli
8 Sulcus calcarinus
9 Gyrus temporalis medius
10 Sulcus occipitalis anterior
11 Gyri occipitales
12 Gyrus occipitotemporalis lateralis
13 Gyrus occipitotemporalis medialis
Fig. 72. Section through the deepest part of the calcarine sulcus (6/ 5 x)
Sections Perpendicular to the Axis of the Brain Stem 79
Fig. 73. Key diagrams showing the level and plane of the sections perpendicular to the axis of the
brain stem in Figures 74-77 (2/3 x)
80 Brain Slices
1 Sulcus lunatus
2 Gyri occipitales
3 Cuneus
4 Sulcus parieto-occipital is
5 Sulcus occipitalis anterior
6 Gyrus temporalis medius
7 Ventriculus lateralis, cornu posterius 20 Radiatio corporis callosi
8 Plexus choroideus ventriculi lateralis 21 Radiatio optica
9 Sulcus temporalis superior 22 Culmen
10 Gyrus dentatus 23 Colliculus inferior
11 Gyrus parahippocampalis 24 Aqueductus cerebri
12 Gyrus intralimbicus 25 Brachium colliculi inferioris
13 Limbus Giacomini 26 Tegmentum mesencephali
14 Sulcus hippocampi 27 Pedunculus cerebellaris superior
15 Gyrus uncinatus 28 Substantia nigra
16 Incisura unci 29 Pedunculus cerebri
17 Gyrus ambiens 30 Pons
18 Ventriculus lateralis, cornu inferius 31 Cornu ammonis
19 Sulcus rhinalis 32 Corpus amygdaloideum
Fig. 74. Section through the middle of the midbrain (6/5 x)

1 Sulcus lunatus 9 Stria Gennari

2 Gyri occipitales 10 Area striata
3 Cuneus 11 Radiatio optica
4 Sulcus calcari n us 12 Culmen
5 Sulcus occipitalis anterior 13 Lobulus centralis
6 Gyrus occipitotemporalis medialis 14 Lobulus quadrangularis
7 Gyrus occipitotemporalis lateralis 15 Ala lobuli centralis
8 Gyrus temporalis inferior 16 Pedunculus cerebellaris superior
17 Velum medullare superius
18 Tegmentum pontis
19 Pons
20 Nervus trigeminus
Fig. 75. Section through the pons at the level of the entrance of the trigeminal nerve (6/5 x)
82 Brain Slices
1 Sulcus calcarinus 16 Nucleus fastigii

2 Gyri occipitales 17 Nucleus globosus
3 Sulcus occipitalis anterior 18 Nucleus emboliformis
4 Gyrus temporalis inferior 19 Nucleus dentatus
5 Declive 20 Pedunculus cerebellaris inferior
6 Lobulus semilunaris superior 21 Pedunculus cerebellaris medius
7 Lobulus simplex 22 Tegmentum pontis
8 Nodulus 23 Tractus corticospinalis
9 Plexus choroideus ventriculi quarti
11 Flocculus
13 Nervus facial is
14 Pons
15 Nervus abducens
Fig. 76. Section through the transitional area of pons and medulla oblongata (6/5 x )
1 Sulcus calcarinus 17 Nucleus dentatus

2 Gyri occipitales 18 Pedunculus cerebellaris medius
3 Gyrus occipitotemporalis lateralis 19 Tegmentum myelencephali
4 Gyrus occipitotemporalis medialis 20 Nucleus olivaris inferior
5 Declive 21 Tractus corticospinalis
6 Lobulus simplex
7 Lobulus semilunaris superior
8 Pyramis vermis
9 Tonsilla cere belli
10 Uvula vermis
12 Tela choroidea ventriculi quarti
13 Recessus lateral is ventriculi quarti
14 Flocculus
15 Plexus choroideus ventriculi quarti
16 Nervus glossopharyngeus
Fig. 77. Section through the medulla oblongata (6/ 5 x)

84 Brain Slices
82 81 8079 79 80 81 82
Fig. 78. Key diagrams showing the level and plane of the sagittal sections in Figures 79-82 (2/3 x)
Fig. 79. Section through the mamilJary body, r./l
1 Sulcus postcentral is !l'
the red nucleus and the fornix (6/5 x ) t{9.
2 Sulcus parieto-occipitalis ........
3 Cuneus e:..
4 Area striata r./l
(1)
5 Sulcus calcarinus $:a.
6 Gyrus occipitotemporalis o·
medialis ::l
rJl
7 Lamina quadrigemina
8 Tegmentum mesencephali
9 Locus coeruleus 19 Sulcus paracentralis 29 Columna fornicis
10 Pedunculus cerebellaris superior 20 Corpus callosum 30 Commissura anterior
t t Ventriculus quartus 21 Corpus fornicis 3 t Nucleus ruber
12 Tela choroidea ventriculi quarti 22 Stria medullaris thalami 32 Decussatio pedunculorum
13 Tractus tegmentalis centralis 23 Ventriculus lateralis cerebellarium superiorum 00
Vl
14 Tonsilla cerebelli 24 Nucleus anterior thalami 33 Substantia nigra
15 Pyramis 25 Nucleus medialis thalami 34 Corpus mamillare
16 Nucleus ol.ivaris inferior 26 Nuclei habenulae 35 Nucleus oculomotorius
17 Nucleus olivaris accessorius dorsalis 27 Tractus mamillotegmentalis 36 Chiasma opticum
18 Nucleus olivaris accessorius medialis 28 Tractus mamillothalamicus 37 Pons
1 Sulcus paracentralis
2 Sulcus centralis
Fig. 80. Section through the head of the
3 Sulcus postcentralis caudate nucleus, the middle part of the 00
0'1
~ thalamus and the cerebral peduncle (6/ 5 x)
5 Gyrus postcentral is
6 Lobulus parieLalis superior
7 Sulcus parieto-occipitalis
8 Gyri occipitales
9 Area stria La
10 Sulcus calca rinus
--
-y
11 Gyrus occipito-
Lemporalis medialis 22 Ventriculus lateralis 33 Lemniscus medialis
12 Nervus trochlearis 23 Crus fornicis 34 Area tegmentalis
t3 Nucleus denLatus 24 Stria terminalis 35 Nucleus subtha lamicus
14 Pedunculus cerebellaris inferior 25 Nucleus venLralis latera lis 36 Capsula interna, genu
15 Pedunculus cerebellaris medius 26 Nuclei pulvinares 37 Commissura anterior
16 Pons 27 Lamina medullaris interna
..,to
38 Globus pallid us Pl
17 Tractus pyramidalis 28 Pedunculus thalami anterior 39 Substantia nigra S·
18 Tonsilla cercbclli 29 Caput nuclei caudati 40 Pedunculus cerebri ~
19 Recessus lateralis ventriculi quarti 30 Nucleus centromedianus 41 Tractus opticus g.
20 Nervi craniales 31 Brachium colliculi superioris 42 Uncus on
21 Lobulus biventer 32 Brachium colliculi inferioris 43 Tractus olfactorius
1 Sulcus centraJis
2 Sulcus parieto-occipitaJis Fig. 81. Section through the caudate nu-
en
3 Corpus callosum cleus, the globus pallidus and the medial III
4 Ventriculus lateraJis geniculate body (6/5 x ) C!9.
....
5 Crus fornicis
....
e:.
6 Gyrus fasciolaris en
7 Gyri occipitales g
8 Radia tio optica
....
o·
:l
9 Sulcus calcarinus en
10 Area striata
26 Corpus genicul atum mediale

11 Gyrus occipitotemporalis 27 Globus paUidus
medialis 28 Commissu ra anterior
12 Fissura prima 29 Putamen
13 Gyrus parahippocampalis 30 Pedunculus cerebri
14 Fissura horizontalis 31 Tractus opticus
15 Nucleus dentalus 32 Limbus Giacomini
16 Pedunculus cerebellaris med ius 21 N ucleus caudatus 33 Gyrus uncinalus
17 Nervus trigeminus 22 Nucleus lateralis posterior 34 Gyrus semilunaris 00
18 Flocculus 23 Capsula interna 35 Ventriculus lateralis. co rnu inferius -.J
19 Fissura posterolateralis 24 Nuclei pulvinares 36 Cornu amm onis
20 Fissura prebiventeris 25 Nucleus vent ralis posterolateralis 37 Corpus amygdaloideum
1 Sulcus centralis
2 Gyrus postcentralis
Fig. 82. Section through the hippocampus, the
3 Sulcus postcentralis putamen and the amygdaloid body (6/5 x)
00
4 Lobulus parietalis superior 00
5 Lobulus parietalis inferior

6 Radia lio corporis callosi
7 Radialio optica
8 Sulcus calcarinus
9 Ventriculus latcralis
10 Area striata
II Gyri occipilales
12 Gyrus occipilO-
temporalis lateralis
22
26 Capsula inlerna
27 Radialio acustica
28 Radiatio oplica
29 Corpus gcniculatum latera Ie
31 Putamen
32 Pedunculus nuclei lentiformis
13 Gyrus dentatus 33 Commissura anterior tl:i
...,
14 Fimbria bippocampi 20 Gyrus precenlralis 34 Claustrum ~
15 Subiculum 21 Sulcus precen lralis 35 Limen insulae S·
16 Laminae albae cerebclli 22 Gyri frontalis 36 Gyri orbilales ~
17 Corpus medullare cerebclli 23 Corona radiata 37 Polus insulae §"
18 Ventriculus lateralis, cornu inferius 24 Cauda nuclei caudali 38 Corpus amygdaJoideum
19 Gyrus occipitotemporalis lateralis
'"
25 Stria terminalis 39 Gyrus temporalis superior
Horizontal Sections 89
85
86
87
Fig. 83. Key diagrams showing the level and plane of the horizontal sections in Figs. 84-87 (2/3 x).
The planes of sectioning are about parallel to the bicommissural line of Talairach (broken line BeL,
just above 86), corresponding to the CT 0° direction
90 Brain Slices
1 Forceps minor
2 Fasciculus occipitofrontalis
superior
3 Corona radiata 19 Gyrus postcentral is
4 Fasciculus longitudinalis superior 20 Sulcus postcentralis
5 Ventriculus lateralis, par centralis 12 Sulcus cinguli 21 Sulcus lateralis ramus posterior
6 Truncus corpori callosi 13 Gyrus cinguli 22 Sulcus subparietalis
7 Vena thalamostriata 14 Gyri frontales 23 Sulcus parieto-occipitalis
8 Lamina affixa 15 Sulcus precentralis 24 Cuneus
9 Plexus choroideus ventriculi lateralis 16 Gyrus precentralis 25 Sulcus occipitalis anterior
10 Forceps major 17 Sulcus centralis 26 Gyri occipitales
11 Stratum sagittale J8 Corpus nuclei caudati 27 Sulcus lunatus
Fig. 84. Section through the corpus callosum and the body of the caudate nucleus. The thickness
of this slice is twice the standard of 2 mm (6/5 x)
1 Fasciculus occipito-
frontalis superior
2 Genu corporis callosi 24 ucleu ventralis lateralis
3 Cavum septi pellucidi 25 ucleu s mediali thalami
4 Ventriculus lateralis, 26 ucleu laterali po terior
cornu aoteriu 27 uclei habenulae
5 Cap ula interna, crus anteriu 28 Nuclei pulvinare
6 Fornix 15 Ventriculus latera lis, cornu inferius 29 Colliculus superior
7 Stria terminali 16 Radiatio corporis callosi 30 Cauda nuclei caudati
8 Capsula intema, genu 17 Gyri frontales 31 Fasciola cinerea
9 Cap ula interna, crus posterius 18 Caput nuclei caudati 32 Gyrus fasciolaris
10 Ventriculu tertius 19 Claustrum 33 Gyri Andreae Retzii
11 Recessu uprapinealis 20 Sulcu latcralis, ramus asccndens 34 Corpus pineale
12 Fasciculus longitudinali superior 21 Sulcu laterali, ramus posterior 35 Vermi cerebelli
13 Radialio oplica 22 ucleus lent iformis 36 Sulcu calcarinu
14 Fimbria hippocampi 23 ucleu anterior thalami 37 Gyri occipitales
Fig. 85. Section through the striate body, the thalamus and the internal capsule. The thickness of
thi slice is twice the standard of 2 mm (6/ 5 x )
92 Brain Slices
cornu anterius
2 Genu corpori callo i
3 Commi ura anterior
4 Pedunculus thalami anterio r
5 Columna fornici
6 Ventriculus tertiu 27 Nucleus ventral.i posterolateralis
7 Lamina medullaris lateralis 17 Caput nuclei caudati 28 Corpus geniculatum lateraJe
8 Lamina medullaris medialis 18 Putamen 29 Corpus gen iculatum mediale
9 Capsula interna, crus posterius 19 Sulcus circularis insulae 30 Area pretectaJis
10 Tractus mamillothalamicus 20 Gyri breves insulae 31 Colliculus superior
11 Capsula interna, pars retrolentiformis 21 Sulcus centralis insu.lae 32 Nuclei pulvinares
12 Commissura posterior 22 Gyrus longus insulae 33 Cauda nuclei caudati
13 Brachium colliculi superioris 23 Globus paUidus, pars la teralis 34 Hippocampus
14 Radiatio optica 24 Globus paUidus, pars medialis 35 Gyrus parahippocampalis
15 Stria terminalis 25 Zona incerta 36 Gy'rus occipitotemporalis medialis
16 Ventriculus lateralis, cornu inferius 26 Nucleus reticularis thalami 37 Lobus anterior cere belli
Fig. 86. Section through the striate body, the anterior commissure and the superior colliculus (6/5 x)
(~
1 Lamina terminalis 24 Nervus oculomotorius

2 Ventriculus tertius 25 Substantia nigra
3 Tractus opticus 13 Gyrus cinguli 26 Corpus amygdaloideum
4 Commissura anterior 14 Sulcus parolfactorius anterior 27 Cornu ammonis
5 Capsula interna, pars sublentiformis 15 Area subcallosa 28 Limbus Giacomini
6 Stria terminal is 16 Insula 29 Gyrus dentatus
7 Ventriculus lateralis, cornu inferius 17 Claustrum 30 Gyrus parahippocampalis
8 Fimbria hippocampi 18 Putamen 31 Sulcus collateralis
9 Pedunculus cerebri 19 Nucleus accumbens 32 Gyrus occipitotemporalis
10 Decussatio pedunculorum cerebellarium 20 Substantia perforata anterior lateralis
superiorum 21 Gyrus diagonalis 33 Lobus anterior cerebelli
11 Pedunculus cerebellaris superior 22 Hypothalamus 34 Fissura prima
12 Aqueductus cerebri 23 Corpus mamillare 35 Lobus posterior cerebelli
Fig. 87. Section through the mamillary body and the cerebral peduncle (6/5 x)
Part IV Microscopical Sections
Coronal Sections Through the Basal Part

of the Prosencephalon
Transverse Sections Through the Brain

Stem and Spinal Cord
96 Microscopical Sections
89 9091 92 93 94 95
Fig. 88. Key diagrams showing level and plane of the sections illustrated in Figures 89- 95. Above:
medial view; below: deep structures exposed from the same side
(l
o..,
o
::s
!::.
\/'J
(1)
1 Gyrus cinguli ~
2 Indusium griseum 0'
3 Corpus nuclei caudati ::srJJ
4 Pontes grisei >-l
caudatolenticulares P'
5 Septum pellucidum a
~
6 Nuclei septi (JQ
P'
7 Putamen
~.:
.' .. 8 Claustrum ~
.. :: .....
9 Insula t:t)
10 Nucleus gyri diagonalis rJJ
" ' ; '~i~l! } 11 Nucleus accumbens '"
!::.
:: '~:i::: : , :;: :;:,~W;:{ 12 Area subcallosa 'i:I
~!:. ' :~ . 13 Substantia innominata ..,....
:'. 14 Nucleus olfactorius a nterior
'o"
'.:..=:::.~:...~ ....,
.... .15 Cortex prepiriformis
.... 16 Cortex entorhinalis ~
;':;g\.. ...,'i:I
o
rJJ
(1)
~
't:l
P'
fl;,?;:~, /"""
' ,' 17 Cingulum
'0""
::s
18 Stria longitudinalis lateralis
19 Stria longitudinalis medialis
superior
22 Capsula externa
23 Capsula extrema
24 Fasciculus uncinatus cerebri
25 Bandeletta diagonalis
26 Stria olfactoria medialis
27 Stria olfactoria lateralis
Fig. 89. Section through the septal area (5/2 x)
\0
-.J
\0
00
1 Nuclei septi
4 Substantia innominata
5 Nucleus preopticus lateralis
6 Nucleus preopticus medialis
....
7 Nucleus gyri diagonalis
8 Lamina terminalis
." " ,
9 Nucleus supraopticus
. :~. 10 Recessus opticus
!{:; 11 Gyrus semilunaris
,";
12 Nucleus anterior )
13 Nucleus corticalis C
. orpus
, ,' 14 Nucleus .basahs
': amygd a-
.~ . : accessorIus . loideum
, 15 Nucleus basalIs
",
•2 '" • 16 Nucleus lateralis
,~
: .
~ j~h , . ,"
•, 1 • •\ 3
JI ' ~ • ~ ..
' 0' " 17 Radiatio corporis callosi
... . 18 Truncus corporis callosi
:::. ::,,:.. 19 Corona radiata
... . 20 Capsula interna, crus anterius
':':;~':~:::
:;
21 Fibrae caudatopallidales
22 Pedunculus thalami anterior
23 Lamina medullaris lateralis
24 Lamina medullaris medialis
25 Fasciculus lenticularis
26 Columna fornicis
28 Stria tenninalis
~
inferior n'
30 Bandeletta diagonalis o
rJl
31 Stria olfactoria lateralis (')
32 Fasciculus uncinatus cerebri o
"0
33 Chiasma opticum n'
E:.
Fig. 90. Section through the anterior commissure and the optic chiasm (5/2 X) CIl
o
(')
....
0'
::s
rJl
(1
0
....,
0
1 Cavum septi pellucidi
:::
e:-
2 Nucleus ventralis anterior if)
(j)
3 Foramen interventriculare (")
.....
4 Claustrum 0
0:::
5 Nucleus paraventricularis
6 Area lateralis hypothalami '">-3
7 Nucleus anterior hypothalami i:l"
....,
8 Nucleus supraopticus 0
s::
(JQ
.... :. 9 Substantia innominata
i:l"
10 Bed nucleus of stria terminalis .....
::•.
:\:. ,
,.,/' ·':j'i~j1"§l(
,~:;~ .~: ~: : . ' 11 Gyrus semilunaris i:l"
(j)
12 N"","" m,di,li, t:C

13 Nucleus corticalis C ~
o orpus
14 Nucleus basahs
accessonus
o
15 Nucleus basalts
0
I d
amyg a-
loideum
'"e:-
'"t:I
~
....,
.....
16 Nucleus lateralis 0
17 Gyrus ambiens
....,
.....
18 Gyrus parahippocampalis i:l"
(j)
'"t:I
....,
0
(j)
19 Fasciculus occipitofrontalis '"
:::
(")
superior (j)
20 Corpus fornicis '"0
i:l"
21 Stria terminalis ~
22 Fibrae caudatopallidales 0
:::
24 Columna fornicis
.: ::.~.:~~:: ' ... 26 Lamina medullaris lateralis
/t~:i? 27 Lamina medullaris medialis
28 Pedunculus thalami inferior
inferior
.... 31 Ansa lenticularis
:. : .' .:,.::"
32 Ansa peduncularis
"':.,'
33 Fibrae amygdalofugales
34 Tractus opticus
Fig. 91 Section through the interventricular foramen, the hypothalamus and the amygdaloid body (5/2 x)
'-0
'-0
......
0
0
1 Indusium griseum
3 Nucleus anterior thalami
4 Nucleus reticularis thalami
5 Nucleus ventralis anterior
6 Putamen
9 Nucleus posterior hypothalami
10 Nucleus ventromedialis
11 Area lateralis hypothalami
12 Nuclei tuberales
13 Nucleus infundibularis
15 Nucleus centralis
16 Nucleus basalis
accessorius ~ Corpus
17 Nucleus medialis amygda-
18 Nucleus corticalis loideum
19 Nucleus basalis
20 Nucleus lateralis
21 Cornu ammonis
22 Cingulum
24 Stria longitudinalis medialis
25 Lamina medullaris externa
.,'.:.:,':
26 Lamina medullaris interna
.' 27 Stria medullaris thalami
28 Capsula interna.
crus posterius
29 Capsula externa
30 Capsula extrema
32 Lamina medullaris medialis (S
~.
....
33 Fasciculus thalamicus 0
en
()
34 Fasciculus lenticularis 0
35 Columna fornicis "0
(S.
36 Ansa lenticularis
37 Tractus opticus
e:-
Fig. 92. Section through the rostral part of the thalamus, the amygdaloid body and the rostral pole of the hippocampus VJ
38 Commissura anterior C1l
()
(5/2 x) .....
o·
;:l
en
(J
0
....
0
::s
e:.
2 Nucleus medialis thalami U'J
('I)
3 Nucleus ventralis lateralis (")
..,.
4 Nucleus reticularis thalami O·
5 Nuclei intralaminares ::sen
6 Nuclei mediani thalami ...,
7 Zona incerta ::s-
....
.~ .~ , 8 Nucleus posterior hypo- 0
t:
..... :: (JQ
thalami
..'. 9 Area lateralis hypothalami
::s-
'?'::;. '. ..,.
.' 10 Nucleus subthalamicus ::s-
.:' . , ('I)
11 Corpus mamillare, nucleus 0;,

lateralis Il>
en
12 Corpus mamillare, nucleus e:.
medialis 'i:I
Il>
13 Substantia nigra ;:;.
14 Corpus amygdaloideum.
': : ~:?~,0Ni;,:.;: ~:'~:'im,':~::Zi~ nucleus basalis accessori us
0.....,
15 Pedunculus nuclei lentiformis S-

('I)
...:::.:.::.<:. . :;.:~~. 7 . 16 Cornu ammonis
, .. 17 Fascia dentata
....'i:I
0
'" en
." 18 Incisura unci ('I)
...~ ~~ ......,'" .~. .. ..: ·:~:.:j~.~;~i::\\~.:.:~: 9;: 8 .. ::s
(")
...
.... ..'. .. :... : 19 Sulcus hippocampi ('I)
.. 20 Subiculum '0
, . '. ·~;1.:4 21 Cortex entorhinalis ::s-
13 Il>
0-
::s
superior
23 Stria terminalis (1)
25 La mina medullaris medialis
26 Fasciculus thalamicus
27 Area tegmentalis H
29 Tractus mamillothalamicus
30 Fasciculus mamillaris princeps
32 Tractus optic us
35 Alveus hippocampi
Fig. 93. Section through the centre of the third ventricle, the mamillary body and the hippocampus (5/2 x)
....
0
....
0
Cauda nuclei caudati IV
2 Nucleus lateralis dorsalis
-
3 Nucleus lateralis posterior
5 Claustrum
6 Putamen
8 Nucleus ventralis
posterolateralis
9 Nucleus centromedianus
10 Nucleus parafascicularis
11 Nucleus ventralis
posteromedialis
12 Nucleus ventralis
'?'::' posteromedialis.
'. pars parvocellularis
.::~::. 13 Globus pallid us, pars lateralis
' ;'; .' ~ 4::_ ~ : ~
.. ... , 14 Zona incerta
7 15 Nucleus ru ber
"?:~, Wf' 16 Nucleus pregeniculatus
-:,::.',:/~:" 18 Substantia nigra
::,~::, . :'. 19 Cornu ammonis
20 Fascia dentata
,::().;:;!(~ ;~ 12 21 Subiculum
16 14':·:::: : : .. ' " ~~,,::>,,'::
:.~: : '::,
~#'. ~...., ' .•• : : : . ' 22 Lamina medullaris interna
.... ...,}. ..
23 Lamina medullaris externa
" 18 ,., .. 24 Capsula interna,
a., .•. crus posterius
....... ~~!'.... .... -- .. 25 Fasciculus longitudinalis
medialis
26 Pedunculus cerebellaris
superior
28 Capsula interna, ~
(i'
pars sublentiformis ...,
oVl
29 Decussatio pedunculorum ()
Fig. 94. Section through the thalamus and the caudal end of the putamen (5/2 X) cerebellarium superiorum o
'1:l
30 Pedunculus cerebri (i '
31 Tractus opticus :::..
32 Fimbria hippocampi C/:J
(1)
()
33 Alveus hippocampi 0,
o
::l
Vl
1 Gyrus cinguli
(')
2 Indusium griseum 0
3 Cauda nuclei caudati ....
0
4 Nucleus lateralis dorsalis ::s
5 Nucleus reticularis thalami e:..
IZl
6 Nucleus lateralis posterior (J)
()
7 Nucleus medialis thalami ......
8 Nuclei habenulae
o·
::s
[/l
10 Nucleus ventralis >-l
::s-
....
posterolateralis 0
11 Pontes grisei C
fJQ
· ...... caudatolenticulares ::s-
......
<:; 12 Nucleus interstitialis ::s-
(J)
;.:.:,::.: 13 Nucleus accessorius nervi
1:0
oculomotorii po
[/l
\~.~ '
.. : .. : ~'~'''''.": : 14 Substantia nigra e:..
15 Corpus genicula tum mediale ""0
,',
po
16 Corpus geniculatum laterale ....
.:. ......
17 Cornu ammonis
0
....,
:.'
18 Fascia dentata
......
t,: 19 Subiculum ::s-
(J)
.,: ,:/1:: 20 Cortex entorhinalis
""0
'~ :.:: ....
, :~ ;. 0
[/l
(J)
,'; 11
21 Cingulum ::s
()
~ , (J)
, 22 Stria longitudinalis lateralis
..' ..':.:' "0
23 Stria longitudinalis medialis ::s-
po
\', ,f 14 24 Stria terminalis (1) 0-
'," .. '.1 .J' . 25 Crus fornicis ::s
if , " .. "t If 26 Stria med ullaris thalami
.'\~ 27 Capsula interna,
.... P....... . pars retrolentiformis
28 Tractus habenulo-
interpeduncularis
29 Fasciculus longitudinalis
dorsalis
medialis
Fig. 95. Section through the medial and lateral geniculate bodies (5/2 x) 31 Pedunculus cerebellaris
superior
32 Radiatio optica
35 Decussatio pedunculorum
cerebellarium superiorum
36 Lemniscus medialis 0
w
-
~::::..[!::i~h~\-----I02
-4-.f::::L;~iP.l!!~---j---l03
~~--T---- l~
~!-H}-~---- 105
,*,A~~~::3r---- 106
-------.:~~~~f--!-;~HfI~---- 1 07
------~~\_f_-+-7fI~----- 108
------~~~~~------ 1~
A
Brain Stem 105
97 ________~~--~~~~~~~~~~~~~~N~II~
98----------~~~~~~~~~----~~=-
99 --------~~~~~~~~~--------
l00 ----------~~~~~~~~--------
101 ----------~~~~~~~~~-------
102------------~~~~--*+~7_~-----
103 ------~__:_'_~f3!:
104 -----,:+-=:--:;;---;-:>:'-:':-:-:-;:,....:.,.~~
105 --;""-'-'----,.-~.."..;,;~fJ'-_':4::~
106-{~::~·~~~-------r~~~~~T---------
107 ----------------H;'¥,~~~:;;;_-------
108 - - - -- -----";-'F-[-'
TR ~?INOTHAL
109 ----------~~~~--------
Fig. 96A, B. Key diagrams showing level and plane of the sections illustrated m Figures 97- 109.
Left: dorsal view ; right: lateral view
..... ~ -,-.
1 Nuclei pu lvinares
2 Colliculus superior
3 Area pretectalis
4 Corpus geniculatum mediale, pars dorsalis
5 Corpus geniculatum media Ie, pars ventrali
6 Corpus geniculatum laterale, laminae magnocellulares
7 Corpus geniculatum laterale, laminae parvocellulares
8 ucleus ventralis posteromedialis
9 Nucleus peripeduncularis
10 Griseum centrale mesencephali
11 Nucleus Darkschewitschi
12 Nucleus ruber, pars parvocellularis
14 Substantia nigra , pars reticuJata
15 Substantia nigra, pars compacta
16 Area tegmental is ventral is
17 Corpus mamillare, nucleus Jateralis
18 Corpus mamillare, nucleus medialis
Brain Stem 107
21
19 Brachium colliculi superioris

20 Radiatio optica
21 Commissura posterior
22 Tractus spinothalamicus
24 Tractus trigeminothalamicus dorsalis
25 Fasciculus longitudinalis dorsalis
26 Fasciculus longitudinalis medialis
27 Tractus tegmentalis centralis
28 Pedunculus cerebellaris superior
29 Tractus pallidoreticularis
30 Fibrae strionigrales
31 Tractus perietotemporopontinus
32 Tractus opticus
33 Tractus habenulointerpeduncularis
35 Tractus frontopontinus
36 Commissura supramamillaris
Fig. 97. Section through the posterior commissure, the medial and lateral geniculate bodies and the
mammillary body (7/1 x )
1 Stratum zonale )
2 Stratum griseum superficiale C II' I .
3 Stratum griseum medium 0 ICU us supenor
4 Stratum griscum profundum

6 Nucleus mesencephalicus nervi trigemini
7 Nucleus cuneiformis
8 Corpus geniculatum media Ie, pars dorsalis
9 Corpus geniculatum media Ie, pars ventralis
10 Nucleus interstitialis
11 Nucleus accessorius nervi oculomotorii
12 Nucleus nervi oculomo torii
15 Substantia nigra, pars reticulata
Brain Stem 109
16 Stratum zonale )
17 Stratum optic~~ Colliculus superior
18 Stratum lemmscl
19 Stratum album profundum
20 Commissura colliculi superioris
22 Tractus mesencephalicus nervi trigemini
23 Tractus tectospinalis
26 Brachium colliculi inferioris
34 Tractus parietotemporopontinus
Fig. 98. Section through the red nucleus and the medial geniculate body (7/1 x )
o.
"
., • "
J
: •
"
4 Nucleus paraiemniscaiis
6 Nucleus nervi oculomotori i
10 Nucleus ruber, pars magnocellularis
11 Nucleus interpeduncularis
Brain Stem 111
12 Commissura colliculi superioris

14 Fasciculus anterolateralis
24 Decussatio tegmentalis dorsalis
27 Decussatio tegmentalis ventralis
28 Pedunculus corporis mamiUaris
Fig. 99. Section through the superior colliculus and the oculomotor nuclei (7/1 x)
1 Nucleus intercollicularis
3 Nucleus paralemniscalis
4 Nucleus centralis colliculi inferioris
6 Nucleus nervi trochlearis
8 Nucleus tegmentalis pedunculopontinus, pars dissipata
11 Nuclei pontis
Brain Stem 113
12 Commissura colliculi inferioris

15 Tractus mesencephalic us nervi trigemini
21 Tractus tcgmentalis centralis
24 Decussatio pedunculorum cerebellarium superiorum
25 Pedunculus mamillaris
29 Fibrae pontocerebellares
Fig. 100. Section through the intercollicular area and the trochlear nucleus (7/1 x)
1 Nucleus intercollicularis
2 Colliculus inferior, nucleus centralis
3 Colliculus inferior, zona lateralis
5 Locus coeruleus
8 Corpus parabigeminum
9 Nucleus tegmentalis pedunculopontinus, pars compacta
10 Nucleus centralis superior
13 Nuclei pontis
Brain Stem 115

18 Lemniscus lateralis
19 Tractus tectopontinus
25 Decussatio pedunculorum cerebellarium superiorum
26 Fibrae corticotegmentales
27 Pedunculus mamillaris
Fig. 101. Section through the inferior colliculus and the decussation of the superior cerebellar peduncles
(7/1 x)
" .. .
'
..
....... . =-."
• , .. t • ••• .•
4 - , :; 5 •.
. ....
1 Nucleus mesencephalic us nervi trigemini

3 Locus coeruleus
4 Nucleus lemnisci lateralis
5 Nucleus parabrachialis lateralis
6 Nucleus parabrachialis medialis
7 Nucleus reticularis pontis oralis
9 Nucleus reticularis tegmenti pontis
10 Nuclei pontis
Brain Stem 117
"
II ervu trochleari
12 Decussatio nervorum trochJearium
14 Pedullculu cerebellaris superior
15 Lemniscus laterali
16 Fasciculus longitudinali medialis
17 Tractus tegmenlalis cenlralis
18 Fasciculus anterolaleralis
19 Tractus teclO pinali
21 Pedunculus cerebellaris superior, ramus dcscendells
22 Fibrae corlicotegmenlale
23 Fibrae pontoccrebellares
24 Pedunculus cerebellaris medius
25 Tractu pa rieto tcmporopontinus
26 TraClu pyramidalis
27 Tractu fronlopon tinus
Fig. 102. Section through the decussation of the trochlear nerves (7/1 x)
1 Nucleus dentatus
2 Nucleus emboliformis
3 Nucleus globosus
4 Nucleus fastigii
5 Nucleus vestibularis superior
6 Griseum centrale metencephali
8 Nucleus sensorius principalis nervi trigemini
9 Nucleus motorius nervi trigemini
10 Nucleus reticularis pontis caudalis
11 Formatio reticularis lateralis
12 Nucleus lemnisci lateralis
13 Nucleus raphes pontis
15 Nuclei pontis
Brain Stem 119
16 Commissura cerebelli
17 Decussatio fasciculorum uncinatorum
cerebelli
19 Pedunculus cerebellaris inferior
20 Fasciculus ovalis
21 Tractus vestibulomesencephalicus
22 Genu nervi facialis
26 Nervus trigeminus, radix motoria
27 Nervus trigeminus, radix sensoria
31 Decussatio tractuum trigemino-
thalamicorum ventralium
Fig. 103. Section through the principal sensory and motor nuclei of the trigeminal nerve (7/1 x)
1 Nucleus dentatus
3 Nucleus globosus
4 Nucleus fastigii
6 Nucleus vestibularis lateralis
7 Nucleus vestibularis medialis
8 Nucleus nervi abducentis
11 Nucleus nervi facialis
12 Nucleus gigantocellularis
13 Nucleus raphes magnus
14 Nucleus lateralis olivae superioris
15 Nucleus medialis olivae superioris
16 Nucleus corporis trapezoidei
17 Nuclei pontis
Brain Stem 121

19 Fibrae cerebellovestibulares
21 Tractus vestibulomesencephalicus + tractus
vestibulospinalis
22 Genu nervi facialis
24 Nervus abducens
26 Nervus facialis
27 Tractus spinalis nervi trigemini
28 Corpus trapezoideum
32 Lemniscus medialis + corpus trapezoideum
Fig. 104. Section through the abducens nucleus, the superior olive and the trapezoid body (7/1 x)
Nucleus dentatus
2 Nucleus globosus
5 Nucleus vestibularis inferior
6 Nucleus prepositus hypoglossi
7 Nucleus ovalis
9 Nucleus spinalis nervi trigemini
pars oralis
13 Oliva superior
15 Corpus pontobulbare
16 Nuclei pontis
Brain Stem 123

18 Pedunculus flocculi
20 Tractus vestibulomesencephalicus +
tractus vestibulospinalis
22 Fibrae nervi facialis
28 Nervus facialis
30 Nervus abducens
til''''''·'· .
! ,,,,,., .
. ,'Iio J.I···'IIIIII..~
~r·'.'·~
Fig. 105. Section through the vestibular nuclei and the motor nucleus of the facial nerve (7/1 x )
1 Nucleus cochlearis dorsalis 15 Striae acusticae dorsales

2 Nucleus vestibularis inferior 16 Striae medullares ventriculi quarti
3 Nucleus vestibularis medialis 17 Nervus vestibularis, ramus descendens
4 Nucleus prepositus hypoglossi 18 Pedunculus cerebellaris inferior
5 Nucleus solitarius 19 Tractus solitarius
6 Nucleus spinalis nervi trigemini, pars oralis 20 Fasciculus longitudinalis medialis
7 Formatio reticularis lateralis 21 Fibrae olivocerebellares
8 Nucleus gigantocellularis 22 Fasciculus anterolateralis
9 Nucleus raphes magnus 23 Nervus glossopharyngeus
10 Nucleus cochlearis ventralis 24 Tractus tegmental is centralis
11 Corpus pontobulbare 25 Lemniscus medialis
12 Nucleus ambiguus 26 Tractus pyramidalis
13 Nucleus olivaris inferior 27 Fibrae arcuatae externae
14 Nuclei arcuati
Fig. 106. Section through the cochlear nuclei (7/1 x)

Brain Stem 125
1 Nucleus cuneatus medialis 17 Pedunculus cerebellaris inferior

2 Nucleus cuneatus latera lis 18 Tractus solitarius
3 Nucleus vestibularis inferior 19 Tractus spinalis nervi trigemini
4 Nucleus solitarius 20 Nervus vagus
5 Nucleus dorsalis nervi vagi 21 Fibrae arcuatae internae
6 Nucleus intercalatus 22 Fasciculus longitudinalis medialis
7 Nucleus nervi hypoglossi 23 Fibrae olivocerebellares
8 Nucleus spinalis nervi trigemini , pars interpolaris 24 Fasciculus anterolateralis
9 Nucleus medullae oblongatae centralis 25 Amiculum olivae
10 Nucleus funiculi anterioris 26 Nervus hypoglossus
11 Nucleus raphes obscurus 27 Lemniscus medialis
12 Nucleus ambiguus 28 Tractus pyramidalis
13 Nucleus funiculi lateralis
14 Nucleus olivaris accessorius dorsalis
15 Nucleus olivaris accessorius medialis
Fig. 107. Section through the middle part of the inferior olive (7/1 x)
22
1 Nucleus gracilis 17 Fasciculus gracilis

2 Nucleus cuneatus lateralis 18 Fasciculus cuneatus
3 Nucleus cuneatus medialis 19 Pedunculus cerebellaris inferior
4 Nucleus solitarius 20 Tractus solitarius
5 Nucleus dorsalis nervi vagi 21 Tractus spinalis nervi trigemini
6 Nucleus intercalatus 22 Nervus accessorius, radices craniales
7 Nucleus nervi hypoglossi 23 Fasciculus anterolateralis
8 Cellulae marginales } Nucleus spinalis 24 Fibrae arcuatae internae
9 Substantia gelatinosa nervi trigemini, 25 Fasciculus longitudinalis medialis
10 Nucleus proprius pars caudalis 26 Decussation of fibrae arcuatae internae
11 Nucleus ambiguus 27 Lemniscus medialis
12 Nucleus funiculi lateralis 28 Nervus hypoglossus
13 Nucleus funiculi anterioris 29 Amiculum olivae
14 Nucleus olivaris inferior 30 Fibrae arcuatae externae
15 Nucleus olivaris accessorius medialis 31 Tractus pyramidalis
16 Nuclei arcuati
Fig. 108. Section through the dorsal column nuclei (7/1 x)

Brain Stem 127
1 Nucleus gracilis 9 Fasciculus gracilis

2 Cellulae marginales } Nucleus spinalis 10 Fasciculus cuneatus
3 Substantia gelatinosa nervi trigemini, 11 Tractus spinalis nervi trigemini
4 Nucleus proprius pars caudalis 12 Nervus accessorius, radices craniales
5 Nucleus cuneatus medialis 13 Tractus spinocerebellaris posterior
6 Nucleus retroambiguus 14 Tractus spinocerebellaris anterior
7 Nucleus medullae oblongatae centralis 15 Fasciculus anterolateralis
8 Nucleus supraspinalis 16 Decussatio pyramidum
18 Nervus spinalis cervicalis I, radix ventralis
Fig. 109. Section through the pyramidal decussation (7/1 x)

4 Tractus tectospinal is
5 Nucleus + tractus mesencephalicus nervi
trigemini .
8 Corpus geniculatum mediale
11 ucleus accessorius nervi oculomotorii
12 Nucleus nervi oculomotorii
14 Tractus tegmental is cent ralis
15 ucleus ru ber
17 Tractus pallidoreticularis
20 Decussatio tegmentalis dorsalis
21 Decussatio tegmentalis ventralis
23 Tractus rubrospinalis
24 Substantia nigra
26 Tractus opticus
Fig. 110. Diagrammatic section through the superior colliculus, showing the course and direction of
fibre tracts. Explanatory diagram to Figures 97- 99
ascending tracts sensory tracts
descending tracts motor tracts
Brain Stem 129
1 Commissura colliculi inferioris 10 Colliculus inferior

2 Decussatio nervorum trochlearium 11 Brachium colliculi inferioris
3 Nervus trochlearis 12 Nucleus + tractus mesencephalic us nervi
4 Tractus tectopontinus trigemini
5 Fasciculus longitudinalis medialis 13 Lemniscus lateralis
6 Tractus tegmentalis central is 14 Nucleus nervi trochlearis
7 Pedunculus cerebellaris superior 15 Tractus trigeminothalamicus dorsalis
8 Decussatio pedunculorum cerebellarium 16 Fasciculus anterolateralis
superiorum 17 Tractus pallidoreticularis
9 Nucleus interpeduncularis 18 Lemniscus medialis
19 Pedunculus cerebellaris superior, ramus
descendens
Fig. 111. Diagrammatic section through the inferior colliculus, showing the course and direction of
fibre tracts. Explanatory diagram to Figure 101. Symbols as in Figure 110.
13
1 Nucleus dentatus 16 Tractus corticovestibularis

2 Nucleus emboliforrnis 17 Fasciculus uncinatus cere belli
3 Nucleus globosus 18 Pedunculus cerebellaris inferior
4 Nucleus fastigii 19 Tractus spinocerebellaris anterior
5 Pedunculus cerebellaris superior 20 Nucleus reticularis tegmenti pontis +
6 Tractus fastigiobulbaris nucleus raphes pontis
7 Nucleus mesencephalicus nervi trigemini 21 Lemniscus medialis
8 Nuclei vestibulares 22 Lemniscus lateralis
9 Nucleus sensorius principalis nervi trigemini 23 Fasciculus anterolateralis
10 Nucleus motorius nervi trigemini 24 Tractus corticobulbaris
11 Fasciculus longitudinalis medialis 25 Radix motoria nervi trigemini, fibrae
12 Tractus tegmentalis centralis motoriae
13 Radix sensoria nervi trigemini 26 Tractus parietotemporopontinus
14 Radix motoria nervi trigemini 27 Tractus frontopontinus
(fibrae proprioceptivae) 28 Tractus pyramidalis
15 Nuclei pontis
Fig. 112. Diagrammatic section through the cerebellar peduncles, showing the course and direction
of fibre tracts. Explanatory diagram to Figure 103. Symbols as in Figure 110
Brain Stem 131
1 Nucleus cochlearis dorsalis 14 Pedunculus cerebellaris inferior

2 Striae acusticae dorsales 15 Nucleus nervi abducentis
3 Nuclei vestibulares 16 Fasciculus longitudinalis medialis
4 Nucleus prepositus hypoglossi 17 Nucleus nervi facialis
5 Nucleus solitarius 18 Lemniscus lateralis
6 Tractus + nucleus spinalis nervi trigemini 19 Tractus spinocerebellaris anterior
7 Nucleus cochlearis ventralis 20 Fasciculus anterolateral is
8 Nervus vestibulocochlearis 21 Lemniscus medialis
9 Nervus intermedius 22 Nervus facialis
10 Oliva superior 23 Tractus pyramidalis
11 Tractus tegmentalis centralis 24 Nervus abducens
13 Nuclei pontis
Fig. 113. Diagrammatic section through the cochlear nuclei and the trapezoid body, showing the
course and direction of fibre tracts. Explanatory diagram to Figures 104-106. Symbols as in Figure 110
1 Nucleus gracilis 14 Pedunculus cerebellaris inferior

2 Nucleus cuneatus medialis 15 Nucleus cuneatus lateralis
3 Nucleus solitarius 16 Nucleus dorsalis nervi vagi
4 Tractus solitarius 17 Nucleus nervi hypoglossi
5 Tractus + nucleus spinalis nervi trigemini 18 Tractus spinocerebellaris posterior
6 Fibrae arcuatae internae 19 Tractus reticulocerebellaris
7 Fasciculus longitudinalis medialis 20 Tractus reticulospinalis
8 Nervus vagus, fibrae sensoriae 21 Nucleus ambiguus
9 Tractus tegmentalis centralis 22 Nervus vagus, fibrae motoriae
10 Amiculum olivae 23 Tractus spinocerebellaris anterior
11 Nucleus olivaris inferior 24 Fasciculus anterolateralis
12 Lemniscus medialis 25 Tractus pyramidalis
13 Tractus olivocerebellaris 26 Nervus hypoglossus
Fig. 114. Diagrammatic section through the medulla oblongata, showing the course and direction
of fibre tracts. Explanatory diagram to Figures 107 and 108. Symbols as in Figure 110
Brain Stem 133
10
1 Fasciculus gracilis 9 Tractus spinocerebellaris posterior

2 Fasciculus cuneatus 10 Radix spinalis nervi accessorii
3 Tractus spinalis nervi trigemini 11 Tractus spinocerebellaris anterior
4 Nucleus spinalis nervi trigemini 12 Fasciculus anterolateralis
5 Tractus pyramidalis lateralis 13 Nucleus retroambiguus
6 Formatio reticularis 14 Nucleus supraspinalis
7 Nucleus supraspinalis 15 Fasciculus longitudinalis medialis
8 Fasciculus longitudinalis medialis 16 Tractus pyramidalis
17 Radix ventralis nervi spinalis cervicalis 1
Fig. 115. Diagrammatic section through the pyramidal decussation, showing the course and direction
of fibre tracts. Explanatory diagram to Figure 109. Symbols as in Figure 110
t Cellulae marginales 7 Fasciculus gracilis

2 Substantia gelatinosa 8 Fasciculus cuneatus
3 Nucleus proprius 9 Fasciculus dorsolateralis
4 Processus reticularis 10 Radix spinalis nervi accessorii
5 Substantia intermedia 11 Tractus spinocerebellaris posterior
6 Cellulae motoriae 12 Tractus pyramidalis la teralis
13 Canalis cen tralis
14 Commissura alba
15 Tractus spinocerebellaris anterior
18 Tractus pyramidalis anterior
19 Radix ventralis
Fig. 116. Section through the first cervical segment (9/ 1 x)

Spinal Cord 135
1 Cellulae marginales 8 Radix dorsalis

2 Substantia gelatinosa 9 Fasciculus gracilis
3 Nucleus proprius 10 Fasciculus cuneatus
4 Processus reticularis 11 Fasciculus dorsolateralis
5 Substantia intermedia 12 Tractus spinocerebellaris posterior
6 Cellulae motoriae laterales 13 Tractus pyramidalis lateralis
7 Cellulae motoriae mediales 14 Tractus spinocerebellaris anterior
18 Radix ventralis
Fig. 117. Section through the transition of fifth to sixth cervical segment (9/1 x)
1 Cellulae marginales 8 Fasciculus gracilis

2 Substantia gelatinosa 9 Fasciculus cuneatus
4 Nucleus intermediolateral is 11 Tractus spinocerebellaris posterior
5 Nucleus thoracicus 12 Tractus pyramidalis lateralis
6 Substantia intermedia 13 Tractus spinocerebellaris anterior
7 Cellulae motoriae 14 Fasciculus anterolateral is
15 Canalis centralis
16 Commissura alba
17 Radix ventralis
Fig. 118. Section through the fifth thoracic segment (9/1 x)

Spinal Cord 137
14
1 Cellulae marginales 8 Funiculus posterior

2 Substantia gelatinosa 9 Radix dorsalis
4 Processus reticularis 11 Funiculus posterolateralis
5 Substantia intermedia 12 Funiculus anterolateralis
6 Nucleus cornucommissuralis 13 Funiculus anterior
7. Cellulae motoriae laterales 14 Radix ventralis
Fig. 119. Section through the fifth lumbar segment (9/1 x)

1 Cellulae marginales 9 Radix dorsalis

2 Substantia gelatinosa 10 Funiculus posterior
4 Substantia intermedia 12 Funiculus posterolateralis
5 Cellulae motoriae laterales 13 Funiculus anterolateralis
6 Commissura grisea posterior 14 Commissura alba
7 Commissura grisea anterior 15 Funiculus anterior
8 Cellulae motoriae mediales 16 Radix ventralis
Fig. 120. Section through the transition of fourth to fifth sacral segment (9/ 1 x)
Spinal Cord 139
1 Sulcus medianus posterior

2 Radix dorsalis
3 Funiculus posterior 16 Fasciculus gracilis 29 Cellulae motoriae mediales
4 Septum medianum posterius 17 Fasciculus cuneatus 30 Tractus pyramidalis anterior
5 Cornu posterius 18 Fasciculus dorsolateralis 31 Fasciculus anterolateralis:
6 Funiculus posterolateralis} Funiculus 19 Tractus spinocerebellaris posterior Tractus spinotectalis
7 Funiculus anterolateralis lateralis 20 Tractus pyramidalis lateralis Tractus spinothaJamicus
8 Cornu anterius 21 Cellulae margin ales Tractus spinoanularis
9 Commissura grisea posterior 22 Substantia gelatinosa Tractus spino-olivaris
10 Canalis centralis 23 Nucleus proprius 32 Fasciculus longitudinalis medialis :
11 Commissura grisea anterior 24 Processus reticularis Tractus vestibulospinalis medialis
12 Commissura alba 25 Tractus spinocerebellaris anterior Tractus vestibulospinalis lateralis
13 Funiculus anterior 26 Fasciculi proprii Tractus reticulospinaJis
14 Fissura mediana anterior 27 Cellulae motoriae laterales Tractus tectospinalis
15 Radix ventralis 28 Substantia intermedia Tractus interstitiospinalis
Fig. 121. The subdivision of the white and grey matter in the spinal cord
Fig. 122: The grey matter of the spinal cord, subdivided according to Rexed [1118], at the level of
the sixth cervical segment
3 2
4 eI!!~~!!!31
5 ---'-'-L=--""
cervical
t Funiculus posterior
2 Fasciculus dorsolateralis
thoracic 3 Tractus spinocerebellaris posterior
4 A-fibres of dorsal root
5 C-fibre of dorsal root
7 Tractus spinoreticularis
9 Nucleus thoracicus
10 • Bordercell '
lumbar
7 7
Fig. 123. The origin and localisation of the ascending fibre tracts at three levels of the spinal cord
Spinal Cord 141
4 1 Fasciculus longitudinalis
medialis dexter
medialis sinister
5 Decussatio tegrnentalis ventralis
(mesencephali)
6 Decussatio tegmentalis
metencephali
7 Decussatio pyramidum
8 Tractus pyramidalis lateralis
10 A-fibres of dorsal root
11 Nucleus propri us
12 Substantia intermedia
13 Cellulae motoriae laterales
10~'I-;;;;;"""I...:sb': 14 Cellulae motoriae mediales
15 Radix ventralis
Fig. 124. Somatic reflex paths and descending supraspinal paths in the spinal cord; cervical level
1 Visceral afferent fibre

2 Nucleus propius
3 Nucleus intermediolateralis
5 Visceral efferent fibre
5
Fig. 125. Visceral reflex paths in the spinal cord. Thoracic level
Part V Functional Systems
Cranial Nerve Nuclei in the Brain Stem
General Sensory Systems and Taste

Special Sensory Systems
Ascending Reticular Systems
Cerebellum
Thalamocortical and Corticothalamic
Connections
Motor Systems
Descending Reticular Systems
Olfactory and Limbic Systems
Long Association and Commissural
Connections
Cranial Nerve Nuclei in the Brain Stem
The truncus cerebri or brain stem harbours addition nerve fibres related to special struc-
the centres of origin and termination of ten tures that occur in the head region. These
(III-XII) of the twelve cranial nerves fibres can be classed in the following three
(Fig. 126). At first sight the arrangement of additional categories:
these cranial nerve nuclei does not show a
5. Special somatic afferent fibres, which are
definite pattern; however, the classical inves-
associated with the receptors in the retina
tigations of Gaskell [386, 387], Herrick [513]
and in the cochlea.
and many others (for a review, see Nieuwen-
6. Special visceral afferent fibres, which sup-
huys [995]) have revealed that these centres
ply the visceral sense organs, i.e., the or-
form part of functional zones, each of which
gans of taste and smell.
is specifically related to one of the fibre cate-
7. Special visceral efferent fibres, which in-
gories of which the cranial nerves are com-
nervate muscles derived from the mesen-
posed. Before elucidating this zonal pattern
chyme of the visceral, branchial arches.
the fibre categories or nerve components of
peripheral nerves in general deserve some Whereas a spinal nerve usually contains
comment. fibres of all four of the' general' categories,
A single nerve fibre in a spinal or cranial there are wide variations between the cranial
nerve may be afferent or efferent and be con- nerves as regards the types of fibres which
cerned with the innervation of either somatic they carry. Some of them have fibres of only
or visceral structures. Combination of these one type, but in others fibres of two or more
two subdivisions yields the following four categories are present.
categories of peripheral nerve fibres: Returning now to the brain stem, the
organisational pattern discovered by Gaskell
1. General somatic afferent fibres, which
and Herrick and their followers is that the
transmit impulses from the skin, skeletal
cranial nerve nuclei are essentially arranged
muscles, joints and ligaments.
in seven longitudinal zones and that each of
2. General visceral afferent fibres, which
these columns is specifically related to fibres
convey impulses from receptors in visceral
of one of the categories mentioned above.
organs and blood vessels centrally.
Thus, as a cranial nerve composed of fibres
3. General visceral efferent fibres, which
of more than one type enters the brain, the
supply the smooth musculature of internal
fibres of its constituent types sort themselves
organs, the cardiac muscle and glands.
out and pass to 'their own' specific zone.
4. General somatic efferent fibres, which
The zonal or columnar pattern displayed
innervate skeletal muscles derived from
by the cranial nerve nuclei in the brain stem
myotomes.
is diagrammatically represented in Figure
The denotation 'general' has been added 127. As in the spinal cord, the afferent centres
because fibres belonging to these four cate- are situated in the alar lamina, whereas the
gories occur in both spinal and cranial efferent centres are located in the basal la-
nerves. The cranial nerves may contain in mina. The sulcus limitans, which in the
Cranial Nerve Nuclei 145
embryonic neuraxis marks the boundary of the ninth, the tenth and the eleventh (cra-
between these two fundamental subdivisions, nial root) nerves. The spinal nucleus of the
is in the adult only recognisable over a short eleventh nerve, which is situated in the lateral
extent. It will be noted that most of the zones part of the base of the ventral horn of the
are only partly occupied by cranial nerve upper four cervical segments, also belongs to
nuclei. This may be related to the reduction this zone. As with the general somatic affer-
of some components of some nerves during ent nuclei, the cell masses of the branchio-
foetal development. The various zones and motor zone have migrated away from their
their constituent primary afferent or efferent original periventricular position.
centres will now be briefly reviewed, passing The general somatic efferent (GSE) zone,
from lateral to medial. finally, may be considered a rostral continua-
The special somatic afferent (SSA) zone tion of the anterior horn of the spinal cord.
contains the nuclei of termination of the It comprises the nuclei of origin of the
cochlear and vestibular division of the eighth twelfth, sixth, fourth and third nerves. All
nerve. four of these nuclei are located near the medi-
The general somatic afferent (GSA) zone an plane of the brain stem.
includes the three sensory nuclei of the tri-
geminal nerve, i.e., the mesencephalic, the
princeps or chief and the spinal nuclei. The
latter nucleus, which also receives some fibres
from the seventh, ninth and tenth nerves, is
caudally continuous with the apical part of
the dorsal horn of the spinal cord. The senso-
ry trigeminal nuclei have shifted ventrolater-
ally during development, hence the princeps
nucleus and the rostral part of the spinal
nucleus lie in the adult ventral rather than
medial to the vestibular nuclei.
The special visceral afferent (SVA) and
general visceral afferent (GVA) zones are in
the adult brain represented by a single cell
mass, which receives the corresponding com-
ponents of the seventh, ninth and tenth
nerve. The latter unite in a well-defined fibre
system, the tractus solitarius.
The general visceral efferent (GVE) zone
contains four nuclei, the nucleus dorsalis of
the tenth nerve, the nuclei salivatorii inferior
and superior of the ninth and seventh nerve,
respectively, and the accessory nucleus of the
third nerve. These nuclei represent together
the cranial division of the parasympathetic
system. They give rise to preganglionic fibres
that terminate in various autonomic ganglia.
The special visceral efferent (SVE) or
branchiomotor zone contains the motor nu-
clei of the fifth and seventh nerve as well
as the nucleus ambiguus, which gives rise to
fibres that pass peripherally as components
146 Functional Systems
1 Tractus mesencephalicus nervi trigemini Vm Radix motoria nervi trigemini

2 Nucleus mesencephalicus nervi trigemini 9 Nucleus accessorius nervi oculomotorii
3 Nucleus sensorius principalis nervi trigemini 10 Nucleus nervi oculomotorii
4 Tractus spinalis nervi trigemini 11 Nucleus nervi trochlearis
5 Nucleus spinalis nervi trigemini 12 Nucleus motorius nervi trigemini
6 Nuclei vestibulares 13 Genu nervi facialis
7 Nuclei cochlea res 14 Nucleus nervi abducentis
8 Nucleus solitarius j 5 Nucleus nervi faciali.s
16 Nuclei salivatorii
17 Nucleus ambiguus
18 Nucleus dorsalis nervi vagi
19 Nucleus nervi hygoglossi
20 Nucleus radicis spinalis nervi accessorii
Fig. 126. The cranial nerve nuclei as viewed from the dorsal side (5/3 x). Left : Sensory nuclei; right:
motor nuclei. Roman numerals indicate the corresponding cranial nerves
Cranial Nerve Nuclei 147
-nr-----+ nucleus accessorius

nerv i oculomotori i
mesencephalon
metencephalon
(pons)
myelencephalon
(medulla oblongata)
-n;-ji-- ___ -+-nuc leus ambiguus
H--t- - - --f nucleus rad icis spinal is

medulla spina lis nervi accessorii
lam ina alaris lamina basalis
Fig. 127. Position of the cranial nerve nuclei in longitudinal columns. The initials at the top indicate
the functional system to which each column belongs. The parts of the brain stem are indicated at
left
General Sensory Systems and Taste
Introduction is the first link in the epicritic pathway from

the face. It synapses in the principal sensory
Sensory pathways within the central nervous nucleus of the trigeminal nerve and decus-
system connect primary afferents with specif- sates at the level of the pons to join the me-
ic parts of the contralateral cerebral cortex. dial lemniscus in its course to the thalamus.
These so-called lemniscal systems synapse in Epicritic systems are somatotopically orga-
nuclei of the spinal cord and lower brain stem nized, e.g. lamination of primary afferents in
and in the sensory relay nuclei of the thala- the posterior funiculus and representation of
mus. Multisynaptic pathways through the re- the body in the ventral posterior nucleus of
ticular formation are arranged in parallel the thalamus and in the primary and second-
with the lemniscal systems. Descending path- ary somatic sensory cortex.
ways from the cerebral cortex, the central The protopathic pathways arise from the
grey matter, the reticular formation and the dorsal horn of the spinal grey matter and
raphe nuclei terminate in the main relay from the pars caudalis of the spinal nucleus
stations of the sensory projection systems. of the trigeminal nerve. Both these relay nu-
Protopathic systems, subserving pain and clei have the same laminated structure. They
temperature and yielding ungraded, diffuse receive thin primary afferents that conduct
impressions of an all-or-none character, can pain and temperature sensation and collater-
be distinguished from epicritic systems als from thicker primary afferents that inner-
concerned with the mediation of tactile and vate mechanoreceptors. The spinothalamic
kinesthetic information of a discriminative tract is the protopathic pathway for the trunk
type [164]. However, one should be cautious and limbs. Its fibres decussate within the cord
in attributing isolated functions to certain at the level of their origin from the dorsal
ascending pathways, which for all we know, horn and ascend in the anterolateral funicu-
may participate simultaneously in most lus together with the spinoreticular and ven-
sensory functions [1010]. tral spinocerebellar tracts. The lateral tri-
Sensation from the trunk and limbs and geminothalamic tract conveys pain and tem-
that from the face are mediated by two differ- perature sensation from the face. It decus-
ent sets of sensory pathways. The posterior sates in the caudal medulla, at the level of
funiculus-medial lemniscus pathway sub- its origin from the pars caudalis of the spinal
serves epicritic sensation from the trunk and trigeminal nucleus. Both the spinothalamic
limbs. Its primary afferents ascend in the pos- and the lateral trigeminothalamic tracts ter-
terior funiculus and synapse in the dorsal col- minate, together with the medial lemniscus,
umn nuclei. These nuclei are the origin of in the ventral posterior nucleus of the thala-
the medial lemniscus, which decussates in the mus. Their termination also includes adjoin-
caudal medulla and ascends, dorsal to the ing parts of the intralaminar nuclei, the pos-
pyramid and along the ventral border of the terior group and the ventral lateral nucleus,
tegmentum, to the thalamus. The sensory where they overlap with other afferent sys-
root (portio major) of the trigeminal nerve tems.
Most of the relay nuclei, in~luding the so- firmed Wall's [1472] observation of a physio-
matosensory cortex, give origin to descend- logical laminar arrangement of the input to
ing systems, which reciprocate the ascending the dorsal horn and the intermediate grey
connections. A descending inhibitory path- matter. The C- and AD-fibres, which inner-
way which selectively influences pain conduc- vate most of the receptors responding to nox-
tion in the dorsal horn and the pars caudalis ious stimuli (including the thermoreceptors
of the spinal trigeminal nucleus includes the and certain mechanoreceptors), terminate on
central grey matter and the raphe nuclei. marginal cells and in the superficial part of
Primary afferents in the facial, glossopha- the substantia gelatinosa [729, 786, 787, 788,
ryngeal and vagal nerves are the first link 1109]. A -fibres also terminate at the ventral
in the gustatory and general viscerosensory border of the dorsal horn. Collaterals of
pathways. These fibres descend in the medul- thicker myelinated (Af3 and AI') fibres, inner-
la oblongata in the solitary tract, which is vating touch receptors in the skin, terminate
medial to the spinal tract of the trigeminal in the deep layers of the substantia gelatino-
nerve, near the floor of the fourth ventricle. sa, the nucleus proprius and the adjoining
They terminate in the nucleus of the solitary intermediate grey matter. Visceral and so-
tract. Connections from this nucleus to the matic primary afferents also converge on
telencephalon are interrupted in the medial, cells in this region [125]. Joint afferents rami-
parvocellular part of the ventral posterior nu- fy in more ventral parts of the intermediate
cleus of the thalamus. Another pathway from grey matter. The collaterals of the large ca-
the solitary tract nucleus synapses in the libre (AIX) myelinated fibres which innervate
parabrachial nuclei and bypasses the thala- muscle spindles and Golgi tendon organs
mus to reach the gustatory cortex in the par- penetrate the ventral horn, where they termi-
ietal operculum and the insula. Telencephalic nate on motoneurons.
targets of the taste pathway include the later- Most of the neurons that are contacted
al nucleus of the amygdala. by primary afferents have long axons, which
enter the funicular white matter; short axon
cells are rare in the spinal grey matter [1214].
Primary Afferents The great majority of small neurons in the
and the Spinal Grey Matter substantia gelatinosa belong to an intrinsic
(Figs. 123 and 124) system of the dorsal horn. It caps the dorsal
horn as a thin cell layer, poor in myelin, lo-
Most primary afferents enter the spinal cord cated between the superficial marginal layer
through the dorsal roots. A certain number and the nucleus proprius. It contains the ter-
of unmyelinated fibres enter through the ven- minal arborizations of thin primary afferents,
tral roots [243], but their exact distribution which contact relay cells of the marginal
remains unknown at present. Dorsal root layer, and also the flame-shaped arborisa-
fibres bifurcate on entering the cord. They tions of thicker dorsal root fibres, which ter-
segregate into thin myelinated (AD) and un- minate on relay cells of the nucleus proprius.
myelinated (C) fibres, which enter the lateral The substantia gelatinosa is often considered
part of the dorsolateral fasciculus and cap to be a site of interaction between the thin
the dorsal horn, and thicker myelinated (AIX, fibres that conduct pain and temperature and
Af3 and AI') fibres, which course through the the larger cutaneous afferents from the dorsal
posterior funiculus. Collaterals of these fibres root [681, 753, 970, 1003, 1473]. The micro-
enter the grey matter of the cord, where they circuitry of the substantia gelatinosa has not
terminate in the dorsal horn, the intermediate been completely resolved.
grey matter and the ventral horn.
Recent physiological and anatomical stu-
dies [115, 116, 178, 347] have generally con-
General Sensory Systems and Taste 151
The Anterolateral System nucleus, the termination is somatotopically

(Figs. 128 and 129) organized. In primates and humans, the
fibres terminate in bursts or rod-like aggre-
The anterolateral funiculus contains some of gates, overlapping the projections from the
the main ascending fibre systems of the spinal dorsal column nuclei. Rostrally, the termina-
cord. The spinothalamic tract, the protopath- tion of the spinothalamic tract extends to the
ic pathway conducting pain and temperature junction with the ventral lateral nucleus,
from the spinal cord, originates from the cells which also receives proprioceptive, vestibular
of the marginal layer, from the nucleus pro- and cerebellar afferents. Spinothalamic fibres
prius of the dorsal horn and from the inter- terminate diffusely in the medial part of the
mediate grey matter (laminae I, IV, V, VI, posterior group, an ill-defined region of the
VII and VIII of Rexed [1118]). Most spino- thalamus located between the posterior ven-
thalamic fibres cross in the white commissure tral nucleus and the lateral nuclear group.
of the cord. Propriospinal and spinoreticular The spinothalamic tract also terminates in a
fibres arise from the same regions of the grey diffuse, non-somatotopic manner in some of
matter. However, these projections are bilat- the intralaminar thalamic nuclei, mainly the
eral and originate mainly in the intermediate central lateral nucleus [935]. The projection
grey matter. Fibres of the ventral (or anteri- to the intralaminar nuclei is partially bilater-
or) spinocerebellar tract occupy the periph- al: some of its fibres cross in the posterior
ery of the anterolateral funiculus. Their ori- commissure. Most of these intralaminar nu-
gin partially overlaps with that of the pro- clei also receive projections from the reticular
priospinal and spino reticular systems [116, formation, the raphe nuclei and the cerebel-
272,347,1078,1385,1516, 1517, 1518]. The lum, some of which subserve motor rather
ascending fibres in the anterolateral fascicu- than sensory functions. Hence, not all regions
lus display a roughly somatotopic organiza- that receive spinothalamic fibres are part of
tion. Fibres originating from lower levels of a pain-conducting system.
the cord are located lateral to fibres from Nociceptive units have been identified in
higher levels. the posterior ventral nucleus, the medial part
In the brain stem, the anterolateral fasci- of the posterior group, the nucleus subme-
culus is located lateral to the reticular forma- dius and the zona incerta [48, 263, 401]. The
tion. Spino reticular fibres terminate at differ- lateral thalamus is concerned with the discri-
ent levels in the medial reticular formation. minative aspects of pain, the medial thalamus
They are the first link in a multi synaptic with the motivational and arousal aspects of
pathway that has been postulated to ascend a painful stimulus. Spinothalamic projections
within the reticular formation and terminate to the medial thalamus arise from the inter-
mainly in the intralaminar nuclei of the thala- mediate grey matter, together with the spino-
mus. In addition, spinal fibres terminate in reticular fibres, and from the marginal layer
the central grey matter of the mesencephalon and nucleus proprius of the dorsal horn,
and in the deep layers of the mesencephalic together with the spinothalamic tract fibres
tectum. At the level of the superior colliculus, to the lateral thalamus [116, 263, 400, 401,
most fibres of the spinothalamic and lateral 1516].
trigeminothalamic tracts lie concentrated in
a small bundle at the dorsal tip of the medial
lemniscus.
The spinothalamic tract terminates in the
posterolateral ventral nucleus, posterior
group and certain intralaminar nuclei of the
thalamus [110, 111, 138, 263, 682, 830, 852,
884, 886, 887]. In the posterolateral ventral
1 Nuclei intralaminares thalami

2 Nucleus ventralis posterolateralis
3 Corpus genicula tum mediale
4 Formatio reticularis medialis
7 Radix dorsalis nervi spinalis
Fig. 128. The anterolateral fasciculus. The position of the ascending spinal tracts and related nuclei
in a dorsal view (5/3 x)
i
I
I
I,
I
7 I Gyrus po tcentrali
I 9
2 Tractus pyramidali
a;1'
3 uclci intralaminare thalami
4 ucleus ventralis po terol aterali s
5 Corpus genicula tum medi ale
6 Griseum centrale me encephali
11 Tractus pyramidalis laterali
12 Nucleus proprius
13 Fasciculus anterolaterali
15 Funiculus anterolaterali
'-----' 14
13 A C-libre
C A-fibre} Ra d IX" d
orsa I"IS nerVI"spma
" I"IS
L.......-...-J
15
Fig. 129. The neuronal connections of the spinoreticular tract (darkly shaded) and of the spinothalamic
tract (solid line)
The Medial Lemniscus System the cerebellum through the restiform body.
(Figs. 130 and 131) Efferents from the diffusely organized ventral
and rostral parts of the dorsal column nuclei
The first link in the epicritic conduction path- are more widely distributed than efferents
way from the cord comprises branches of from the cell clusters in the dorsal parts of
thick myelinated dorsal root fibres that as- the nuclei. Both regions project to the contra-
cend in the posterior funiculus, where they lateral thalamus, but the ventral and rostral
are somatotopically organized, fibres from parts also project to the cerebellum, the infe-
sacral and lumbar roots ascend medially in rior olive and back to the dorsal horn [112,
the gracile fascicle; those from cervical roots 135, 232, 395, 749].
ascend laterally in the cuneate fascicle. A From the gracile and the medial cuneate
small contingent of thoracic fibres takes an nuclei arise internal arcuate fibres. These
intermediate position (Fig. 123). The gracile cross the medial plane and ascend in the
and cuneate fascicles terminate in corre- medial lemniscus to terminate in the postero-
sponding dorsal column nuclei in the caudal lateral ventral nucleus, the posterior group,
end of the medulla oblongata. Along the the magnocellular part of the medial genicu-
course of the dorsal root fibres in the posteri- late body and the zona incerta [110,111,137].
or funiculus, and at their termination in the The projection of the dorsal column nuclei
dorsal column nuclei, the initial laminar pat- to the posterolateral ventral nucleus is orga-
tern becomes reorganized so that fibres, from nized in a "core-and-shell" fashion. Tactile
adjacent dorsal roots, that innervate the same projections from the cell clusters of the dorsal
cutaneous field are bundled together. As a column nuclei occupy the core of the nucleus
result of this convergence the overlap be- and proprioceptive afferents terminate in the
tween neighbouring dermatomes is elimi- shell. The fibres of the medial lemniscus ter-
nated, but the original laminar organization minate in a series of parallel laminae, which
of the posterior funiculus becomes indistinct extend anteroposteriorly, throughout the
[1503]. shell and the core of the nucleus. Each lamina
Posterior funiculus fibres innervating the represents a specific region of the body, but
distal parts of the limbs terminate somato- includes different sensory modalities along its
topically in cell clusters which occupy the anteroposterior axis [362, 636, 642, 1504].
dorsal parts of the gracile and medial cuneate
nuclei. In the ventral and rostral parts of
these nuclei the somatotopic organization is
The Somatosensory Cortex
much less precise. The posterior funiculus
contains an important system of intrinsic
fibres, originating from the nucleus proprius Different somatosensory areas have been dis-
of the dorsal horn. These intrinsic fibres ter- tinguished in the cerebral cortex. The first
minate in the ventral and rostral parts of the (Sl), which occupies the postcentral gyrus,
dorsal column nuclei [485, 1172]. Other consists of four cytoarchitectonic fields (3a,
fibres, which ascend in the posterolateral fun- 3b, 1 and 2 of Brodmann [177]), which extend
iculus, terminate bilaterally in the ventral and parallel to the central sulcus. The Sl cortex
rostral parts of the dorsal column nuclei, and can be divided into narrow, regionally specif-
also in a cell group (group Z [171]) located ic columns, which extend perpendicular to
at the rostral end of the gracile nucleus, the central sulcus through all four cytoarchi-
which serves as a spinothalamic relay nucleus tectonic fields. The cortical columns receive
for muscle afferents of the lower limb [13, their afferents from the thalamic cells located
422, 1172]. A similar proprioceptive relay for within the laminae of lemniscal fibres in the
the upper limb is located in the lateral cun- ventral posterior nucleus. The core of this
eate nucleus [139], which mainly projects to nucleus projects to fields 3b and 1, the shell
to fields 3a and 2. Each of the four cytoarchi- Small somatosensory components of the fa-
tectonic fields of S1 therefore contains a com- cial, glossopharyngeal and vagal nerves also
plete representation of the body surface. Pro- join the spinal trigeminal tract.
prioceptive afferents from the shell region The pars caudalis of the spinal trigeminal
terminate in the anterior field (3a) and the nucleus consists of a layer of marginal cells,
posterior field (2), which surround the central a substantia gelatinosa and a nucleus pro-
tactile fields (3b and 1). The latter receive prius, which are continuous with the same
their thalamic projections from the core of layers of the spinal dorsal horn. The A- and
the ventral posterior nucleus [362, 629, 635, C-fibres of the spinal trigeminal tract and
636,642, 898,959, 1316, 1504]. those of the spinal roots have essentially simi-
The ventral posterior nucleus also projects lar synaptic relations. As well, the afferent
to the second somatosensory area (S2), which connections of the pars caudalis and the dor-
is located in the parietal operculum. Nocicep- sal horn are essentially similar [189,380,528].
tive projections to the somatosensory cortex The epicritic and protopathic pathways
have not been documented and painful sensa- from the trigeminal nerve arise from the ros-
tions cannot be evoked by stimulation of the tral and caudal parts, respectively, of the tri-
somatosensory cortex in humans [1064]; see geminal nuclear complex. Sjoqvist's tracto-
however [36]. Nociceptive units are present tomy of the spinal tract at the level of the
at the level of the posterior ventral nucleus rostral border of the pars caudalis [730, 1257]
of the thalamus, and the projection of the therefore effectively abolishes pain and tem-
medial part of the posterior group to the re- perature sensibility in the ipsilateral face,
troinsular cortex has been implicated in the leaving the tactile sense intact. The proto-
transmission of pain in monkeys [190, 630], pathic pathway from the trigeminal nerve,
see also [1365]. which joins the spinothalamic tract as the lat-
eral trigeminothalamic tract, originates from
the pars caudalis of the spinal trigeminal nu-
cleus and crosses in the caudal medulla ob-
The Trigeminal System longata [1247]. The pars caudalis also gives
(Figs. 132 and 133) rise to an intranuclear ascending system,
which terminates in the principal sensory nu-
Somatosensory fibres of the trigeminal nerve cleus [189, 380, 585, 1049]. This system has
enter the pons in the sensory root (portio been implicated in pain conduction from the
major) and are distributed to the principal face, by behavioural experiments which
sensory nucleus and the spinal trigeminal nu- showed that a midsagittal cut at the level of
cleus. A-fibres terminate in the principal sen- the principal sensory nucleus is more effective
sory nucleus and in the different subdivisions in abolishing pain sensation in the face than
of the spinal trigeminal nucleus (pars oralis, a more caudal cut at the level of the pars
pars interpolaris and pars caudalis, [1027]). caudalis [457]. The lateral trigeminothalamic
Thin A-f> and C-fibres descend in the spinal tract ends in clusters of terminals in the con-
tract (which continues in the dorsolateral fas- tralateral posteromedial ventral nucleus of
ciculus of the dorsal horn) and terminate in the thalamus, in the adjoining part of the
the pars caudalis and upper cervical dorsal posterolateral ventral nucleus, in the border
horn [283]. Fibres of the mandibular division region of these nuclei with the ventral lateral
of the trigeminal nerve descend in the dorsal nucleus, in the posterior group, and bilateral-
part of the tract, whereas the ophthalmic di- ly in some of the intralaminar nuclei [110,
vision fibres occupy the ventral part. Fibres 189,221,380].
of the maxillary nerve occupy an intermedi- The ventral part of the principal sensory
ate position and descend less far caudally nucleus gives rise to the crossed epicritic
than the other two divisions [730, 1173]. pathway, which joins the medial lemniscus

3 Fibrae arcuatae internae
4 Nucleus cuneatus medialis
5 Nucleus gracilis
6 Fasciculus cuneatus
7 Fascicul us gracilis
9 Ganglion spinale
Fig. 130. The medial lemniscus. Position of tracts and nuclei in a dorsal view (5/3 x)
1 Gyrus postcentralis
11 4 Lemniscus medialis
5 Fibrae arcuatae internae
6 Nucleus gracilis
8 Fasciculus gracilis
9 Fasciculus cuneatus
11 Ganglion spinale
Fig. 131. The neuronal connections of the dorsal column nuclei and the medial lemniscus
V1 Nervus ophthalmicus
V2 Nervus maxillaris
VJ Nervus mandibularis
1 Nucleus ventralis posteromedialis
3 Ganglion trigeminale
4 Radix motoria nervi trigemini
5 Radix sensoria nervi trigemini
6 Nucleus mesencephalicus nervi
trigemini
7 Nucleus sensorius principalis nervi
trigemini 13 Nucleus ventralis posteromedialis
8 Nucleus spinalis nervi trigemini 14 Lemniscus trigeminalis
9 Tractus spinalis nervi trigemini 15 Lemniscus medialis
10 Substantia gelatinosa 16 Tractus spinothalamicus
11 Radix dorsalis nervi spinalis 17 Fasciculus tegmentalis ventralis
12 Nucleus proprius 18 Tractus trigeminothalamicus lateralis
Fig. 132. The central connections of the trigeminal nerve. Position of nerves, tracts and nuclei in
a dorsal view (5/3 x). Roman numerals indicate the corresponding cranial nerves
I Gyrus postcentralis
2 Fibrae corticonucleares
3 Nucleus ventralis posteromedialis
4 Lemniscus trigeminalis
8 Fibrae prOPriocePtivae} Radix motoria
9 Fibrae motoriae nervi trigemini
10 Ganglion trigeminale
11 Radix sensoria nervi trigemini
13 Nucleus sensorius principalis nervi
trigemini
15 Pars OraliS}
. terpo Ians
' Nucleus . .
16 Pars ID . . splDabs
. .
17 Pars caudalis nervi tngemlDl
18 Nucleus proprius
19 Substantia gelatinosa
Fig. 133. The neuronal connections of the trigeminal nerve

as the trigeminal lemniscus. The uncrossed, Descending Connections

dorsal trigeminothalamic tract arises from to Somatosensory Relay Nuclei
the dorsal part of the principal sensory nucle- (Fig. 134)
us [187, 865, 1262, 1382, 1539]. The trigemin-
allemniscus terminates in the posteromedial Descending fibres from the deep layers of
ventral nucleus, where it overlaps with the pyramidal cells in the first and second soma-
projection of the lateral trigeminothalamic tosensory areas (S1 and S2) terminate soma-
tract. The termination of the uncrossed tri- totopically in the ventral posterior nucleus.
geminothalamic tract in the cat differs from A more extensive area including the adjoin-
that of the crossed fibres and occupies the ing part of the motor cortex in the precentral
lateral part of the parvocellular portion of gyrus (area 4), the premotor cortex (area 6)
the posteromedial ventral nucleus, lateral to and the sensory association area (area 5) in
the gustatory subdivision of the nucleus the superior parietal lobule contributes fibres
[1538]. that join the pyramidal tract and terminate
Investigations into the central nervous sys- in the contralateral sensory nuclei of the tri-
tem of rodents have not been systematically geminal nerve, the dorsal column nuclei and
considered in this chapter. An exception the dorsal horn [11, 13, 221, 259, 671, 738,
should be made for the discovery that in mice 739, 740, 1366, 1488, 1489]. Descending
there is a discrete projection of the facial fibres from the cortex preferentially termi-
vibrissae to distinct cell aggregates in the nate outside the clusters or laminae which
fourth layer of the somatosensory cortex contain the main sensory relay cells of the
[1527]. This has promulgated a series of ventral posterior nucleus of the thalamus, the
investigations which have improved our principal trigeminal sensory nucleus and the
understanding of information processing in dorsal column nuclei. In the pars caudalis of
a sensory system [11, 100,332,484,685]. the trigeminal spinal tract nucleus and the
Unlike to all other primary afferents, the dorsal horn, the fibres terminate both in the
proprioceptive muscle-spindle afferents, deep layers and in the marginal zone or in
which enter the brain stem in the trigeminal the substantia gelatinosa [232].
nerve, arise from cells located within the cen- Other descending systems, which originate
tral nervous system. These cells constitute the from the raphe nuclei and the reticular for-
mesencephalic nucleus of the trigeminal mation of the pons and medulla oblongata,
nerve, located alongside the central grey mat- terminate on the cells that give rise to the
ter of the mesencephalon. The axons of these spinothalamic and the lateral trigeminotha-
cells descend in the mesencephalic tract of lamic tracts in the spinal grey matter and in
the trigeminal nerve, and have collaterals the pars caudalis of the trigeminal spinal tract
which terminate on the cells of the trigeminal nucleus. Fibres from raphe magnus nucleus,
motor nucleus. The main axons of the mesen- some of which are serotonergic, and from the
cephalic root, together with the axons of the adjoining reticular formation terminate bila-
motoneurons, constitute the portio minor of terally in the marginal layer, the substantia
the trigeminal nerve. More distally, the por- gelatinosa of the pars caudalis and dorsal
tio minor is continuous with the mandibular horn and in the intermediate grey matter.
division of the trigeminal nerve, which dis- This system inhibits both nociceptive and
tributes both sensory and motor components non-nociceptive units. The raphe-spinal sys-
to the muscles of mastication. tem is often considered to be the final link
in a descending pathway from the central
grey matter of the mesencephalon, which me-
diates the analgesia produced by stimulation
or opiates [81, 116, 312, 347, 401, 851, 1163].
The role of monoaminergic and peptidergic
1 Lobulus parietalis superior : area 5 (S III)

2 Gyrus postcentraJis : areae 3,1,2 (S J)
3 Gyrus precentralis: area 4
4 Premotor cortex : area 6
5 Area S II
6 ucleus ventralis posterolateraJis thalami
7 Nucleus ventralis posteroOledialis thalami
10 Nucleus sensorius principal is nervi trigemini
11 ucleus raphes magnus
12 ucleus reticularis gigantocellularis
13 ucleus spinalis nervi trigemini, pars oral is
14 ucleus spinalis nervi trigemini, pars interpolaris
16 Nucleus gracilis
17 Nucleus spinalis nervi trigemini, pars caudalis
J8 Decussatio pyramidum
19 Raphe-spinal fibres in funiculus posterolateralis
20 Tractus spinothaJamicus
21 ucleus proprius corous posterioris
22 Cellulae marginales cornus posterioris
24 Nervus trigeminus, radix sensoria
27 Tractus pyramidalis lateralis
Fig. 134. Descending systems to sensory relay nuclei

1 Nucleus ventralis posteromediaLis

3 Nucleus tegmentalis dorsalis
6 Nucleus ovalis
8 Nucleus solitarius
11 Tractus solitarius
12 Obex
13 Nucleus ambiguus
Fig. 135. The tractus solitarius. Position of nerves, tracts and nuclei in a dorsal view (5/3 x). Roman
numerals indicate the corresponding cranial nerves
'lIl~~~=~-'"
lX+{§~~"""""'f::::\
X *~~~~-
1 Operculum frontoparietale
2 Limen insulae
3 Nucleus ventralis posteromedialis,
pa rs parvocellularis
4 Tractus trigm inothalamicus dorsalis
7 ucleus parabrachialis medialis
8 ucleus ovalis
9 ucleus solitarius, pars gustatoria
10 ucleus dorsalis nervi vagi
11 ucleus olitarius, pars
cardiorespiratoria
12 ucleus ambiguus
13 Area postrema
14 Tractus solitariospinalis
Fig. 136. The neuronal connections of the visceral afferent system. The nuclei of the special visceral
afferent part of the system (taste) are darkly shaded
cell groups and pathways in regulating pain parvocellular part of the posteromedial ven-
transmission is considered more fully in tral nucleus of the thalamus, in the amygdala
Nieuwenhuys' monograph on the chemical and directly in the gustatory neocortex. The
anatomy of the central nervous system [997]. gustatory neocortex and the lateral nucleus
The descending connections from the spi- of the amygdala are reciprocally intercon-
nal tract nucleus of the trigeminal nerve [189, nected. Both project to the parvocellular part
191, 1165] and the ventral region of the dor- of the posteromedial ventral nucleus [765,
sal column nuclei [221, 749] to the dorsal 766, 1001, 1007, 1008, 1538]. The cortical
horn can be considered as part of an intra- projection area for taste is found in the fron-
nuclear pathway. Their function has not yet tal and parietal operculum and in the limen
been elucidated. insulae [105].
The dorsal longitudinal fascicle, located in
the nucleus praepositus hypoglossi and the
central grey matter, connects the caudal part
The Visceral Afferent Systems of the solitary nucleus with the dorsal teg-
(Figs. 135 and 136)
mental nucleus and more rostral structures
[948]. Crossed, solitariospinal fibres descend
Visceral afferent fibres and small contingents from the caudal part of the nucleus [1383].
of somatosensory fibres enter the medulla
oblongata in the facial, glossopharyngeal and
vagus nerves. The somatosensory fibres join
the spinal tract of the trigeminal nerve. The
visceral afferents descend more medially, in
the solitary tract, to terminate on the solitary
nucleus. Special visceral afferents subs erving
taste terminate in the rostral, gustatory part
of this nucleus. General visceral afferents de-
scend more caudally to terminate in the cau-
dal, cardiorespiratory part of the solitary nu-
cleus and the area postrema. Some of these
fibres cross caudal to the obex and ascend
for some distance in the contralateral solitary
tract, others terminate in the ventral part of
the lateral cuneate nucleus and in the dorsal
nucleus of the vagus nerve [98, 656, 657,
1000]. Ascending root fibres of the facial
nerve terminate in the oval nucleus, a rostral
prolongation of the solitary nucleus located
dorsal to the sensory nuclei of the trigeminal
nerve [58, 98, 1120].
In humans, fibres from the dorsal tri-
geminothalamic tract originate from the oval
nucleus and the principal trigeminal sensory
nucleus. In rats and cats, the ascending taste
pathway contains an extra synapse in the me-
dial parabrachial nucleus. From here, fibres
ascend ipsilaterally (and in the rat also bilat-
erally) in the region of the dorsal trigemino-
thalamic tract, to terminate in the medial
Special Sensory Systems
The Vestibular System Deiters receives only a few primary vestibular

(Figs. 137-140) fibres, in its ventral part [207].
Although single vestibular nerve fibres
The Vestihular Nerve and Nuclei have been observed to distribute their termi-
nal branches to the superior, medial and infe-
The vestibular system provides information rior vestibular nuclei [597, 848] experimental
about the position and motion of the head studies have shown that individual receptor
in space. The receptive elements are the hair components of the labyrinth project preferen-
cells of the membranous labyrinth. These tially to particular portions of the vestibular
cells are located in the cristae ampullares of nuclear complex. Cells in the vestibular gan-
the semicircular canals and in the maculae glion that innervate the cristae of the semi-
of the utricle and the saccule. The first-order circular canals project primarily to the supe-
elements are bipolar neurons, the somata of rior and medial vestibular nuclei, while those
which constitute the vestibular ganglion, lo- that innervate the maculae of the utricle and
cated in the internal auditory meatus. The saccule have central terminations mainly in
peripheral processes of these cells are distrib- the inferior vestibular nucleus [207, 369,
uted to the hair cells. The central processes 1287].
of the bipolar cells constitute the vestibular Some primary vestibular fibres project
part of cranial nerve VIII, which enters the beyond the vestibular nuclei to terminate
brain stem at the level of the pontomedullary ipsilaterally in the cerebellum, the reticular
junction. Within the vestibular nuclear com- formation and the lateral cuneate nucleus.
plex the fibres of the vestibular nerve bifur- The primary vestibulocerebellar fibres
cate into short ascending and longer descend- reach the cerebellum via the juxtarestiform
ing branches [597,848]. The vestibular nucle- body and terminate profusely in all parts of
ar complex is situated beneath the floor of the nodulus and in the ventral folia of the
the lateral part of the fourth ventricle. It com- uvula. Much smaller numbers of fibres reach
prises four cell masses, the superior, lateral, the flocculus, deep folia of vermal lobules V
medial and inferior (or descending) vestibu- and VI and the lingula. All primary vestibu-
lar nuclei (Figs. 104-107). Most ascending locerebellar fibres terminate as mossy fibres
branches of the vestibular nerve fibres termi- in the granular layer of the cerebellar cortex
nate in the superior vestibular nucleus, but [207, 706]. The vestibuloreticular fibres
some ascend to the cerebellum (see below). terminate in the parts of the gigantocellular
The descending branches of the vestibular reticular nucleus situated adjacent to the
nerve constitute a conspicuous bundle in the vestibular nuclear complex, and particularly
inferior vestibular nucleus. The fibres in this in a cell group lying immediately caudoven-
bundle issue numerous transversely orientat- tral to the abducens nucleus [207]. There is
ed collaterals, which pass to the inferior and experimental evidence suggesting that this re-
medial vestibular nuclei [597, 848]. The large- ticular cell group projects primarily to the
celled lateral vestibular nucleus or nucleus of contralateral abducens nucleus [372, 434,
1 Nucleus interstitialis
8 Nervus vestibularis
10 Tractus vesti bulospinalis lateralis
11 Tractus vestibulospinalis medialis
Fig. 137. The vestibular system. Position of nerve, tracts and nuclei 10 a dorsal view (5/3 x). The
lateral vestibulospinal tract is shown on the right
Special Sensory Systems 167
13 14
1 Nucleus interstitialis (Cajal)

2 Nucleus accessorius nervi
oculomotorii
3 Cerebellum
A 4 Fasciculus uncinatus cerebelli
5 Fasciculus longitudinalis media.1is
8 Nucleus reticularis gigantocellularis
10 Nucleus vestibularis latera lis
13 Fibrae afferentes nervi vestibularis
(VUn
14 Fibrae efferentes nervi vestibularis
(VUn
16 Nucleus cuneatus lateralis
17 Fibrae spinovestibulares
18 Tractus spinocerebeUaris posterior
Fig. 13SA, B. The connections of the vestibular system. A Afferent and efferent fibres of the vestibular
nerve and central afferents of the vestibular nuclear complex. B Internuclear and commissural connec-
tions of the vestibular complex
837]. The primary vestibular fibres which inferior and lateral vestibular nuclei. Spino-
pass to the lateral cuneate nucleus originate vestibular fibres, arising largely from caudal
from macular regions and terminate in the levels of the spinal cord, ascend ipsilaterally
rostrolateral portion of the nucleus. and terminate in the lateral vestibular nucle-
It is noteworthy that the vestibular nerve, us and in the caudal parts of the medial and
in addition to afferent fibres, also carries a inferior vestibular nuclei [1093]. Further-
number of efferent fibres. The latter arise more, Carpenter and co-workers [206, 216]
from a group of neurons which is interposed recently reported that in the cat the central
between the abducens and the superior ves- cervical nucleus, a cell mass located in the
tibular nuclei and lies embedded in the reticu- intermediate grey matter of the spinal cord,
lar formation. The projection to one laby- projects to the contralateral inferior and su-
rinth has a bilaterally symmetrical origin perior vestibular nuclei. Cells located in the
from the cell groups on each side of the brain caudal half of the nucleus praepositus hypo-
stem [411]. In the cat about 20% of the effer- glossi project to the inferior and medial ves-
ent vestibular neurons give offaxons which tibular nuclei, and these connections are par-
reach both labyrinths [289]. Goldberg and tially reciprocated by efferents from those
Fernandez [411] have demonstrated that the vestibular nuclei [206]. The only vestibular
efferent fibres have a predominantly exicita- nuclear afferents descending from higher lev-
tory influence on the afferent activity in the els of the neuraxis arise from the interstitial
vestibular nerve. They suggest that the effer- nucleus of Cajal and from the midline viscer-
ent vestibular system functions to extend the al nuclei of the oculomotor complex
dynamic range of the afferents during the (VOMC). The fibres emanating from the in-
large accelerations accompanying voluntary terstitial nucleus of Cajal descend in the me-
head movements. dial longitudinal fascicle and terminate in the
ipsilateral superior and medial vestibular nu-
clei, whereas those from the VOMC project
Afferents to the Vestibular Nuclei to the medial and inferior nuclei [206, 216].
The vestibular nuclear complexes of the
Apart from afferents from the vestibular gan- two sides are amply interconnected by com-
glion the vestibular nuclei receive input from missural fibres (Fig. 138 B). The superior and
various other sources, among which are the medial nuclei send numerous fibres to their
cerebellum, the spinal cord, the nucleus prae- counterparts the opposite side [371]. More-
positus hypoglossi and the interstitial nucleus over, the medial nuclei project strongly to
of Cajal. The cerebellovestibular fibres arise the contralateral superior nuclei, and vice
from Purkinje cells located in the flocculono- versa. The inferior vestibular nucleus pro-
dular lobe, the uvula and the anterior lobe vides afferents to the opposite superior, me-
vermis, and from the fastigial nucleus [206, dial and inferior nuclei [206]. Extensive inter-
216]. The vermis (nodulus and uvula) and nuclear connections between the individual
the flocculus project in a complementary nuclei of the vestibular complex have also
manner [1456] to the ipsilateral superior, me- been demonstrated [206, 216, 1162]. Experi-
dial and inferior vestibular nuclei. The princi- mental evidence thus indicates that the supe-
pal afferents to the lateral vestibular nucleus rior vestibular nucleus is reciprocally related
are the axons of Purkinje cells which are lo- to the medial and inferior nuclei.
cated in a paramedian strip of the anterior
lobe vermis (Fig. 166). Because of this, the
lateral vestibular nucleus may be considered
a ventrally displaced cerebellar nucleus. The
cerebellovestibular fibres originating from
the fastigial nucleus project bilaterally to the
Efferents from the Vestibular Nuclei terior funiculus of the spinal cord. Fibres
originating from the medial and inferior ves-
The vestibular nuclei distribute their efferents tibular nuclei constitute a descending compo-
more widely in the neuraxis than any other nent of the MLF which is known as the me-
special sensory system [206]. These efferents dial vestibulospinal tract. Fibres from each
can be grouped as follows (Fig. 139): of the two participating vestibular nuclei pass
bilaterally to the cervical part of the spinal
1. Fibres passing to the cerebellum.
cord, where they make monosynaptic con-
2. Projections to the spinal cord and to ocu-
nections with motor neurons innervating the
lomotor centres.
neck muscles [1015]. These connections form
3. Fibres ascending to the thalamus.
part of a set of reflex circuits by which the
The secondary vestibulocerebellar fibres position and movements of the head are cor-
originate from the inferior, medial and supe- related with those of the eyes.
rior nuclei and project to the ipsilateral nodu- Fibres originating in the medial vestibular
lus, uvula and anterior lobe vermis and, bilat- nucleus, which descend with the medial vesti-
erally, to the flocculus. They terminate, like bulospinal tract, have also been observed to
the primary vestibulocerebellar fibres, as terminate in the contralateral central cervical
mossy fibres in the granular layer of the cere- nucleus. This nucleus receives dorsal root
bellar cortex [165, 206, 216]. Certain parts fibres conveying impulses from joint surfaces
of the vestibular nuclear complex are able of the upper cervical vertebrae and projects
to influence the activity of cerebellar climbing crossed fibres to the superior and inferior
fibres. However, this occurs via an indirect vestibular nuclei as well as to the cerebellum
vestibulocerebellar pathway which is synapti- [206].
cally interrupted in the inferior olive. Vesti- The ascending vestibular contributions to
bulo-olivary fibres originate from the inferi- the MLF originate mainly from the superior
or, medial and superior vestibular nuclei and and medial vestibular nuclei. The fibres from
terminate in specific parts of the inferior olive the superior nucleus remain largely on the
(in the cat these are the ipsilateral subnucleus ipsilateral side, whereas those from the me-
p, and bilaterally the dorsomedial cell col- dial nucleus ascend bilaterally. Together
umn and the caudal medial accessory olive these fibres constitute the vestibulomesen-
[206, 207, 1177]. In view of the known topo- cephalic projection. They terminate in the nu-
graphy of the olivocerebellar projections, it clei of the extraocular muscles, i.e. the abdu-
is likely that the climbing fibre-mediated ves- cens, trochlear and oculomotor nuclei, the
tibular information mainly influences the interstitial nucleus of Cajal, and in the rostral
caudal vermis. interstitial nucleus of the MLF. The intersti-
The projections to the spinal cord and tial nucleus of Cajal gives rise to a small inter-
oculomotor centres from the lateral vestibu- stitiospinal tract that, by way of the MLF
lar nucleus form the lateral vestibulospinal and the anterior funiculus, passes throughout
tract, an ipsilateral, somatotopically ,organ- the length of the brain stem and spinal cord.
ised bundle that descends in the lateral part The fibres that connect the vestibular nu-
of the anterior funiculus. Its fibres are distrib- clei with the oculomotor centres are links in
uted throughout the spinal cord, exerting fa- elementary three-neuron reflex paths. The
cilitatory influences on spinal reflex activity impulses conveyed by these paths help stabil-
and on extensor muscle tone. The remaining ise retinal images by producing eye move-
three vestibular nuclei discharge efferents ments compensatory for head movements.
into the medial longitudinal fascicle (MLF), According to the recent experimental studies
a conspicuous bundle extending in a dorso- of Carpenter and co-workers [206, 216] all
medial position throughout the brain stem. four vestibular nuclei project to the nuclei
Caudally this bundle passes over into the an- of the extraocular muscles. The main results
1 Area 2v
2 Area 3a
4 Nucleus ventralis posterior inferior
5 Nucleus interstitialis roslralis of
the FLM
6 Nucleus interstitialis of Cajal
7 Tractus interstitiospinalis
9 Tractus vestibulothalamicus
11 Tractus vestibulomesencephaLicus
(FLM)
13 Cerebellum
21 Nucleus oLivaris inferior
22 Tractus vestibulospinalis medialis
23 Tractus vestibulospinalis lateralis
Fig. 139. The connections of the vestibular system: efferent connections of the vestibular nuclei
t Musculus obliquus superior

2 Musculus rectus medialis
3 Musculus rectus superior
4 Musculus obliquus inferior
5 Musculus rectus inferior
6 Musculus rectus lateralis
8 Nervus oculomotorius sinister
9 Nervus oculomotorius dexter
10 Nervus abducens
11 Nucleus nervi II I
12 Nucleus nervi IV
13 Pedunculus eerebellaris superior
15 Nucleus nervi VI
18 Ganglion vestibulare
19 Canalis semicircularis anterior
20 Canalis semicircularis lateralis
21 Canalis semicircularis posterior
Fig. 140. Connections between the semicircular canals and the nuclei of the extraocular muscles. Excita-
tory connections are indicated in black; inhibitory connections with open contours. The synaptic rela-
tions of the inhibitory vestibuloocular relay cells are not indicated
of these studies may be summarised as fol- lothalamic fibres remains ipsilateral and fol-
lows. Projections from the medial vestibular lows the so-called ascending tract of Deiters,
nucleus supply both abducens nuclei and as- a bundle located immediately dorsolateral to
cend bilaterally and asymmetrically in the the MLF. The vestibulothalamic fibres termi-
MLF; crossed fibres project to the contralat- nate bilaterally in the pars oralis of the ven-
eral trochlear nucleus, the oculomotor com- tral posterolateral nucleus (VPLo), the ven-
plex and the interstitial nucleus of Cajal, tral posterior inferior nucleus (VPI) and the
while a small number of uncrossed fibres pro- pars caudalis of the ventral lateral nucleus.
ject exclusively to the oculomotor complex. In the monkey physiological experiments
The inferior vestibular nucleus issues exclu- have revealed two separate vestibular cortical
sively crossed ascending fibres in the MLF, areas, area 2v at the rostral tip of the intra-
which terminate contralaterally in the troch- parietal sulcus and area 3a in the floor of
lear nucleus and in the oculomotor complex. the central sulcus. It seems likely that fibres
Ascending projections from the ventral part originating in the VPLo represent the link
of the lateral vestibular nucleus terminate in the vestibulocortical pathway to area 3a,
sparsely in the ipsilateral abducens nucleus, and that the VPI is the source of the vestibu-
and substantially in the ipsilateral oculomo- lar thalamocortical fibres terminating in area
tor complex. The superior vestibular nucleus 2v [248,360,757,840,1017,1112].
projects via the MLF to the following ipsilat-
eral centres: trochlear nucleus, oculomotor
complex, interstitial nucleus of Cajal and nu-
The Auditory System
cleus of Darkschewitsch. Contralateral pro-
(Figs. 141-143)
jections of the superior vestibular nucleus de-
cussate in the isthmus region and terminate
contralaterally in the oculomotor complex Auditory Centres
and, sparsely, in the trochlear nucleus, the
interstitial nucleus of Cajal and the nucleus The central auditory system includes, in addi-
of Darkschewitsch. The manner in which im- tion to several rhombencephalic cell masses,
pulses from the three semicircular canals in- a mesencephalic, a diencephalic and a telen-
fluence, via the vestibular nuclear complex, cephalic centre (Fig. 141). The rhombence-
the centres for the extraocular muscles, is in- phalic cell masses are the cochlear nuclei, the
dicated diagrammatically in Fig. 140. nuclei together constituting the superior oli-
vary complex and the nuclei of the lateral
lemniscus.
Projections from the Vestibular Nuclei The human cochlear nuclei are composed
to the Cerebral Cortex of a ventral and a dorsal nucleus, which are
comparable though not identical in their
The pathways along which vestibular stimuli cytoarchitecture to those of other mammals
reach the level of "conscious experience" are [938]. The ventral cochlear nucleus comprises
still imperfectly known. However, there is ev- a rostral area of spherical cells, a central area
idence that sparsely scattered cells located of multipolar and globular cells, a caudal
within the superior, medial and lateral vesti- area of so-called octopus cells and a latero-
bular nuclei give rise to fibres which termi- dorsal cap of small neurons. The dorsal coch-
nate in the thalamus. Most of these fibres lear nucleus is well developed, but does not
ascend in the area of the ipsilateral lateral show the characteristic laminated pattern
lemniscus. They then decussate in the isthmus seen in other mammals.
region, after which they pass diffusely The superior olivary complex is largely
through the midbrain tegmentum and the embedded in the trapezoid body, a large,
fields of Forel. A smaller number of vestibu- transversely oriented fibre stream. It com-
prises the medial and lateral superior olivary Auditory Pathways

nuclei and the nucleus of the trapezoid body.
The medial and lateral superior olivary nuclei Figure 141 shows that the centres mentioned
are elongated cell masses, the former of above are interconnected by distinct fibre
which is much better developed than the lat- streams. The cochlear nuclei give rise to two,
ter. The nucleus of the trapezoid body is, just more or less transversely orientated bundles,
as the lateral superior olivary nucleus, poorly the trapezoid body and the dorsal acoustic
developed in man [1029]. These three cell stria. The trapezoid body crosses in the ven-
masses, i.e. the medial and lateral superior tral part of the tegmentum. After having
olivary nuclei and the nucleus of the trape- reached the lateral part of the pons, the bun-
zoid body, are surrounded by a zone contain- dle makes an abrupt rostral turn and con-
ing small groups of cells that vary in size tinues as the lateral lemniscus.
and shape. The latter cell groups are collec- The nuclei of the superior olivary complex
tively designated as the periolivary nuclei. and the cell mass bearing its name lie embed-
They were formerly believed to be involved ded in the trapezoid body. The dorsal acous-
only descending pathways. However, it has tic stria passes over the inferior cerebellar pe-
recently been established that the cell groups duncle. Arching ventrally, this bundle tra-
of the peri olivary nuclei also give rise to verses the reticular formation to reach the
ascending projections [3]. region of the contralateral superior olivary
The nuclei of the lateral lemniscus consti- complex where it continues into the lateral
tute an elongated strand of cells which lies lemniscus. The bundle just mentioned is situ-
embedded in the fibre bundles of the lateral ated in the dorsolateral part of the pons and
lemniscus as it ascends through the pontine the caudal midbrain. It ascends to the inferior
tegmentum to the midbrain. Two nuclei, a colliculus, in which most of its fibres termi-
ventral and a dorsal, can be distinguished. nate. The brachium colliculi inferioris, a
Experimental studies [407, 726, 1501] have prominent bundle visible on the surface of
shown that these nuclei represent important the midbrain, connects the inferior colliculus
links in the ascending auditory pathway. with the medial geniculate body. The inferior
At the mesencephalic level the auditory colliculi of both sides are, in addition, inter-
system is represented by the inferior collicu- connected by commissural fibres. The final
Ius. This structure, which is a most important link in the auditory system is formed by the
relay station in both the ascending and de- auditory radiation through which the medial
scending auditory projections, consists of a geniculate body is connected with the cere-
large, compact central nucleus and a more bral cortex. Passing laterally, this radiation
diffuse, mainly laterally situated zone of grey traverses the sublenticular portion of the in-
matter. The specific diencephalic nucleus for ternal capsule.
hearing is the medial geniculate body, which It is noteworthy that from the level of
forms part of the dorsal thalamus. This nu- the inferior colliculus onwards the ascending
cleus comprises three main divisions, medial, auditory projection can be subdivided into
dorsal and ventral. Part of the ventral divi- a 'core' projection and a 'belt' projection
sion has a laminar organisation. The telen- [164]. The final target of the core projection
cephalic auditory centre occupies the posteri- is the primary auditory cortex, whereas the
or portion of the cortex covering the upper target of the belt projection is formed by the
surface of the temporal lobe. auditory cortical fields surrounding the pri-
mary auditory area. The core and belt projec-
tions include separate relay stations in the
inferior colliculus and in the medial genicu-
late body. Within the inferior colliculus the
central nucleus and the zona lateralis repre-
1 Planum temporale
2 Gyrus temporalis transversus (HeschJ)
3 Radiatio acustica
7 Colliculus inferior
8 Lemniscus la teralis
9 Nuclei lemnisci lateralis
10 Nucleus olivaris superior lateralis
11 Nucleus olivaris superior medialis
14 Stria acustica dorsalis
15 Nucleus cochJearis ventralis
16 Nucleus cochJearis dorsalis
18 Nervus cochJearis (VIII)
19 Planum polare
20 Limen insulae
21 Sulcus rhinalis
22 Gyrus uncinatus
23 Outline of insula
24 Sulcus temporalis transversus
25 Plane of sectioning
A
Fig. 141A, B. The auditory system. A Position of nerve, nuclei and tracts in a dorsal view (5/3).
The transverse gyrus of Heschl and the planum temporale have been drawn in the true position
relative to the brain stem and the thalamus; B Dorsal view of the temporal lobes, to show the position
of the transverse gyri of Heschl and the temporal plane (1 / 1 x)
1 Planum temporale
2 Gyri temporales transversi
3 Radiatio acustica
4 Corpus geniculatum media Ie, pars dorsalis
5 Corpus geniculatum mediale, pars ventralis
6 Corpus geniculatum mediale, pars medialis
9 Colliculus inferior, zona lateralis
11 Colliculus inferior, nucleus centralis
12 Nucleus lemnisci lateralis dorsalis
13 Nucleus lemnisci lateralis ventralis
14 Decussatio lemniscorum lateralium
16 Nuclei periolivares
17 Nucleus olivaris superior medialis
18 Nucleus olivaris superior lateralis
21 Nervus cochlearis
22 Nucleus cochlearis ventralis
23 Nucleus cochlearis dorsalis
24 Stria acustica dorsalis
Fig. 142. The connections of the auditory system: ascending projections

10
13-----.... ~--- 13
20 ---f~~------' ~----~~~-~ 20
1 Radiatio acustica 13 Fibres to m. tensor tympani

2 Colliculus inferior, zona lateralis 14 Radix motoria nervi trigemini
3 Colliculus inferior, nucleus centralis 15 Nucleus motorius nervi trigemini accessorius
4 Commissura colliculi inferioris 16 Nucleus motorius nervi trigemini
5 Decussatio lemniscorum lateralium 17 Nucleus olivaris superior lateralis
6 Nucleus lemnisci lateral is dorsalis 18 Nucleus nervi facialis
7 Lemniscus lateralis 19 Nucleus nervi facialis, cellulae accessoriae
8 Nuclei periolivares 20 Fibres to m. stapedius
9 Fasciculus olivocochlearis 21 Nervus facialis
10 Nervus cochlearis
11 Nucleus cochlearis ventralis
Fig. 143A, B. The connections of the auditory system. A commissural and descending connections;
B pathways for the middle ear reflexes
sent the' core' and 'belt' centres, respective- The other secondary auditory projection, the
ly. The final link in the auditory core projec- trapezoid body, arises from the ventral coch-
tion arises from the laminated portion of the lear nucleus. Some of its fibres pass, like
ventral nucleus of the medial geniculate those of the dorsal auditory stria, directly to
body, whereas the remaining divisions of that the inferior colliculus, but many other fibres
thalamic centre project to the belt area of are interrupted in one of the nuclei that lie
the auditory cortex [201]. embedded in the trapezoid body or in the
The primary auditory cortex corresponds nuclei of the lateral lemniscus. The following
roughly to the transverse gyrus of Heschl. indirect channels are among those connecting
Part of the belt area of the auditory cortex the ventral cochlear nucleus with the inferior
occupies the planum temporale, i.e. the re- collicuI us:
gion on the superior temporal plane lying
posterior to the transverse gyrus and extend- 1. The spherical cells in the ventral coch-
ing back to the end of the Sylvian fossa. lear nucleus project to the ipsilateral lateral
There are considerable variations in the size superior olivary nucleus and to the medial
and convolution of the supratemporal plane superior olivary nucleus of both sides. Both
in the two hemispheres. Thus, Heschl's gyrus superior olivary nuclei project in turn to the
is usually solitary on the left, but double on inferior colliculus [503]. The cells in the me-
the right side, and the left planum temporale dial superior olivary nucleus are bipolar and
is commonly much larger than the right [374, orientated horizontally. From both the me-
376, 377]. These gross anatomical asymme- dial and the lateral pole of these cells a den-
tries may well represent the morphological dritic tuft arises. The laterally extending den-
substrate for language lateralisation [374]. drites receive their input from fibres of the
ipsilateral side, whereas the medially extend-
ing dendrites receive input from the contra-
The Ascending Auditory Projection lateral side. This highly specific spatial or-
ganisation of afferents to cells in the medial
The primary neurons of the auditory system superior olivary nucleus has to do with the
are the bipolar cells that constitute the spiral role played by these cells in interpreting inter-
ganglion. Their peripheral processes make aural differences in phase and intensity for
contact with the auditory receptors, i.e. the localisation of sound.
outer and inner hair cells in the organ of Cor- 2. The globular cells in the ventral coch-
ti. The central processes of the bipolar cells lear nucleus project to the contralateral nu-
constitute the cochlear division of cranial cleus of the trapezoid body. The latter sends
nerve VIII, which enters the central nervous fibres to the ipsilateral lateral superior oli-
system just caudal to the vestibular division vary nucleus, which in turn projects to the
of the same nerve. The primary auditory ipsilateral inferior colliculus [408, 1279].
fibers bifurcate immediately and are distrib- There is physiological and morphological ev-
uted to both the dorsal and the ventral coch- idence to indicate that auditory information
lear nuclei. Within the cochlear nuclei these reaches the cerebellum via portions of the
bifurcating fibers show profuse collateral pontine nuclei. These portions receive corti-
branching and establish specific types of syn- copontine fibres originating from the audito-
aptic contacts with various types of neurons ry cortex and tectopontine fibres arising from
present in the cochlear nuclear complex. the superior and inferior colliculi. The superi-
Secondary auditory fibres originating or colliculus, it should be noted, receives
from the dorsal cochlear nucleus constitute fibres from the auditory cortex, as well as
the dorsal acoustic stria and pass, via this from the lateral zone of the inferior colliculus
bundle and via the lateral lemniscus, to the [164, 728]. Apart from the indirect pathways
contralateral inferior colliculus e.g. [1024]. just discussed, the cerebellar auditory area re-
ceives a limited number of fibres arising di- in this descending system is formed by the
rectly from cells situated in different parts olivocochlear bundle, which originates from
of the cochlear nuclear complex [561]. the peri olivary nuclei. Most of the fibres of
this bundle decussate in the tegmentum. They
Fibres belonging to the acoustic system
enter the vestibular nerve and join the coch-
decussate in the median plane at various lev-
lear nerve via a vestibulocochlear anastomo-
els (Figs. 142, 143). The components of the
sis. After having entered the cochlea, they
dorsal acoustic stria and the trapezoid body
terminate in the organ of Corti. The higher
have already been discussed. It should, how-
links in the corticocochlear projection are
ever, be added that by way of these fibre sys-
constituted by (1) fibres passing from the pri-
tems several commissural connections be-
mary auditory cortex to the lateral zone of
tween the right and left cochlear nuclear
the inferior colliculus, and (2) fibres descend-
complexes are established [204].
ing (mainly ipsilaterally) to the peri olivary
1. The olivocochlear fascicle, or bundle of nuclei. The exact site at which these fibres
Rasmussen, is a fibre system originating from originate is unknown. Although the projec-
the peri olivary nuclei. The fascicle consists tions from the medial geniculate body to the
largely of decussating fibres, which traverse auditory cortex are reciprocated by descend-
the median plane, passing through the med- ing corticogeniculate fibres, the medial gen-
ullary reticular formation. iculate body is generally not considered to
2. The commissure of the lateral lemniscus be a link in the corticocochlear projection.
nuclei, also known as Probst's commissure, It is, however, worthy of note that Adams
passes through the brachium conjunctivum [2] has produced evidence to suggest that the
and the most rostral part of the pontine teg- medial geniculate sends fibres to the inferior
mentum. It contains true commissural fibers colliculus.
between the right and left dorsal nuclei of The studies of Warr and collaborators
the lateral lemniscus and fibers passing from [459, 1474, 1475, 1476, 1497] indicate that
those nuclei to the contralateral inferior colli- the olivocochlear bundle consists of two sep-
culi [726]. arate efferent systems, which differentially in-
3. The commissure of the inferior collicu- nervate the two types of hair cells situated
Ius consists primarily of true commissural in the organ of Corti. One system originates
fibres interconnecting the two central nuclei from large cells in the ventromedial part of
[179], but also contains other fibres. These the peri olivary area. Their coarse axons pro-
pass from the central nucleus and lateral zone ject mostly to the contralateral cochlea,
of the inferior colliculus to the core and belt where they form large synaptic terminals at
portions, respectively, of the medial genicu- the bases of the outer hair cells. The other
late body [33, 728]. olivo cochlear system arises from smaller neu-
4. The medial geniculate bodies are not rons which are situated more laterally, in the
joined by commissural connections, but the vicinity of the lateral superior olivary nucle-
various auditory cortical areas of the two us. Unlike the large medial cells, these small
sides are reciprocally connected via the cor- elements project mostly to the ipsilateral
pus callosum. cochlea and make "en passage" synapses
with the primary afferent fibres, just beneath
the inner hair cells. Adams [3] also reported
The Descending Auditory Projection the presence of two different types of olivo-
cochlear neurons. However, he remained un-
Parallel to the pathway from the organ of able to confirm Warr's categorisation of these
Corti to the auditory cortex, there is an unin- elements into medial and lateral groups. The
terrupted chain of neurons conducting im- olivocochlear neurons form a system by
pulses in the opposite direction. The final link which the brain can influence its own audi-
tory input. Physiological experiments have in the pathway providing for reflex turning
shown that these neurons inhibit sensory out- of eyes and head in response to auditory stim-
flow from the cochlea. uli. (It has already been mentioned that these
Apart from the corticocochlear path, two fibres also form part of an auditory projec-
other descending auditory projections de- tion to the cerebellum).
serve brief mention. These projections, which Finally, pathways passing from the ventral
arise from the inferior colliculus and from cochlear nuclei to the motor nuclei of the
the peri olivary nuclei, both terminate in the trigeminal and facial nerves constitute reflex
cochlear nuclei. As we have seen, the cochlear arcs that link the organ of Corti with the
nuclei receive their main input from the coch- tensor tympani and the stapedius muscles. In
lea, but do not send fibres back to that organ. response to sounds of high intensity, these
The colliculo-cochleonuclear projection muscles (Fig. 143B) contract reflexly and
consists of an ipsilateral component originat- dampen the vibration of the ear ossicles. In
ing from the lateral zone of the inferior colli- this way the organ of Corti is protected from
culus, and a bilateral component originating damage by excessive stimulation. Borg [144]
from the central nucleus. Both components provided experimental anatomical and
terminate in the dorsal cochlear nucleus physiological evidence that the reflex arcs in-
[249]. dicated are closed by fibres passing from the
The periolivary cochleonuclear projection ventral cochlear nucleus to regions of the ip-
originates, as its name implies, from the cell silateral and contralateral medial superior
groups situated in the peri olivary area. These olivary nuclei, which in turn project to the
cell groups may well receive impulses from tensor tympani and stapedius motoneurons
the ventral cochlear nuclei and from fibres in the motor trigeminal and facial nuclei. He
descending from the inferior colliculus. considered it likely that the stapedius moto-
Adams [3] mentions two groups of perioli- neurons also receive a direct projection from
vary cells which impinge upon the cochlear the ventral cochlear nucleus. Recent studies
nuclear complex, a lateral group of multi- with the retrograde tracer technique [643,
polar cells, whose members are located 674, 828, 923, 1236, 1276, 1356] have shown
around the ipsilateral lateral superior olivary that the stapedius and the tensor tympani
nucleus, and a medial group of small neurons motoneurons both form separate cell groups,
situated ventral to the nucleus of the trape- situated close to but clearly beyond the con-
zoid body. This medial group projects bila- fines of the facial and motor trigeminal nu-
terally to the cochlear nuclei. clei.
Auditory Reflex Pathways The Visual System
The auditory cell masses in the brain stem The Visual Pathway (Figs. 144 and 145)
serve not only as relay nuclei in ascending
and descending projections, but also as reflex The visual pathway begins at the retina, a
centres. Thus, efferents from the cochlear nu- thin, transparent lamina of tissue that is on-
clei enter the reticular formation, where they togenetically derived from the wall of the
synapse with neurons of the ascending reticu- diencephalon. The retina is a laminated
lar activating system. Impulses entering the structure that contains photoreceptor cells,
reticular formation along this path give rise neurons and glia. The photo receptors, i.e. the
to the auditory evoked startle response. rods and cones, constitute the outer layer of
Fibres passing from the lateral zone of the the retina. Impulses from the photoreceptors
inferior colliculus to the superior colliculus are transmitted to bipolar cells that form an
[9, 728] may be considered as important links intermediate zone. The bipolar cells represent
the primary afferent neurons of the visual decussation the axons of the retinal ganglion
system. Unlike the corresponding elements in cells continue without interruption behind
other sensory systems, the retinal bipolar the chiasm as two diverging optic tracts.
cells have only very short axons. These axons These arch around the lateral sides of the
terminate on the ganglion cells, large ele- diencephalon until they reach the lateral gen-
ments that occupy the inner zone of the reti- iculate bodies, where most of their fibres ter-
na. minate. However, some fibres continue in a
The retinal ganglion cells can be subdi- mediocaudal direction and pass via the bra-
vided on both structural and functional chium of the superior colliculus to the superi-
grounds into several categories. The best- or colliculus, the pretectal region and the ter-
known functional subdivision is that into X-, minal nuclei of the accessory optic system
Y - and W -cells, a categorisation which was (Fig. 145). These fibres link the visual system
based on receptive field properties and con- with the centres that regulate the activity of
duction velocities of ganglion cells in the cat the intrinsic and extrinsic musculature of the
[1138]; cf, however, [1139]. Studies of the cat eye. Moreover, the retinotectal fibres form
retina have also revealed the presence of part of an alternative (" extrageniculate")
three principal morphological classes of gan- projection to the cerebral cortex.
glion cells, rx, f3 and y, which are distinguish- In the monkey the A- and B-cells, which
able according to their soma size and axon together account for about 90% of all retinal
calibre and the disposition of their dendrites. ganglion cells, project to the lateral genicu-
There are now compelling reasons to assume late body, whereas not more than 10% of
that the rx, f3- and y-cells correspond to the all retinal ganglion cells, among which are
physiologically distinguished Y-, X- and W- the C-cells, project to the superior colliculus
groups, respectively (cf. [780, 1299] for re- [1070,1071,1299]. The axons of some A-cells
view). Corresponding elements observed in branch and terminate in the lateral geniculate
the monkey retina have been designated A-, body as well as in the superior colliculus
B- and C-cells [783]. [405].
The A- and B-cells are assumed to be the It is noteworthy that some retinofugal
counterparts of the physiologically distin- fibres or collaterals leave the optic chiasm,
guished Y - and X-like cells, whereas the C- enter the anterior part of the hypothalamus,
cells probably correspond to a subgroup of then terminate in the suprachiasmatic nucle-
W-like ganglion cells [1299]. The three types us. This is a small peri ventricular cell mass
of ganglion cells have different terminal tar- that lies, as its name implies, directly above
gets [1070, 1071, 1299]. the chiasm (Fig. 192; [251, 694, 939, 1373]).
The axons of the ganglion cells pass over Via this retinohypothalamic projection the
the inner surface of the retina and converge visual system participates in the regulation
towards the posterior pole of the eye, where of various behavioural rhythms [939]. In the
they pierce through foramina in the sclera rat some of the retinohypothalamic fibres
and then constitute the optic nerve. Only have been observed to extend beyond the su-
after having left the retina do the axons of prachiasmatic nucleus into the lateral hypo-
the ganglion cells acquire a myelin sheath. thalamic area [694].
The optic nerves pass to the optic chiasm, As has already been mentioned, most
which is situated at the base of the brain in fibres of the optic tract end in the lateral geni-
the most rostral part of the hypothalamus culate body. This thalamic centre has a dis-
(Fig. 144). Within the optic chiasm a partial tinct laminated cellular structure consisting
decussation takes place: fibres from the nasal of six more or less concentric layers. These
halves of the retinae cross to the opposite layers are usually numbered 1-6, beginning
side; those from the temporal halves of the from the ventromedial hilar region. Layers
retinae remain uncrossed. After this partial 1 and 2 are composed of larger cells than
in the remaining layers (Fig. 97). The retinal the primary visual cortex below the calcarine
fibres terminate in a highly orderly fashion sulcus, while the upper halves of the retinae
on the layers of the lateral geniculate body. project to the cortex above that sulcus.
Layers 2, 3 and 5 receive fibres from the ipsi- 3. The maculae, i.e. the retinal areas con-
lateral eye, and layers 1,4 and 6 receive fibres cerned with central vision, project to a rela-
from the contralateral eye. Moreover, type-A tively very large area, which forms the poste-
ganglion cells project to the magnocellular rior part of the primary visual cortex. The
laminae, type-B cells to the parvocellular lam- parts of the retinae concerned with the pe-
inae [1071, 1299]. The efferent fibres of the riphery of the binocular field project to a
lateral geniculate body constitute the optic smaller, intermediate part. The nasal periph-
radiation or geniculocalcarine tract, which ery of the retinae, concerned with the tem-
terminates mainly in the primary visual area poral periphery of the visual field (in which
of the cerebral cortex, i.e. Brodmann's area vision is monocular), project to the extreme
17, but also in the surrounding secondary vis- anterior part of the visual cortex.
ual areas 18 and 19. The primary visual area
The two main divisions of the lateral geni-
is also designated the striate cortex (area
culate body project to different cortical lam-
striata), because it contains the highly charac-
inae and activate different groups of cortical
teristic line of Gennari, a macroscopically
cells. Neurons in the magnocellular layers of
visible layer of myelinated fibres bisecting the
the lateral geniculate body, which are mainly
internal granular layer (layer 4; Figs. 72, 79-
innervated by type-A retinal ganglion cells,
82, 85). The striate cortex surrounds the cal-
reach cortical laminae 4 AB and 6. Neurons
carine sulcus on the medial side of the occipi-
in the parvocellular geniculate layers, which
tal lobe (Fig. 11). The fibres of the optic radi-
are mainly innervated by type-B retinal cells,
ation first traverse the retrolenticular part of
terminate in cortical laminae 4 C and 6 [1299,
the internal capsule, then arch around the
1300]. The primary visual cortex (area 17)
lateral ventricle and finally pass posteriorly
projects to the surrounding secondary visual
towards the occipital cortex. Figure 144
(visual association) cortex (areas 18 and 19).
shows that many fibres of the optic radiation
The retinogeniculocalcarine projection is par-
do not reach their destination by the shortest
allelled by a second pathway. Because its
route. Those arising from the lateral part of
fibres do not synapse in the lateral geniculate
the lateral geniculate body and terminating
body, the second pathway is called the extra-
ventral to the calcarine fissure sweep forward
geniculate visual pathway. Its successive
into the temporal lobe and pass laterally over
components are:
the inferior horn of the lateral ventricle be-
fore turning backwards. There is a precise
1. Retinotectal fibres.
point-to-point projection from the retina to
2. Fibres that originate from the superficial
the lateral geniculate body and from the lat-
layers of the superior colliculus and pro-
ter to the visual cortex. Figure 145 illustrates
ject to certain parts of the pulvinar.
some features of these projections in relation
3. Fibres that pass from the pulvinar to both
to the projection of the external world (" vi-
the primary and secondary visual cortical
sual field") on the retina, as follows:
areas. The extrageniculate pathway is in-
1. As a consequence of the partial decussa-
volved in visual orientation and attention.
tion of the retinofugal fibres in the chiasm,
all impulses from the right halves of both Fibres from the visual cortex project to
retinae (representing the left half of the visual various subcortical centres. Efferents, from
field) are transmitted to the right occipital the striate cortex in particular, extend
lobe, and vice versa. through all layers of the lateral geniculate
2. The lower retinal halves (representing body and also project to several other tha-
the upper half of the visual field) project to lamic nuclei, including the pulvinar and the
1 Bulbus oculi
2 Nervus optlcus
3 Chiasma oplicum
4 Tractus opticus
5 Radiatio optica, genu temporale
6 Ventriculus lateralis, cornu inferius
7 Tractus opticus, radix lateralis
8 Tractus opticus, radix medialis
9 Corpus genicula tum laterale
to Lobus temporalis
11 Radialio optica
12 Pulvinar thalami
15 Ventriculus lateralis, pars centralis
16 Splenium corporis callosi
17 Radialio corporis callosi
18 Stratum sagittale
19 Ventriculus latera lis, cornu posterius
20 Radiatio optica, genu occipitale
21 Area striata
22 Sulcus calcarinus
Fig. 144. The visual system I: the retinogeniculocortical projection in a ventral view (1 /1 x). Of the
cerebral hemispheres only the right temporal lobe has been depicted
R L
1 Central part }
2 Binocular part visual field
3 Monocular part
4 Retina
5 Macu la lutea
6 Nervus opticus
7 Chiasma opticum
8 Tractus opticus
9 Tractus opticus, radix medialis
10 Tractus oplicus, radix lateralis
11 ucleus terminalis medialis of the accessory optic system
12 ucleus terminalis lateralis of the accessory optic system
13 Tractus opticus accessorius, fasciculus superior
14 ucleus terrninalis dorsalis of the accessory optic system
15 Regio pretectalis
17 Area 17, primary visual co rtex
:~ ~~: :~} secondary visual cortex
20 Brachium colliculi superio ris
22 Pulvinar thalami
23 Radiatio optica
24 Sulcus calcarinus
Fig. 145. The visual system I: The retinogeniculocortical projection, the extrageniculate visual pathway,
the retinopretectal projection and the accessory optic system. The position of tracts and nuclei corre-
sponds to that in Fig. 144. The infracalcarine part of area 17 is shaded
thalamic reticular nucleus. Other substantial superior fasciculus of the accessory optic
subcortical efferents pass from the visual cor- tract. This fascicle descends from the main
tex to the pretectal region and the superior optic tract between the superior colliculus
colliculus. and the medial geniculate body, proceeds su-
The pretectal region or complex is imme- perficially over the cerebral peduncle, and
diately rostral to the superior colliculus at terminates ventromedially near the exit of the
the level of the posterior commissure. Ac- oculomotor nerve. There are three terminal
cording to a recent analysis in the monkey nuclei:
[579], this complex can be subdivided into
1. The dorsal terminal nucleus is ventral
five different cell groups: the optic tract nu-
to the rostral portion of the superior collicu-
cleus, the pretectal olivary nucleus and the
Ius, close to where the superior fascicle of
medial, anterior and posterior pretectal nu-
the accessory optic tract emerges. It merges
clei. Fibres from the retina pass to the pretec-
with the optic tract nucleus, one of the pre-
tal region, predominantly to the contralateral
tectal cell masses.
side. The pretectal olivary nucleus and the
2. The lateral terminal nucleus is ventro-
optic tract nucleus receive dense bilateral
caudal to the medial geniculate body, at the
projections, whereas the posterior and medial
dorsal edge of the cerebral peduncle.
pretectal nuclei are more sparsely innervated
3. The medial terminal nucleus is in the
by the retina [113, 579]. The involvement of
mediobasal portion of the midbrain, close to
the pretectal complex in visual reflexes and
the medial margin of the substantia nigra.
eye movements will be considered below. Suf-
The three terminal nuclei receive their retinal
fice it to mention here that:
input principally from the contralateral eye.
1. The rodent equivalent of the medial pre- It has long been assumed that the medial ter-
tectal nucleus contains cells which represent minal nucleus is lacking in primates (e.g.
the source of a (mainly contralateral) centri- [1373]); however, its presence has recently
fugal projection to the retina [598, 599]. been demonstrated in two primate species
2. Elements in the optic tract nucleus form [1484].
a relay in pathways connecting the retina
The terminal nuclei of the accessory optic
with the cerebellum. Fibres originating from
tract are reciprocally connected with the nu-
the latter nucleus have been traced in the rat
cleus of the optic tract. This is of functional
and rabbit to the following three "precerebel-
interest because the AOS and the optic tract
lar" nuclei: the reticular nucleus of the pon-
nucleus are both involved in processing direc-
tine tegmentum, the pontine nuclei, and the
tion-selective visual information [535, 1254].
dorsal cap of the inferior olivary nucleus
In rodents and cats, the terminal nuclei
[535, 1367]. Fibres connecting the posterior
of the accessory optic tract, along with the
pretectal nucleus with the caudal dorsal ac-
nucleus of the optic tract, have been shown
cessory olive have been described in the cat
to project differentially to the dorsal cap of
[1 ].
the inferior olive. This part of the inferior
olive sends climbing fibres to the vestibulo-
The Accessory Optic System cerebellum, i.e. the flocculonodular lobe [535,
846, 1358, 1470]. Physiological experiments
The so-called accessory optic system (AOS) have shown that the AOS is intimately in-
is a number of primary optic fibres which volved in visual-vestibular interaction, play-
deviate from the optic tract at various levels ing an important role in processing and dis-
and project to three small mesencephalic cell tributing visual signals subserving compensa-
masses, the dorsal, lateral and medial termi- tory eye and head movements [1254].
nal nuclei of the AOS (cf. [1255]). The largest
contingent of these fibres is known as the
Visual Reflexes cord synapse in the sympathetic ciliospinal

centre. The origin of these fibres is unknown.
The light reflex, i.e. constriction of the pupil The pathway for the accommodation re-
on illumination of the eye, is mediated by flex, i.e. increase in the curvature of the lens
a reflex arc that involves the following links for near vision, is (as far as its mesencephalic
(Fig. 146): centres and efferent limb are concerned) clo-
sely comparable to the pathway for the light
a) Axons of retinal ganglion cells which
reflex. In both pathways the pretectal region,
pass via the optic nerve and tract to the pre-
the accessory oculomotor nucleus and the ci-
tectal region, particularly the olivary pretec-
liary ganglion are successive relay stations.
tal nucleus [158, 1384].
However, the afferent limb of the accommo-
b) Axons of pretectal neurons, projecting
dation reflex includes the visual cortex and
to the accessory oculomotor (Edinger-West-
is thus much longer and much more complex
phal) nuclei of both sides [103].
than that of the light reflex (Fig. 147). The
c) Axons of the parasympathetic pregan-
final link on the efferent side of the accom-
glionic neurons of the accessory oculomotor
modation pathway is formed by the post-
nuclei, which pass with the oculomotor
ganglionic fibres to the ciliary muscle.
nerves to the ciliary ganglia, where they syn-
apse.
d) Postganglionic neurons whose axons in-
Eye Movements
nervate the sphincter pupillae muscle of the
iris.
Eye movements are effected by the coordi-
The pretectal efferents to the ipsilateral ac- nated activity of the six extraocular muscles.
cessory oculomotor nucleus provide for the These muscles are innervated by the third,
direct pupillary light reflex, those to the con- fourth and sixth cranial nerves, the nuclei of
tralateral nucleus for the consensual pupilla- which are located in the medial part of the
ry light reflex. pontine and mesencephalic tegmentum. We
In addition to the response to light, pupil- still lack a coherent picture of the central neu-
lary constriction also occurs following the ral mechanism for the control of eye move-
initiation of ocular convergence. The neural ments; however, the following features of the
pathway for this convergence-induced pupil- circuitry involved are well established:
lary constriction is believed to be indepen-
dent of that for light-induced pupillary con- 1. The abducens and oculomotor nuclei
striction, since the two reflexes are dissociat- contain, in addition to motoneurons, "pre-
ed in certain clinical conditions (the Argyll motor" neurons, which provide reciprocal in-
Robertson sign). The centre for pupillary di- ternuclear pathways [71, 434, 439, 838]
latation is located in the intermediolateral (Fig. 148A: open circles). Axons of internu-
cell column of the upper thoracic cord. The clear neurons in the abducens nucleus cross
preganglionic sympathetic fibres originating the midline and ascend in the contralateral
from this so-called ciliospinal centre ascend medial longitudinal fasciculus (MLF) to the
through the sympathetic trunk and synapse oculomotor complex, where they selectively
in the superior cervical ganglion with post- excite motoneurons of the contralateral me-
ganglionic elements. The axons of the latter dial rectus muscle [195, 520, 877, 969, 1284,
accompany the branches of the internal caro- 1285]. This" six-to-three pathway" is essen-
tid artery and traverse the ciliary ganglion tial for adduction of the eye in horizontal
before they innervate the dilatator pupillae conjugate gaze. Lesions of the MLF between
muscle of the iris. the abducens and oculomotor nuclei produce
Fibres descending through the lateral part the syndrome of internuclear ophthalmople-
of the medulla oblongata and of the spinal gia. The internuclear neurons innervating the
1 Pupilla
2 Iris
3 Retina
4 Nervi ciliares breves
5 Nervus opticus
6 Ganglion ciliare
7 Ramus sympathicus
9 Nucleus accessorius nervi
oculomotorii
10 Regio pretectalis
11 Commissura posterior
13 Ganglion cervicale superius
14 Ramus communicans
15 Centrum ciliospinale
Fig. 146. The visual system II: the neural reflex arcs of the visual system; the pupillary light reflex
"........... ..
,,
,,
, 1 Lens crystallina
,, 2 Corpus ciliare
I
I
I
3 Retina
I
I 4 Nervi ciliares breves
I
5 Nervus opticus
6 Ganglion ciliare
,:' 8 Nucleus accessorius nervi
I
, I
I oculomotorii
, I
I 9 Regio pretectalis
" ... _'\"
I
\
, 10 Commissura posterior
11 Corpus geniculatum
........... \\'''' laterale
...._ ....... - ...... -
12 Radiatio optica
13 Area striata
Fig. 147. The visual system II: the neural reflex arcs of the visual system; the accomodation reflex
medial rectus motoneurons are affected, so This projection is organised so that the ipsi-
that during attempted gaze to the contralat- lateral connections are inhibitory and the
eral side, the adducting eye shows a paresis contralateral connections are excitatory [69,
[71,504,877]. 71]. These connections form part of the vesti-
2. Physiological and clinical evidence sug- bulo-ocular reflex pathways, which help sta-
gests that a group of cells located in the para- bilise the visual image on the retina by pro-
median pontine reticular formation (P P RF), ducing compensatory eye movements in re-
which forms part of the oral and caudal pon- sponse to head movements.
tine reticular nucleus (Fig. 153), plays a cru- The different vestibulo-ocular pathways
cial role in the control of rapid (saccadic) always address pairs of synergistic muscles
conjugate eye movements [690, 1285]. The from the two eyes, which exert their actions
PPRF projects to the abducens nucleus of in one of the three planes of the semicircular
both sides [198, 434]. Physiological experi- canals ([427], Fig. 140). Through these path-
ments have shown that the fibres passing to ways excitation and inhibition of two oppo-
the ipsilateral abducens nucleus activate mo- site canals by a movement of the head in
toneurons and internuclear neurons [432, space, leads to a conjugate (compensatory)
505], whereas those passing to the contralat- eye movement in the opposite direction.
eral abducens nucleus exert an inhibitory ac- Movements in the plane of the horizontal ca-
tion [432]. The PPRF also projects to the ipsi- nals are effectuated through crossed, excit-
lateral prepositus hypoglossal nucleus and to atory connections from the medial vestibular
the ipsilateral rostral interstitial nucleus of nucleus to the abducens nucleus, which inner-
the MLF (RIMLF). Both nuclei send fibres vates the lateral rectus muscle, and the inter-
to the oculomotor nucleus. The fibres passing nuclear pathway which recrosses to terminate
from the PPRF the RIMLF ascend close to, on medial rectus motoneurons in the ipsilat-
but outside, the MLF [1218]. Afferents to the eral oculomotor nucleus [206, 370, 372, 522,
PPRF include: 760, 839, 1546]. The vertical canals are ex-
cited by a movement of the endolymphe
(a) bilateral projections from the frontal eye
away from the ampullae. Excitation of a pos-
fields, discussed below;
terior canal leads, through a crossed excitato-
(b) projections from the contralateral superi-
ry pathway in the medial longitudinal fascicle
or colliculus;
from the medial vestibular nucleus to the
(c) ipsilateral projections from numerous
trochlear and oculomotor nuclei, to contrac-
brain-stem centres such as the interstitial
tion of the ipsilateral superior oblique and
nucleus of Cajal, the mesencephalic retic-
the contralateral inferior rectus muscles [373,
ular formation, the vestibular nuclei and
426, 518, 519, 521]. Excitation of an anterior
the nucleus prepositus hypoglossi [777,
canal results in contraction of the ipsilateral
1218].
superior rectus (which is innervated from the
The PPRF is also called the "pontine gaze contralateral oculomotor nucleus [1098] and
centre". A unilateral lesion of this centre the contralateral inferior oblique muscles.
leads to a loss of all rapid eye movements This excitatory connection relays in the supe-
towards the ipsilateral side. A bilateral ros- rior vestibular nucleus and/or the portion of
tral PPRF lesion produces horizontal gaze this nucleus located within the floccular pe-
palsy with intact vertical gaze, but a bilateral duncle (the group Y of Brodal and Pom-
caudal PPRF lesion leads to total loss of ra- peiano [171] and passes through the superior
pid eye movements in both the horizontal cerebellar peduncle [1532]. The ipsilateral in-
and the vertical plane [505]. hibitory pathways ascend in the mediallongi-
3. The vestibular nuclear complex sends a tudinal fascicle. Those to the abducens nucle-
massive bilateral projection to the nuclei of us take their origin from the medial vestibu-
the extraocular muscles [206, 373] (Fig. 139). lar nucleus; the inhibitory connections from
the vertical canals relay through the superior of the targets of the prepositus projections
vestibular nucleus. contain neurons whose activity is related to
4. As its name implies the nucleus pre- eye movements. They suggest that the func-
positus hypoglossi (prepositus nucleus) is situ- tion of the prepositus nucleus may be to gen-
ated directly in front of the hypoglossal nu- erate an efference copy of oculomotor activi-
cleus (Fig. 127). It occupies a periventricular ty, and to distribute this signal to centres in
position, immediately adjacent to the MLF the brain stem which are involved in various
(Figs. 105, 106). This nucleus sends fibres to aspects of gaze control.
all of the external eye muscle nuclei, both 5. The interstitial nucleus of Cajal, which
ipsilateral and contralateral, and therefore is situated in the rostral part of the mesence-
has to be considered an important "preocu- phalic tegmentum (Fig. 98), also represents
lomotor" centre ([70, 373, 439, 837]; a preoculomotor centre (Fig. 149). This nu-
Fig. 148 B). Particularly dense projections are cleus receives afferents from the pretectal re-
directed to the contralateral abducens nucle- gion, the superior colliculus and the vestibu-
us and the ipsilateral medial rectus subdivi- lar nuclei [21, 206, 736]. Cortical afferents
sion of the oculomotor nucleus [875]. emanating from the frontal eye field have
The prepositus nucleus receives afferents also been described [65]; cf, however [779].
from the frontal eye fields of both sides [774], The efferents of the interstitial nucleus pass
the ipsilateral interstitial nucleus of Cajal to the oculomotor and trochlear nuclei of
[218], the rostral interstitial nucleus of the both sides, to the ipsilateral medial vestibular
MLF [874] and the nucleus of the optic tract nucleus and to the spinal cord. The fibres
[535]. It entertains reciprocal connections to the contralateral oculomotor and troch-
with the ipsi- and contralateral vestibular lear nuclei pass via the posterior commissure
complex, particularly the medial, inferior and [206, 1285]. The coarse fibres which descend
ventrolateral nuclei, the PPRF and the cere- to the spinal cord constitute the small inter-
bellum [44, 710, 875, 1467, 1545]. stitiospinal component of the MLF.
Apart from the various extraocular motor 6. The rostral interstitial nucleus of the me-
nuclei, the following cell masses receive in- dial longitudinal fascicle, or MLF (RIMLF)
puts from the prepositus nucleus: the contra- is situated rostral to the interstitial nucleus
lateral inferior olive, the contralateral superi- of Cajal in the transition zone between the
or colliculus and pretectum, and several tha- midbrain and the diencephalon. This centre
lamic nuclei including the intralaminar and is directly involved in the generation of fast
ventrolateral nuclei [711, 875]. It is notewor- vertical eye movements [196, 197]. The
thy that the vestibular nuclei also project to RIMLF receives afferents from the frontal
these thalamic centres. eye field and the superior vestibular nucleus,
It has been established that, as far as exci- and is reciprocally related with the PPRF
tation and inhibition are concerned, the af- [196, 779]. Cells of the RIMLF have predom-
ferents from the vestibular complex to the inantly ipsilateral projections to the oculo-
prepositus nucleus are organised similarly to motor nucleus, particularly to the motoneu-
the vestibulo-oculomotor projections [69]. ron subgroups of the vertical eye muscles
Moreover, the activity of nearly every neuron [197, 1285]. Clinicopathological evidence in-
studied in the prepositus nucleus appears to dicates that Parinaud's syndrome (vertical
be correlated with the position and move- gaze paralysis) results from damage to the
ments of the eyes [814, 874]. McCrea and RIMLF or its efferent fibres [1089].
Baker [874, 875] have pointed out that the 7. The superior colliculus has a laminar
afferents from the prepositus nucleus arise structure consisting of alternate grey and
chiefly from regions in the brain which are white layers (Figs. 98, 99). From superficial
directly involved in the supranuclear control to deep, these layers are:
of eye and head movements, and that most
1°
. 0
5---i-+4
1( 1 1 Area 8
2 Regio pretectalis
3 Nucleus interstitialis rostralis fasciculi
i 0
longitudinalis medialis
10 6 Colliculus superior
I 0 8 Nucleus cuneiformis
11 Formatio reticularis pontis paramedian us
13 Nervus abducens
15 Nuclei vestibulares
17 Tractus interstitiospinalis
Fig. 148A, B. The visual system III: The oculomotor pathways. A intrinsic connections and efferents
of the paramedian pontine reticular formation (PPRF); B afferents of the PPRF and connections
of the interstitial nucleus of Cajal. N.h.: Afferent connections are indicated with interrupted lines
and efferent connections with continuous lines
1 Area 8
3 Nucleus ventralis lateralis thalami
4 Nucleus tractus opticus
5 Regio pretectalis
6 Nucleus interstitialis rostralis fasciculi
longitudinalis medialis
11 Formatio reticularis pontis paramedianus
12 Cerebellum
Fig. 149. The visual system III: The oculomotor pathways: the connections of the prepositus hypoglossi
nucleus. N.h.: afferent connections are indicated with interrupted lines and efferent connections with
continuous lines
I
I
I,
,
jI
I.
5~ 11 ~
.
J I
1 Area 8
2 Nucleus interstitialis rostralis
fasciculi longitudinalis medialis
3 CollicuJus superior
5 Formatio reticularis pontis
~/ lo O
I
paramedianus
7 ucleus prepositus hypoglossi
Fig. 150. The visual system III: The oculomotor pathways: the connections of the rostral interstitial
nucleus of the MLF. N.b.: afferent connections are indicated with interrupted lines and efferent connec-
tions with continuous lines
2
I,
\
1
2
3
Areae 17, 18, 19
Retina
Pulvinar thalami
8---·
4
5
6
Colliculus superior
Colliculus inferior
Corpus parabigeminum
00 I
Fig. 151. The visual system III: The oculomotor pathways: the connections of the superficial zone
of the superior colliculus. N.b.: afferent connections are indicated with interrupted lines and efferent
connections with continuous lines
1 Area 8
2 Area 4 11 Colliculus inferior
3 Area 22 12 Nucleus cuneiformis
4 Nucleus ventralis anterior) 13 Tractus tectobulbaris medialis
Nucleus ventralis lateralis 14 Tractus tectobulbaris lateralis
Nucleus lateralis dorsalis thalami 15 Nucleus reticularis tegmenti pontis
Nucleus medialis 16 Nucleus reticularis pontis
Nucleus parafascicularis 17 Nuclei pontis
5 Regio pretectalis 18 Cerebellum
6 Nucleus interstitialis rostralis fasciculi 19 Nucleus nervi abducentis
longitudinalis medialis 20 Nucleus reticularis myelencephali
7 Colliculus superior 21 Nucleus olivaris accessorius medialis
8 Commissura colliculi superioris 22 Nucleus prepositus hypoglossi
9 Nucleus interstitialis (Cajal) 23 Nucleus spinalis nervi trigemini
10 Substantia nigra, pars reticulata 24 Tractus spinotectalis
Fig. 152. The visual system III: The oculomotor pathways: the connections of the deep collicular
zone. N .h.: afferent connections are indicated with interrupted lines and efferent connections with
continuous lines
1. The mainly fibrous stratum zonale originating from several non-visual cortical
2. The stratum griseum superficiale areas, e.g. the motor cortex (area 4), the fron-
3. The stratum opticum tal eye field (area 8) and the auditory cortex
4. The stratum griseum medium (area 22) [304, 363, 778] ; (b) auditory fibres
5. The stratum album medium or stratum arising from the inferior colliculus; (c) soma-
lemnisci tosensory spinotectal and trigeminotectal
6. The stratum griseum profundum fibres; (d) a highly ordered projection from
7. The stratum album profundum the pars reticulata of the substantia nigra
On the basis of connections and functions [435]; and (e) fibres from the fastigial nucleus
the superior colliculus can be partitioned into [86]. It is also worth mentioning that com-
a superficial zone and a deep zone. The su- missural fibres connect the deeper grey layers
perficial zone, which consists of layers 1-3, of the two superior colliculi, and that these
receives primarily visual afferents and gives deeper layers are also reciprocally related to
rise to ascending fibres. The deep zone, which the pretectal region [113].
encompasses layers 4-7, receives heteroge- The deep collicular zone gives rise to as-
neous multimodal inputs and sends descend- cending fibres which terminate within several
ing fibres to centres in the brain stem and dorsal thalamic centres, including the ventral
spinal cord. anterior, ventral lateral, lateral dorsal, medial
The visual afferents of the superior colli- and parafascicular nuclei [477]. These as-
culus orginate in part from both eyes and cending fibres presumably represent collater-
in part from the ipsilateral visual cortex. The als from axons which, after having descended
retinal fibres pass via the optic tract and into the tegmentum of the midbrain, pass
reach the superior colliculus via its brachium caudally towards the rhombencephalon and
(Figs. 15,144,145), to enter the stratum opti- the spinal cord [431]. The descending collicu-
cum [96]. The cortical fibres, which arise lar fibres constitute two bundles, the larger
from areas 18 and 19 as well as 17, appear contralateral medial tectobulbospinal tract
to be topographically organised [363]. They or predorsal fascicle and the smaller direct
also enter the superior colliculus via its bra- lateral tecto bulbar tract [476]. In the tegmen-
chium, but pass mainly into the stratum zon- tum of the midbrain the efferents of the deep
ale. Physiological experiments have shown collicular layers issue fibres to the interstitial
that the superficial collicular zone contains nucleus of Cajal, the rostral interstitial nucle-
a retinotopic map of the contralateral ho- us of the MLF and the mesencephalic reticu-
monymous halves of the visual field, so that lar formation (Fig. 99) and descend near the
central visual regions are represented rostral- median raphe just ventral to the MLF. Some
ly, peripheral regions caudally, upper visual of these fibres continue to cervical spinal seg-
regions medially and lower regions laterally. ments, occupying a position in the medial
The superficial collicular zone gives rise part of the anterior funiculus. The crossed
to a large, topographically organised projec- tectobulbospinal tract issues fibres to numer-
tion to the pulvinar thalami, which arises ous rhombencephalic areas, including the re-
from neurons situated in a cytoarchitectoni- ticular nucleus of the pontine tegmentum, the
cally distinct sublamina of the stratum dorsolateral pontine grey matter, the pontine
griseum superficiale [572, 1056]. The superfi- and medullary medial reticular formation
cial zone also gives rise to a substantial ipsi- (particularly the PPRF), the medial accessory
lateral projection to the corpus parabigemin- olive, the abducens nucleus and the nucleus
um (Fig. 101), a structure which has regional- praepositus hypoglossi. The tectospinal
ly organised bilateral projections back to the fibres terminate in laminae VI and VII of
superior colliculus [73, 477]. the cervical grey matter. The fibres of the
The deep collicular zone receives a variety direct tecto bulbar tract descend laterally to
of afferent projections, including: (a) fibres those of the predorsal fascicle. These fibres
terminate mainly in the reticular nucleus of which reach this zone and have, Via their
the pontine tegmentum, the dorsolateral pon- axons, direct access to the various premotor
tine grey matter and the pontine reticular for- centres mentioned above [897]. In other
mation [431, 476]. words this set of neurons constitutes an "in-
An orderly representation of the visual terface" by which sensory signals are trans-
world, as present in the superficial collicular formed into commands for orientation re-
zone, has also been found in the deeper zone. sponses [573]. This "interface" is in turn
The auditory space and the surface of the under the control of the corpus striatum via
body are also represented in an orderly man- the nigrocollicular projection, the cerebellum
ner in the deeper zone of the superior collicu- via the fastigiocollicular projection and the
Ius; these representations or maps are topo- frontal eye field via corticotectal fibres. It is
graphically aligned with each other as well noteworthy that the eye and head movements
as with the visual map. Thus, the collicular are topographically represented in the deep
visual, auditory and somatosensory maps are collicular zone, and that this "motor"
all in spatial register (for review cf. [573, organisation is, reportedly, in register with
1286]. The relationships between these var- the sensory representations in this zone [573,
ious sensory representations should not be 1286].
thought of as entirely static (e.g. only valid The superior colliculus is not the only
for the situation in which the eyes occupy centre which relays cerebellar influence on
the primary orbital position). In the monkey the orienting response. Without going into
a shift in eye position produces a correspond- details, the dorsolateral parts of the basal
ing shift in the auditory map [615]. However, pontine nuclei and the reticular nucleus of
it is hard to grasp how the collicular repre- the pontine tegmentum (both of which also
sentation of an inherently dynamic body sur- receive fibres from the cerebral cortex and
face could become and remain aligned with the pretectal region) constitute the first link
the representation of the visual world. in a tectopontocerebellar mossy fibre projec-
One of the principal functions of the supe- tion, which terminates in lobules VI and VII
rior colliculus is to participate in the control and in the flocculus (lobule X). The same
of orienting responses, i.e. the steering of ra- cerebellar cortical areas receive a climbing
pid combined movements of eyes, head and fibre projection from certain parts of the me-
body towards external stimuli. For visual dial accessory olive. These parts are domi-
stimuli such an orienting response leads to nated by the superior colliculus via the
bringing the image of an object onto the fo- crossed tecto bulbar tract. The vestibular nu-
vea centralis. Following this location or "fo- clei occupy a prominent position among the
veation" another important function of the premo tor centres which are influenced by the
colliculus, tracking of the object, comes into cerebellum.
operation. The movements of eyes, head and
8. One of the cell masses in the pretectal
body involved in orientation are under the
complex, the optic tract nucleus is an essential
control of various premotor centres, among
station in the subcortical pathway mediating
which are the rostral interstitial nucleus of
horizontal optokinetic nystagmus [245].
the MLF, the interstitial nucleus of Cajal,
the PPRF, the internuclear neurons in the 9. A small cortical area situated in the pos-
abducens nucleus, the nucleus praepositus terior part of the middle frontal gyrus, which
hypoglossi, the pontine and medullary reticu- roughly corresponds to area 8 of Brodmann,
lar formation and neurons in the medial part is involved in the initiation of the orienting
of the cervical spinal grey matter. The effer- response as well as of voluntary eye move-
ent cells in the deeper zone of the superior ments, not dependent on visual stimuli
colliculus receive and integrate the sensory (' scanning '). The initial part of the pathway
(visual, auditory, and somatosensory) stimuli from this frontal eye field to the nuclei of
the extraocular muscles is formed by fibres 10. A large occipital cortical field, roughly
that terminate in the pretectal region, the su- corresponding to areas 17, 18 and 19, plays
perior colliculus, the PPRF and the preposi- a role in controlling eye movements induced
tus hypoglossal nucleus [736, 773, 774, 778, by visual stimuli (' pursuit movements '). This
1218]. Direct connections with the rostral occipital eye field, like the frontal eye field,
part of the oculomotor nuclear complex have sends efferents to the superior colliculus and
also been reported [772]. the pretectum [72, 1403].
Ascending Reticular Systems
The Reticular Formation The Raphe Nuclei

(Figs. 153, 154)
The raphe nuclei include the nucleus raphes
pallidus, the nucleus raphes obscurus, the nu-
The area which occupies the central portion cleus raphes magnus, the nucleus raphes pon-
of the brain stem is known as the reticular tis, the nucleus centralis superior and the nu-
formation. Throughout most of its extent this cleus raphes dorsalis. Studies with the aid of
area is occupied by aggregations of loosely histofluorescence and immunohistochemical
arranged cells of different types and sizes, techniques have revealed that many cells in
and the fibre systems that pass through its these nuclei contain the indolamine seroton-
territory are likewise mostly diffusely organ- in. Because the raphe nuclei are not entirely
ised. The term reticular formation refers to congruent with the various aggregations of
the fact that the dendrites of the cells in this serotoninergic neurons, Dahlstrom and Fuxe
area are arranged in bundles that together [279, 280] introduced a new classification of
form a net-like pattern [1211]. The traversing the indolamine-containing cells in the brain
fibre systems pass through the interstices of stem of the rat, distinguishing nine cell
this network. The reticular formation is sur- groups numbered B1-B9. Most of these cell
rounded by cranial nerve nuclei and relay groups have also been recognised in primates
centres and also by the long ascending (lem- [343, 344, 564]. In the following synopsis of
niscal) and descending fibre systems of the the raphe nuclei the description of these
brain stem. Caudally the reticular formation centres as presented by Taber et al. [1349] and
is continuous with the intermediate grey mat- Braak [153] will be harmonised as much as
ter or substance, of the spinal cord; rostrally possible with the subdivision introduced by
it continues into the intralaminar nuclei of Dahlstrom and Fuxe.
the thalamus and into certain aggregations The nucleus raphes pallidus, in the ventral
of subthalamic cells, among which the zona medulla oblongata, borders ventrally on the
incerta may be mentioned. On both cytoar- pyramidal tracts. It corresponds largely to
chitectonic and functional grounds the retic- cell-group B1 of Dahlstrom and Fuxe. Sero-
ular formation can be divided into three lon- toninergic neurons within the superficial ar-
gitudinal columns or zones [162]: (1) a medi- cuate nuclei may be considered as a ventro-
an and paramedian zone, which consists of lateral extension of the B1 group [421, 1258].
the raphe nuclei, (2) a medial zone which con- The nucleus raphes obscurus (Fig. 107),
tains many large cells, and (3) a lateral, large- which corresponds to cell group B2, is situ-
ly parvocellular zone. Figure 153 shows that ated at the same level as the nucleus raphes
a sharp boundary between the medial and lat- pallidus, but occupies a more dorsal position.
eral zones can only be drawn at the levels of The nucleus raphes magnus is found in the
the pons and the rostral medulla oblongata. caudal pons (Figs. 104-107). Many of the
serotoninergic neurons of group B3 are situ-
ated within its confines. The rather small
Median group
.:.' ': ... . (nuclei raphes)
. , ' , ..
0, ': .: : : '. ' •• • :
1 Nucleus raphes dorsalis

. . ...
.
'
' ," ... .. 2
3
Nucleus centralis superior
Nucleus raphes pontis
5 Nucleus raphes obscurus
' .. ...: . .
~ '
Medial group
(fonnatio reticularis medialis)
6 Nuclei cuneiformis+sub-
cuneiformis
7 Nucleus reticularis pont is
oralis
8 Nucleus reticularis tegmenti
pontis
9 Nucleus reticularis pontis
caudalis
. ... ..
,'
.; .........".... ~
,
... :.: ,',
::<···::Mhl
. .. : .........
.... ":;.
::: :. .....: :.. :. ..-:.
'
.. ' .... . .
\: ..:.:.:.::: ....\~~;.~:
' : " ... ... ::."
.. ,
.
: . '"
': .,
..'. :" .
' .'
. ,, '
' , ' :.:::: .... . .
Lateral group
(formatio reticularis lateralis)
11 Nucleus tegmentalis pedunculo-
pontinus, pars compacta
t 2 Nucleus parabrachialis lateralis
14 Nucleus medullae oblongatae
centralis
Motor nuclei
t 5 Nucleus motorius nervi
trigemini
t 6 Nucleus nervi facia lis
t 7 Nucleus ambiguus
Fig. 153. The reticular formation. Semidiagrammatic representation in a dorsal view of the brain
stem. Left : cytoarchitecture; right : subdivision. The cranial nerve motor nuclei are darkly shaded
24
A B c
1 Nucleus gracilis Al,A2, Noradrenenergic cell groups, showing no relation
2 Nucleus dorsalis nervi vagi A5, A7 to cytoarchitectonic entities
3 Nucleus solitarius A6 Noradrenergic cells, constituting the locus
4 Nucleus cuneatus medialis coeruleus
5 Nucleus nervi hypoglossi A6 sc Noradrenergic cells confined to the area subcoerulea
6 Nucleus spinalis nervi trigemini B3, B6, B8 Serotoninergic cell groups
7 Nucleus ambiguus arsvl Area reticularis superficialis ventrolateralis
8 Nucleus funiculi anterioris ncs Nucleus centralis superior
9 Nucleus funiculi lateralis nKF Nucleus of Ki:illiker-Fuse
10 Nucleus olivaris accessorius medialis npbl Nucleus parabrachialis lateralis
11 Nucleus olivaris inferior npbm Nucleus parabrachialis medialis
12 Tractus pyramidalis npgl Nucleus paragigantocellularis lateralis
13 Nucleus prepositus hypoglossi nrg Nucleus reticularis gigantocellularis
14 Nucleus vestibularis medialis nrm Nucleus raphes magnus
15 Nucleus vestibularis inferior nrpc Nucleus reticularis parvocellularis
16 Pedunculus cerebellaris inferior nrpo Nucleus reticularis pontis oralis
17 Nucleus cochlearis dorsalis tpl Tegmentum pontis laterale
18 Corpus pontobulbare
19 Velum medullare superius
20 Griseum centrale pontis
23 Nucleus reticularis tegmenti ponlis
24 Nuclei pont is
Fig. 154. The reticular formation. Diagrammatic transverse sections through the caudal (A), intermedi-
ate (B) and rostral (C) rhombencephalon, to show the position of the reticular formation and of
the various monoaminergic (noradrenergic: ... , adrenergic : ... , serotoninergic: .) and cholinergic: _
cell groups located within the confines of that structure
1 eocortex
2 Gyrus cinguli
3 Striae longitudinales + cingulum
4 Nucleus caudatus
5 Corpus callosum
6 Putamen
7 Fornix 29 Bulbus olractorius
8 Stria terminalis 30 Ansa peduncularis + fibrae
9 Tha lamus amygdalofugales ventrales
10 Stria medullaris 31 Corpus amygdaloideum
II Nucleus habenulae lateralis 32 Gyrus parahippocampalis
12 Nucleus habenulae medialis 33 Gyrus dentatus
13 Area tegrnentalis ventralis 34 Cornu Ammonis A
14 Area lateralis hypothalami 35 Subiculum
15 Nucleus dorsomedialis 36 Substantia nigra
16 Fasciculus telencephalicus medialis 37 Substantia nigra, pars compacta
17 Corpus mamillare 38 Tractus habenulo-interpeduncularis
18 Nucleus veotromedialis 39 Griseum centrale mesencephali
19 Nucleus infuodibularis 40 ucleus raphes dorsalis (B7)
20 Nucleus anterior hypothalami 41 ucleus inlerpeduncularis
21 Nucleus preopticus medialis + lateralis 42 Fasciculus longitudinalis dorsalis
22 Area preoptica lateralis 43 Formatio reticularis mesencephali
23 Nucleus septi medialis + lateralis 44 Nucleus centralis superior (B6 + BS)
24 Nucleus gyri diagonalis 54 Nucleus raphes pontis (B5)
2S Nucleus suprachiasmaticus 46 Griseum centrale pontis
26 Nucleus olfactorius anterior 47 Ventriculus quartus
27 Nucleus accumbens 48 Nucleus raphes magnus (B3)
28 Cortex prefrootalis 49 Nucleus solitarius
Fig. 155A, B. The ascending reticular system. A ascending projections from the raphe nuclei; B afferent
connections of the nucleus raphes dorsalis and the nucleus centralis superior
pontine raphe nucleus (Fig. 103), which cor- pontis oralis. It is important to note that the
responds to cell group B5, lies between the mesencephalic cuneiform and subcuneiform
nucleus raphes magnus and the central supe- nuclei, as well as a narrow medial strip of
rior nucleus. The latter is situated in the ros- the nucleus medullae oblongatae centralis
tral part of the pontine tegmentum and ex- functionally form part of the medial reticular
tends rostrally into the tegmentum of the zone. The cells in the medial reticular zone
midbrain. It encompasses parts of cell groups have long, sparsely branching dendrites that
B6 and B8. The large dorsal raphe nucleus, are oriented in the transverse plane. Their
finally, is situated within and ventral to the axons generally bifurcate into a long ascend-
periaqueductal grey matter and roughly cor- ing and a shorter caudal branch or vice versa
responds to cell group B7. [626]. These branches give off numerous col-
The raphe nuclei receive afferents from laterals along their course. The axonal sys-
numerous centres, many of which form part tems of many of the cells in question remain
of the limbic system. Moreover, these nuclei within the confines of the reticular formation
entertain reciprocal connections with the and thus serve as links in multi synaptic as-
noradrenergic cell groups to be discussed be- cending and descending pathways. However,
low. The pontine raphe nucleus is strongly those of many other elements ascend to the
and bidirectionally connected with the cere- diencephalon or descend to the spinal cord.
bellum [172, 1351] but, remarkably, the sero- The afferent systems to the medial reticular
toninergic innervation of the cerebellum does zone include (a) a large number of fibres orig-
not originate from any of the raphe nuclei, inating from all levels of the spinal cord, (b)
but rather from elements lying in the pontine fibres or collaterals from relay centres of
medial reticular formation [120]. The non- all sensory cranial nerves, (c) cerebellar
cerebellar efferents of the raphe nuclei can afferents, arising mostly from the fastigial
be subdivided into ascending and descending nucleus, (d) fibres descending from the hypo-
projections. The ascending projections arise thalamus and from several limbic forebrain
from the dorsal raphe and central superior structures, and (e) a large projection from
nuclei and are distributed to a variety of pro- the premotor area of the cerebral cortex
sencephalic structures (Fig. 155A). The de- (Figs. 156, 184). The fibre connections of the
scending projections, which originate mainly medial reticular formation suggest that this
from the nuclei raphes magnus, raphes palli- area is integrated into both sensory and mo-
dus and raphes obscurus, innervate numer- tor pathways, and this has been confirmed
ous rhombencephalic centres and the spinal by physiological experiments.
grey matter (Fig. 183A). The serotoninergic
neurons in the brain stem are provided with
highly branched fibres which together inner-
The Lateral Reticular Formation
vate virtually the entire central nervous sys-
tem, thus comprising the most extensive cen-
tral neuronal network yet described. The lateral reticular zone, which is confined
to the rhombencephalon, poses many prob-
lems with regard to both its extent and its
subdivision. In the present synopsis the fol-
The Medial Reticular Formation
lowing six entities, several of which show
considerable overlap, will be distinguished:
The medial reticular zone may also be sub- the area reticularis parvocellularis, the area
divided into a number of centres, which are reticularis superficialis ventrolateralis, the
commonly designated as the nucleus reticu- tegmentum pontis laterale, the noradrenergic
laris gigantocellularis, the nucleus reticularis cell groups A1-A7, the adrenergic cell
pontis caudalis and the nucleus reticularis groups C1-C2 and the cholinergic cell groups
Ch5-Ch6. Throughout most of its extent, the groups A1 and A5 and the adrenergic cell
area or nucleus reticularis parvocellularis group C1 are embedded in the superficial
[162] lies directly medial to the sensory nucle- ventrolateral reticular area (see below).
us of the trigeminal nerve. In the lower me- The superficial ventrolateral reticular area
dulla oblongata, the bulk of the reticular for- receives afferents from the spinal cord [41],
mation should be considered to form part the solitary nucleus [805, 1125, 1207, 1362],
of the lateral zone [538]. As indicated in the medullary and pontine reticular forma-
Fig. 154 A, this zone borders here upon the tion [41], the caudal raphe nuclei [41], the
medially located solitary tract nucleus and parabrachial and Kolliker-Fuse nuclei [281,
the dorsal motor vagal nucleus. It has been 292, 367], the periaqueductal grey matter and
suggested [987] that the axons of the cells several hypothalamic areas [41, 340, 523,
in the area reticularis parvocellularis spread 818]. Its efferents include descending projec-
medially into the magnocellular medial zone. tions to respiratory motoneurons in the cervi-
On this account the parvocellular area has cal and thoracic cord, and to autonomic pre-
been designated as the "sensory" or "asso- ganglionic neurons in the thoracic interme-
ciative" part of the reticular formation [162]. diolateral zone [498,807,811,921,1159]; and
However, autoradiographic tracing studies ascending projections to the Kolliker-Fuse
[541] have shown that the neurons of the par- nucleus [883] and to the hypothalamic su-
vocellular area are the main source of projec- praoptic and paraventricular nuclei [882,
tions to the bulbar motor nuclei, several of 1159].
which lie embedded in this zone (Figs. 153 The lateral pontine tegmentum constitutes
and 185). These neurons probably subserve the enlarged rostral end of the lateral reticu-
several bulbar reflexes and are also the recipi- lar zone. Its medial part consists largely of
ents of projections descending from higher scattered, small neurons. Its lateral part is
levels of the brain [541,743]. formed by a complex of cells that surrounds
The area reticularis superficialis ventrola- the superior cerebellar peduncle along its
teralis occupies, as its name implies, a super- course through the dorsolateral metencepha-
ficial position in the ventrolateral region of lon. This complex can be divided into two
the rhombencephalon. It extends from the aggregations of cells, the medial and lateral
level of the facial nucleus to the spinomedul- parabrachial nuclei, and a ventral extension
lary junction. This area does not constitute which is known as the Kolliker-Fuse nucleus
a distinct cytoarchitectonic entity. Its rostral (Fig. 154C). Several noradrenergic cell-
part largely coincides with the lateral para- groups (the locus coeruleus, the area subcoer-
gigantocellular nucleus as delineated by ulea and cell group A7, see below) and a rath-
Olszewski and Baxter ([1029], cf. Fig. 154B), er diffuse cholinergic cell group (Ch5, see be-
whereas its most caudal portion is constituted low) are located within the lateral pontine
by the nucleus retroambiguus. The main rea- tegmentum.
son for considering this superficial reticular The lateral pontine tegmental grey matter
area as a unit is functional rather than mor- is primarily involved in the regulation of re-
phological. Extensive physiological experi- spiratory, cardiovascular and gastrointestinal
mentation, including local cooling, topical activity. It receives substantial ascending pro-
application of drugs to the surface, local mi- jections from the nucleus of the solitary tract
croinjections of drugs, electrical stimulation and from the area reticularis superficialis
and electrolyticallesioning, carried out main- ventrolateralis [97, 688, 1006, 1125] and pro-
ly in cats, have shown that the area in ques- jects heavily to the thalamus, particularly the
tion plays an important role in cardiovascu- nucleus ventralis posteromedialis, pars par-
lar and respiratory control, and is also in- vocellularis and the intralaminar complex
volved in pain suppression [288, 339, 523, [367, 1001, 1191, 1464], the hypothalamus
869, 870, 1159]. The noradrenergic cell [882, 1191, 1548], the nucleus centralis amyg-
dalae [1001, 1361, 1464] and the bed nucleus macroscopically visible blue-black streak of
of the stria terminalis [367, 1191]. The lateral tissue situated in the floor of the fourth ven-
pontine tegmentum also projects directly to tricle at rostral pontine levels (Figs. 79 and
a portion of the insular cortex [1187, 1191, 102). Evidence suggests that all of the neu-
1244, 1540]. Reciprocal descending forebrain rons situated in the central part of this struc-
projections arise from the insular cortex ture are noradrenergic [385, 1321]. Somewhat
[1187, 1540], the amygdala [550, 721, 927, schematically it may be said that the nor-
1103, 1361] and the hypothalamus [41, 108, adrenergic cell group situated within the lo-
1195]. Descending fibres from the lateral cus coeruleus has three extensions: rostral,
pontine tegmentum project to the nucleus of ventral and caudolateral. The rostral exten-
the solitary tract and the ventrolateral medul- sion consist of elements lying in the caudolat-
lary area [1191, 1362], and to respiratory and eral part of the mesencephalic central grey
autonomic preganglionic motoneurons in the matter. The ventral extension is formed by
spinal cord [539, 626, 810, 1191,1204, 1362]. scattered neurons situated within a cytoarchi-
tectonic entity usually referred to as the nu-
cleus subcoeruleus (A6sc). The caudolateral
extension of the noradrenergic cell cluster sit-
Noradrenergic Cell Groups (Fig. 154)
uated within the locus coeruleus is consititut-
ed by the A4 group already mentioned. The
Neurons that synthesise noradrenalin in the A6 group and its three extensions may be
brain are restricted to the pontine and medul- designated together as the (noradrenergic)
lary tegmental regions. Seven noradrenergic locus coeruleus complex. The cells of the A 7
cell groups, designated as Al-A7, have been group are situated in the rostroventral part
described in rodents [279]. Most of these have of the pontine tegmentum. The Al, AS and
also been recognised in primates [343, 344, A 7 groups form a caudorostral continuum,
366, 385, 562, 563, 927], including man [136, which extends throughout the lateral rhomb-
999, 1025, 1061]. encephalic tegmentum. The caudal part of
Groups Al and A2 are both situated in this noradrenergic lateral tegmental complex
the lower medulla oblongata. The cells of is connected by strands of noradrenergic cells
group Al are embedded in the superficial with the A2 group. Rostrally, a comparable
ventrolateral reticular area; those of group string of cells forms a bridge between the
A2 lie in the solitary tract nucleus, the dorsal locus coeruleus complex and the AS and A 7
vagal nucleus and the adjacent reticular par- groups. The entire rostral accumulation of
vocellular area. On the basis of its topogra- noradrenergic neurons lies, partly at least,
phy, the A2 group is also designated as the within the confines of the medial parabra-
noradrenergic dorsal medullary cell group. chial and Kolliker-Fuse nuclei.
Cells corresponding to group A3 in the rat, The noradrenergic cell groups just dis-
lying just dorsal to the inferior olivary com- cussed give rise to extensively branched as-
plex, have not been observed in primates. cending and descending fibre systems that
Group A4 consists of a band of subependy- course the length of the neuraxis and form
mal neurons which extends along the superi- terminal networks in a variety of grisea.
or cerebellar peduncle. This group merges Throughout the rhombencephalon the dorsal
rostromedially with the caudal portion of part of the lateral reticular zone contains a
group A6. Group A5 is situated in the caudo- large collection of catecholaminergic fibres,
lateral part of the pontine tegmentum and designated by Jones and Friedman [624] as
forms part of the lateral paragigantocellular the longitudinal catecholamine bundle. The
nucleus. dorsomedial medullary (A2) and dorsolateral
Group A6 is a densely packed accumula- pontine cell groups (A6 and A6sc) contribute
tion of cells within the locus coeruleus, a ascending and descending fibres to this bun-
dIe. The axons of the ventrolateral tegmental cord, where they innervate mainly the auto-
group (Al, A5 and A 7) also feed into this nomic preganglionic neurons in the interme-
bundle at successive levels through radially diolateral nucleus [530, 1157].
coursing, transverse fibres [624]. Rostrally
the longitudinal catecholamine bundle tra-
verses the midbrain tegmentum in a position
Cholinergic Cell Groups;
ventrolateral to the periaqueductal grey mat-
Locomotor and Relay Centres
ter and then attains the hypothalamus, where
it joins the medial forebrain bundle. Caudally
the bundle continues into the lateral funicu- Mesulam and colleagues [905, 907] distin-
lus to descend into and innervate the spinal guished six cholinergic cell groups in the
cord. brain of the rat and the monkey. They desig-
nated these cell groups as Chl-Ch6. Four
(Chl-Ch4) are located in the basomedial tel-
encephalon; the remaining two (Ch5-Ch6)
Adrenergic Cell Groups
form a continuum in the caudal midbrain and
rostral rhombencephalon (cf. also [1203]).
Adrenergic neurons are confined to the hypo- The rostral part of cell group Ch5 largely
thalamus [355, 1164] and the lower rhomb- coincides with the pedunculopontine tegmen-
encephalon [356, 530, 560, 695, 1164]. In the tal nucleus (see below); its caudal part ex-
latter region, two adrenergic cell groups have tends into the lateral pontine tegmentum,
been distinguished. By analogy with the no- partly encroaching upon the locus coeruleus
menclature of Dahlstrom and Fuxe [279] complex and the medial parabrachial nucle-
these cell groups have been designated as Cl us. Cell group Ch6 is situated within the pon-
and C2 [530]. Cell group Cl, which is the tine central grey matter. The complex of
largest of the two, is located in the ventrolat- cholinergic neurons just outlined gives rise
eral reticular area. Cell group C2 lies within to an ascending fibre system, the dorsal teg-
the solitary tract nucleus, the dorsal efferent mental pathway of Shute and Lewis [1246],
nucleus of the vagus and the adjacent reticu- which projects upon the superior colliculus
lar formation. Hokfelt and colleagues [530], and several parts of the thalamus, among
who first described cell groups Cl and C2, which are the intralaminar nuclei [548, 1246].
pointed out that these cell groups are identi- It has also been reported that the rhombence-
cal to the rostral parts of the noradrenergic phalic cholinergic complex, and particularly
cell groups Al and A2, respectively. Howe cell group Ch6, projects directly to the cere-
and co-workers [560], on the other hand, em- bral cortex [1203]; cf. also [1096].
phasised that the adrenergic cell groups Cl The pedunculopontine tegmental nucleus
and C2 are situated rostral to, rather than constitutes the caudolateral portion of the
within the confines of, cell groups Al and mesencephalic reticular formation. This nu-
A2. Adrenergic fibres originating from the cleus contains, apart from loosely arranged
Al and A2 groups pass by way of the longi- small cells, a compact cluster of larger ele-
tudinal catecholamine bundle through the ments (Figs. 101 and 153). This compact part
lateral reticular zone and the mesencephalic of the pedunculopontine nucleus (TPc)
tegmentum to the hypothalamus [356, 530, roughly corresponds to an area known as the
1164]. These fibres give rise to terminal fields mesencephalic locomotor region. The latter
in a considerable number of centres, among has received this name because its electrical
which are the locus coeruleus, the periaque- stimulation in post-mamillary-decerebrate
ductal grey matter and the dorsomedial and mammals induces coordinated locomotion
paraventricular hypothalamic nuclei [530]. on a treadmill [446, 1238, 1259]. The TPc
Descending adrenergic fibres enter the spinal receives afferents from, and sends efferents
to, numerous extrapyramidal centres. These ing serotoninergic pathway and the much
connections will be dealt within the section smaller dorsal ascending serotoninergic path-
on motor systems. Suffice it to mention here way.
that the pedunculopontine nucleus receives The ventral ascending serotoninergic path-
afferents from the ventral, 'limbic' parts of way arises principally from the dorsal raphe
the striatum [1338] and projects to the medul- and central superior nuclei. Its fibres sweep
lary reticular formation [1283] as well as to ventrally from these nuclei and then curve
the spinal cord [1338]. rostrally through the ventral tegmentum,
Finally it should be mentioned that within after which they enter the medial forebrain
the domain of the reticular formation three bundle in the lateral hypothalamic area. Dur-
well-defined, typical relay nuclei are found ing its course through the midbrain fibres are
which project massively to the cerebellum. given off to the interpeduncular nucleus, the
These" precerebellar" nuclei are the reticular substantia nigra and the ventral tegmental
nucleus of the pontine tegmentum (Figs. 103, area. In the rostral mesencephalon a number
104), the anterior funicular nucleus or para- of fibres leave the main group to ascend via
median reticular nucleus and the lateral funi- the habenulo-interpeduncular tract to the
cular nucleus or lateral reticular nucleus medial habenular nucleus and to various tha-
(Figs. 107, 108). lamic centres, including the parafascicular
Following these introductory notes, the and midline nuclei, the anterior and ventral
ascending reticular pathways will be dis- nuclear complexes, the lateral dorsal nucleus
cussed. For practical reasons the projections and the lateral geniculate body [133,134,252,
of the three longitudinal zones: median and 264,943].
paramedian, medial and lateral, will be dealt Two large fibre contingents leave the ven-
with separately. It should, however, be kept tral ascending serotoninergic pathway along
in mind that these zones are strongly inter- its course through the lateral hypothalamic
connected and that their mutual boundaries area, one directed laterally, the other ventro-
are both structurally and functionally in- medially. The laterally directed fibres follow
distinct in many places. the ansa peduncularis-ventral amygdalofugal
pathway system and continue through the in-
ternal capsule into the amygdala, the stria-
tum and the external capsule to reach the
Ascending Reticular Pathways
lateral and caudal parts of the neocortex
[134, 265, 368]. The medioventrally directed
Ascending Projections from the Raphe Nuclei fibres innervate a large number of preoptic
(Fig. 153) and hypothalamic centres, among which are
the mamillary body, the dorsomedial, ventro-
Although it is well known that all of the medial and infundibular nuclei, the anterior
raphe nuclei contain a certain proportion of and lateral hypothalamic areas and the me-
non-serotoninergic neurons 115iO] and sever- dial preoptic, lateral preoptic and suprachias-
al important studies on the efferent projec- matic nuclei.
tion of these centres have been carried out In front of the hypothalamus, the ventral
with techniques that do not discriminate be- ascending serotoninergic pathway splits up
tween serotoninergic and non-serotinergic into the following fibre groups [253, 262,
axons, [e.g. 67, 82, 132, 133, 134,252, 1180], 265]:
for the sake of simplicity all efferent raphe
projections will be designated here as sero- 1. Some fibres enter the medullary stria
toninergic. to terminate in the peri ventricular region of
The pathways which ascend from the the thalamus and the medial habenular nucle-
raphe nuclei include the large ventral ascend- us.
2. Some fibres pass with the stria termina- and occipital cortical regions; cells innervat-
lis to the amygdaloid complex. ing the frontal cortex appeared to be located
3. Some fibres follow the fornix to the den- more rostrally and laterally than those pro-
tate gyrus and the cornu Ammonis (Am- jecting to either the parietal or the occipital
mon's horn). cortex. In contrast, the central superior nu-
4. Some fibres enter the diagonal band of cleus appeared to project uniformly upon the
Broca to supply the nucleus of the same name neocortex, without exhibiting any topo-
and the medial and lateral septal nuclei [378]. graphical organisation. The findings of
5. Some fibres pass, by way of the rostral O'Hearn and Molliver [1023] with regard to
portion of the external capsule, to the rostral the topical projection of the dorsal raphe nu-
part of the caudate nucleus, to the nucleus cleus on the cerebral cortex were recently
accumbens and to the rostral and lateral confirmed by Waterhouse and collaborators
parts of the neocortex. [1480]. By injecting different retrograde
6. A bundle of fibres which traverses the tracers in the forelimb sensorimotor or visual
rostral septum enters the cingulum bundle to region of the neocortex and in forelimb (crus
eventually reach the hippocampus via a cau- II) or visual (para flocculus) areas of the cere-
dal approach; during its course fibres are giv- bellum, the latter authors also made the inter-
en off to the cingulate and en to rhinal cortex esting observation that many single dorsal
and to the adjacent neocortical areas. In the raphe neurons provide a common input to
hippocampus the supracallosal projection the functionally related neocortical and cere-
fans out into the subiculum and the cornu bellar regions investigated.
Ammonis. The dorsal ascending serotoninergic path-
7. Some fibers proceed rostrally into the way consists of fibres that pass rostrally in
medial olfactory stria to terminate in the an- and around the dorsal longitudinal fascicle
terior olfactory nucleus and the olfactory of Schutz [134, 344, 943]. Its fibres originate
bulb. mainly from the dorsal raphe nucleus but,
according to some authors, also from the cen-
Experimental studies have shown that to tral superior nucleus [123, 344] and the nucle-
some extent the mesencephalic raphe nuclei us raphes magnus [134]. After having distrib-
have preferential sites of termination. Thus, uted fibres to the mesencephalic central grey
the caudate nucleus and the putamen are sup- matter and the posterior hypothalamic area,
plied by the dorsal raphe nucleus, whereas most of the fibres of the dorsal pathway enter
the efferents of the central superior nucleus the medial forebrain bundle to join the ven-
are distributed mainly to more medially situ- tral pathway.
ated structures such as the hypothalamus and
the hippocampus [67, 133, 1411]. A recent
study, based on double retrograde axonal Afferents to the Mesencephalic Raphe Nuclei
tracing [588], has revealed that various struc-
tures which receive raphe projections have The mesencephalic raphe nuclei are largely
their own characteristic representations with- integrated into typical limbic circuitry, which
in the mesencephalic raphe nuclear complex. is why Nauta [977] included these centres in
Overlapping sets of rostrocaudally arranged what he called the limbic midbrain area
neurons within this complex are associated (Fig. 153). The dorsal raphe and central su-
with the basal ganglia, the amygdala and the perior nuclei receive a massive projection
hippocampus, respectively. As regards the in- from the interpeduncular nucleus [447, 841,
nervation of the neocortex, O'Hearn and 977]. This projection has been considered as
Molliver [1023] have reported that overlap- the final link in a multisynaptic hippocam-
ping cell groups within the dorsal raphe nu- pal-raphe conduction route, encompassing
cleus project differentially to frontal, parietal the precommissural fornix, the septum, the
medullary stria, the medial habenular nucleus forebrain, suggest a general modulatory role
and the habenulo-interpeduncular tract [cf. in regulating behaviour [cf. e.g. 605]. How-
447, 509, 510, 977]. Another major afferent ever, the well-established fact that subsets of
projection to the mesencephalic raphe nuclei neurons within the mesencephalic raphe nu-
originates from the lateral habenular nucleus. clear complex show differential efferent pro-
Its fibres descend with the habenulo-inter- jections [588, 1023, 1073] indicates that this
peduncular tract but bypass the interpedun- complex may also be involved in the regula-
cular nucleus and, thence, ascend directly to tion of more specific physiological and
the nuclei mentioned [8, 447, 510, 841]. This behavioural processes.
projection presumably conveys mainly im-
pulses arising from the nucleus of the diago-
nal band and from the lateral preoptico-
Ascending Projections
hypothalamic continuum to the raphe nuclei.
from the Medial Reticular Zone
Fibres originating from these regions have
(Fig. 156)
been traced to the lateral habenular nucleus
[509, 977]. Fibres originating from the nucle-
After having entered the central nervous sys-
us of the diagonal band and from the lateral
tem, stimuli gathered by the various kinds
preoptico-hypothalamic continuum also pro-
of receptors may reach the cerebral cortex
ject directly to the mesencephalic raphe nu-
either via modality-specific lemniscal systems
clei. However, these fibres do not attain the
or via what is called the non-specific afferent
midbrain via the so-called "dorsal dience-
or extralemniscal system. Since the reticular
phalic conduction system" [1317], but rather
formation of the brain stem forms a crucial
via the medial forebrain bundle [253, 254,
part of the latter system it is also known as
977, 1195, 1320]. This projection is aug-
the ascending reticular or, with reference to
mented by a number of fibres originating
one of its essential functions, the ascending
from the prefrontal cortex (areas 6, 9 and
reticular activating system. The structural
10), apparently the only neocortical region
components of this system are: (1) the spino-
which projects directly to the mesencephalic
reticular projection, (2) reticular afferents
raphe nuclei [56]. The only large projection
from centres of the various sensory cranial
ascending to the mesencephalic raphe nuclei
nerves, (3) direct and indirect reticulothalam-
from lower levels of the neuraxis originates
ic fibres, and (4) the "aspecific" thalamo-
from the pontine central grey matter [841,
cortical projection. These components are
1179], an area which, remarkably, is strongly
illustrated in Fig. 156 and will now be
and reciprocally connected with the inter-
discussed.
peduncular nucleus [447]. Finally it should
be mentioned that one group of authors [8] j. The spinoreticuiar projection (Figs. 123
has described a direct projection from the nu- and 129). A large population of cells in the
cleus of the solitary tract to the mesencephal- intermediate grey matter of the spinal cord
ic raphe nuclei. may be characterised as non-specific, because
Since the mesencephalic raphe nuclei are they receive, via interneurons, convergent in-
dominated by afferents arising either directly formation from nearly all types of primary
or indirectly from medial and basal forebrain afferent fibres [151]. The axons of most of
structures associated with the limbic system these cells decussate in the anterior white
it seems likely that the functional modulation commissure, ascend in the anterolateral funi-
of these nuclei reflects fluctuations in the culus and terminate, either directly, or via
functional state of the limbic system [510]. one or more synaptic interruptions at spinal
The widespread distribution of the efferents levels, in the medial reticular formation.
from the midbrain raphe nuclei, which inner- However, a small contingent of these fibres
vate virtually all principal subdivisions of the bypasses the reticular core of the brain stem
1 Neocortex
2 Corpus striatum
3 Nucleus lateralis posterior
4 uclei intralaminares thalami
5 { Nucleus ventralis lateralis
Nucleus ventralis anterior
6 Nucleus ventralis posterior
8 Zona incerta
10 Nuclei septi
J1 Nucleus preopticus
13 Area tegmentalis ventralis
14 Nucleus lateralis corporis mamillaris
17 Formatio reticularis mesencephali
19 Tractus tcctobulbaris + -spinalis
20 Nucleus reticularis pontis oralis
21 Fasciculus uncinatus cerebelli
22 Nucleus fastigii
24 Trigeminal input
25 Acoustic input
26 Vestibular input
27 Solitarian input
29 Nucleus medullae oblongatae cenlralis
Fig_ 156. Ascending fibre systems of the medial reticular formation. For the descending fibres see
Figures 184 and 185
and projects directly to the intralaminar nu- important component of the ascending retic-
clei, which constitute the chief thalamic relay ular activating system [419, 905, 907].
of the ascending reticular system. Although 4. The "aspecific" thalamocortical projec-
spinal ascending fibres terminate throughout tion. Impulses travelling along the ascending
the reticular formation, the majority is dis- reticular system to reach the diencephalon
tributed to particular levels. One area of may be relayed towards the cortex along dif-
maximal termination is situated in the medul- ferent paths. It has been established [152, 637]
la oblongata and coincides with the caudal that the intralaminar nuclei have a wide-
part of the gigantocellular nucleus and with spread, direct projection to the cerebral cor-
the rostral part of the central nucleus of the tex. This projection is usually considered to
medulla oblongata; a second area of maxi- be constituted by collaterals of the fibres
mal termination is formed by the nucleus reti- which connect the intralaminar nuclei with
cularis pontis caudalis, and a third maximal the striatum. However, physiological experi-
terminal region is found in the most rostral ments have recently shown that in the rostral
part of the pontine tegmentum [150, 162, part of the intralaminar complex the projec-
884]. Remarkably, numerous spinoreticular tions to the striatum and the cortex arise in
fibres which terminate at the latter level do separate sets of neurons of different conduc-
so in the area subcoerulea, i.e. well beyond tion velocities [1293]. Although neuron popu-
the limits of the medial reticular zone [164, lations projecting to different cortical fields
887]; Fig. 154C. overlap widely, a coarse topology exists in
2. Relay centres of most sensory cranial that intralaminar neurons projecting to the
nerves. These are sources of input to the retic- posterior cortex are located more rostrally
ular formation [164]. General somatic affer- and those projecting to the anterior cortex
ent stimuli are conveyed by collaterals from are located more caudally in the intralaminar
the spinal trigeminal nucleus. The vestibular complex [180, 834].
and cochlear nuclei and their pathways also Classically, cortically projecting thalamic
project to the medial reticular formation and nuclei are way-stations in other ascending re-
the same holds true for the visual and the ticular paths. It is known that several of these
olfactory system. Optic impulses are mediat- 'specific' nuclei, e.g. the ventral lateral and
ed by retinotectal and tecto reticular fibre sys- the lateral posterior nuclei, receive afferents
tems [428]. Olfactory impulses probably from the reticular formation. These reticular
reach the mesencephalic reticular formation afferents reach the nuclei in question either
by way of the medial forebrain bundle [958]. directly or via the intralaminar cell groups.
3. Reticulothalamic fibres. Fibres preferen- Before closing this synopsis of the ascending
tially originating from areas in which maxi- reticular system, three comments should be
mal termination of spinal afferents occurs as- made:
cend through the reticular core of the brain
stem and project to the intralaminar thalamic 1. It should be emphasised that although
nuclei. This direct reticulothalamic projec- many of the spinal cells that project to the
tion is parallelled by indirect ones, consisting reticular formation receive impulses from
of two or more successive neuronal links various sensory modalities, and'although fur-
[321, 626, 987]. A particularly large portion ther convergence of sensory impulses occurs
of the reticulothalamic projection has been at the brain stem level, this does not mean
shown to be synaptically interrupted in the that all of the neuronal units participating
lateral part of the pontine tegmentum [626], in the ascending reticular system are entirely
i.e. the area where numerous spinoreticular 'non-specific'. Thus it is known that pain
fibres also terminate [737]. The cholinergic and other protopathic stimuli may well be
neurons located in this area give rise to an conducted upwards by this system with main-
ascending projection, which is considered an tenance of their specificity.
2. The medial reticular zone receives, apart Ascending Projections

from afferents from sensory centres, projec- from the Lateral Reticular Zone
tions from a variety of other sources, among (Figs. 157-159)
which are the cerebral cortex, the dorsal and
ventral pallidum, the lateral hypothalamic In the introduction to this chapter, the lateral
area, the substantia nigra and the cerebellum reticular zone was provisionally subdivided
[221,356, 740, 743, 976, 977, 988, 1338]. Sev- into six parts: the area reticularis parvocellu-
eral of these connections will be dealt with laris, the area reticularis superficialis ventro-
in later sections, particularly in the section lateralis, the tegmentum pontis laterale, the
devoted to the descending reticular systems. noradrenergic cell groups Al-A7, the adren-
It is, however, important to note at this junc- ergic cell groups Cl-C2 and the cholinergic
ture that the ascending and descending retic- cell groups Ch5-Ch6 (Fig. 154). It was
ular systems are by no means independent pointed out that several of these centres show
of each other. Rather, the neural network considerable overlap, and it should be added
of the reticular formation allows considerable here that two of the areas mentioned, the
interaction between ascending and descend- area reticularis superficialis ventrolateralis
ing impulses. and the tegmentum pontis laterale contain
3. The ascending reticular system projects a number of cell aggregations which presum-
not only to the cortex but also to a large ably can be more aptly characterised as
number of subcortical centres. At rostral visceral relay nuclei than as reticular centres.
midbrain levels, the ascending reticular path- In the first part of the following synopsis,
way splits up into a dorsal and a ventral com- the ascending afferent and efferent connec-
ponent, the former continuing in the thala- tions of the parabrachial pontine tegmentum
mus, the latter coursing into the subthalamus and of the superficial lateral reticular area
and hypothalamus [152, 321, 626, 987, 1212, will be discussed. The ascending projections
1293]. The ventral component projects to the of the noradrenergic, adrenergic and cholin-
reticular thalamic nucleus, subthalamic nu- ergic centres mentioned will then be dealt
cleus, zona incerta and the lateral hypotha- with. To our knowledge the parvocellular re-
lamic area in the diencephalon, whereas in ticular zone does not give rise to any long
the telencephalon fibres are distributed to the ascending projections.
preoptic area, the caudate-putamen complex, The parabrachial area can be subdivided
the globus pallidus, the bed nucleus of the into a medial parabrachial nucleus, a lateral
stria terminalis and the substantia innomi- parabrachial nucleus and the ventrally situ-
nata [152, 321, 626, 987, 1132, 1293, 1295, ated Kolliker-Fuse nucleus, which differ both
1434]. The functional significance of most of connectionally and functionally (Fig. 157).
these connections remains enigmatic. The Anatomical and physiological evidence in-
substantia innominata (Figs. 89-92) contains dicates that in rat and cat the medial parabra-
numerous large neurons which provide a ma- chial nucleus plays a crucial role in the pro-
jor projection to the entire cerebral cortex cessing of gustatory information. This nucle-
[631, 683, 904, 910]. It has been suggested us receives a strong projection from the ros-
that these neurons may serve as a relay in tral, gustatory portion of the solitary tract
a route passing from the brain stem reticular nucleus [688, 805, 1006, 1125]. Norgren and
formation to the cortex, an alternative route Leonard [1007,1008] have demonstrated that
to the one passing via the thalamus [626]. responses to taste stimuli can be recorded
from an area roughly corresponding to this
nucleus (which was hence designated the
"pontine taste area "). The medial parabra-
chial nucleus sends efferents to the most me-
dial, parvocellular division of the nucleus
ventralis posteromedialis (VPMpc) of the cortex may well be part of the reticular acti-
thalamus [1191, 1464]. The fourth and final vating system.
link in this gustatory projection is constituted The lateral parabrachial nucleus receives
by fibres which pass from the thalamic centre many fibres from the general viscerosensory
mentioned to the taste area of the cortex [186, caudal part of the solitary tract nucleus [97,
1186]. The latter forms a ventral, insular ex- 915,1006,1125]. Its efferents share a number
tension of area 1 of the somatosensory cor- of end stations with the efferents from the
tex. Whether the medial parabrachial nucleus medial parabrachial nucleus, such as the lat-
subserves a gustatory function in primates eral hypothalamic area, the substantia inno-
as well remains to be established. It has been minata and the central amygdaloid nucleus,
reported [97] that in the monkey the rostral but it chiefly innervates the autonomic re-
portion of the solitary tract nucleus projects gions of the medial hypothalamus, including
directly to VPMpc and not to the parabra- the median preoptic, the paraventricular,
chial region. However, the medial parabra- dorsomedial and ventromedial nuclei [367,
chial nucleus also receives afferents from the 791,889, 1001, 1009, 1167, 1191, 1548]. Most
caudal portion of the solitary tract nucleus, target areas of the lateral parabrachial nucle-
i.e. the portion receiving primary afferents us also receive direct innervation from the
carrying respiratory, cardiovascular and gas- caudal part of the solitary tract nucleus
trointestinal information [97]. As regards ef- [1125].
ferents, the medial parabrachial nucleus has The K611iker-Fuse nucleus, which at least
been demonstrated to project not only to the partly corresponds to the functionally de-
VPMpc, but also to the intralaminar thalam- fined pneumotaxic centre, receives afferents
ic nuclei, the lateral hypothalamic area, the from the caudal part of the solitary tract nu-
central amygdaloid nucleus, the basal mag- cleus [1125] and from the area reticularis su-
nocellular nucleus (which forms part of the perficialis ventrolateralis [883]. It has been
substantia innominata) and the lateral pre- reported that this nucleus sends ascending ef-
frontal and insular cortices [131, 367, 1001, ferents to the lateral preopticohypothalamic
1187, 1191, 1244]. There is physiological evi- zone and to the central amygdaloid nucleus
dence to suggest that the projections to the [1191], but its principal projections are di-
central amygdaloid nucleus and the substan- rected caudally to the lower medulla oblon-
tia innominata carry gustatory information gata and the spinal cord (see the section on
[131, 1004]. The insular cortical area inner- descending reticular systems).
vated by the medial parabrachial nucleus is The area reticularis superficiaUs ventrola-
far more extensive than the gustatory cortex teraUs receives afferents from all levels of the
[cf. 707] and may be designated as the cortical spinal cord. The cells from which this spino-
representation of the central autonomic sys- medullary projection originates are located,
tem [367, 1187]. Interestingly, the non-gusta- mainly contralaterally, in laminae IV, V, VII,
tory parts of this insular region project in VIII and X of the spinal grey matter [41].
a topographical manner to the caudal por- However, the principal afferents to the super-
tion of the solitary tract nucleus [367]. The ficial ventrolateral reticular area originate
lateral prefrontal cortex, which receives a from the caudal, gen'eral viscerosensory area
projection from the medial parabrachial nu- of the solitary tract nucleus [97, 805, 812,
cleus, is not connected to other central auto- 1125, 1159, 1207]. The ventrolateral reticular
nomic areas. This cortical area receives a area projects first and foremost to autonomic
strong projection from the intralaminar tha- and respiratory centres in the spinal cord, but
lamic nuclei, which also receive a dense para- a certain proportion of its efferents ascend
brachial input. According to Saper [1187] the to the hypothalamus, where they terminate
direct and indirect projections from the me- in the supraoptic and paraventricular nuclei
dial parabrachial nucleus to the prefrontal [128,240, 882, 1207]. These fibres convey car-
diovascular and in particular baroreceptive [603, 820, 1207], but most noradrenergic
information to the hypothalamus. The cells fibres reach the hypothalamus via the dorsal
from which these fibres originate in the periventricular system to be discussed below.
ventrolateral medulla excite the vasopressin- The main telencephalic areas in which the
producing cells in the supraoptic and para- dorsal noradrenergic bundle and its branches
ventricular nuclei, which results in vasopres- terminate are: (a) the amygdala (mainly the
sin secretion from the neurohypophysis [129, central and basal nuclei [335, 625, 1375] ; (b)
130, 1159]. the olfactory tubercle or anterior perforated
Containing about half of the total number substance, the anterior olfactory nucleus and
of noradrenalin-synthesising neurons, the lo- the olfactory bulb [124, 336, 1333, 1402]; (c)
cus coeruleus complex (Fig. 158) is quantati- the nucleus of the diagonal band, the medial
vely by far the most important noradrenergic septal nucleus and the bed nucleus of the stria
centre of the brain. Its efferents constitute terminalis [796, 800, 940]; (d) the hippo-
two ascending fibre systems, the (large) dor- campal formation (dentate gyrus, cornu Am-
sal noradrenergic bundle and the (much monis and subiculum) [124, 625, 820, 1402];
smaller) rostral limb of the dorsal periventri- and (e) the entire neocortex, including the
cular pathway. Other efferents are distrib- cingulate, retrosplenial and entorhinal corti-
uted to the cerebellum, and still others de- cal areas [388, 794, 796, 1402]. In most of
scend to the lower medulla oblongata and the areas mentioned the afferents from the
to the spinal cord (see pp. 290-292). The dor- locus coeruleus are intermingled with axons
sal noradrenergic bundle or dorsal tegmental originating from other noradrenergic cell
bundle [793, 794], which forms part of the groups. However, the noradrenergic fibres in-
major longitudinal catecholamine bundle, de- nervating the hippocampus and the neocor-
scribed by Jones and Friedman [624], tra- tex seem to originate exclusively from the lo-
verses the midbrain tegmentum in a position cus coeruleus. In several earlier studies [388,
ventrolateral to the periaqueductal grey mat- 1402] the noradrenergic innervation of the
ter. At rostral mesencephalic levels the bun- neocortex is described as diffuse and uniform
dle arches rostroventrally and reaches the hy- throughout. However, more recent investiga-
pothalamus, where it joins the dorsal portion tions have revealed regional variations in pat-
of the medial forebrain bundle complex. tern and density of cortical noradrenergic in-
Within this complex the dorsal noradrenergic nervation, adhering to cytoarchitectonic
bundle proceeds to the septal region. Along boundaries [785, 955, 956].
its course, numerous smaller and larger The paths along which the efferents from
branches emerge to innervate a large number the locus coeruleus ascend to the various tel-
of mesencephalic, diencephalic and telence- encephalic centres will not be discussed here
phalic grisea. The mesencephalic areas of ter- in detail.' It is, however, worth mentioning
mination include the central grey matter that most of these noradrenergic projections
(substance), the dorsal raphe nucleus, the su- attain their targets via pathways known from
perior and inferior colliculi and the interpe- "classical" neuroanatomy, such as the ansa
duncular nucleus [784, 854]. In the rostral peduncularis-ventral amygdalofugal path-
mesencephalon and at various diencephalic way complex, the diagonal band of Broca,
levels, groups of fibres leave the dorsal nor- the fornix, the stria terminalis and the supra-
adrenergic bundle to innervate the epithala- callosal striae [942, 957].
mus and the dorsal thalamus. The medial and The rostral limb of the dorsal periventricu-
lateral habenular nuclei and most thalamic lar pathway arises mainly from the locus
centres receive a substantial coerulean nor- coeruleus complex and ascends to the dience-
adrenergic input along these channels [596, phalon within the ventromedial part of the
798, 942]. The hypothalamus does receive mesencephalic periaqueductal grey matter,
noradrenergic input from the locus coeruleus forming part of the longitudinal dorsal fasci-
---~
1 Cortex prefrontalis, pars lateralis

2 Operculum frontoparietale (" cortex visceralis")
3 Nucleus ventralis posteromedialis, pars parvocellularis
5 Nucleus preopticus lateralis
6 Nucleus para ventricularis, pars magnocellula ris
7 Area lateralis hypothalami 14 Nucleus parabrachialis lateralis
8 Nucleus preopticus medianus 15 Nucleus parabrachialis medialis
9 Substantia innominata 16 Nucleus of K611iker-Fuse
10 Nucleus dorsomedialis 17 Nucleus solitarius, pars gustatoria
11 Nucleus ventromedialis 18 Area reticularis superficialis ventrolateralis
12 Nucleus supraopticus 19 Nucleus solitarius, pars cardiorespiratoria
13 Nucleus centralis amygdalae 20 Fibrae spinoreticulares
Fig. 157. The ascending reticular system. Ascending projections from the area reticularis superficialis
ventrolateralis and the area parabrachialis
eocortex
2 Gyrus cinguli
3 Striae longitudinale
4 Corpus callosum
5 Fornix
6 Stria termin al is
8 Thalamus
9 Nucleus interstitia lis striae term ina lis
10 Nucleus habenulae latera li
11 Nucleus haben ulae medialis
13 a ciculu telencephalicus mediaLis
14 ucleus para ventricularis, pars parvocellularis
15 ucleu epli mediali
16 Ba ndeletla diagonalis 28 Tractus haben uloi nterpeduncul aris
17 Bulbus olfaclorius 29 Fasciculus longi tudinaLis dorsalis
18 ucleus olfactorius anterior 30 Collicul u uperior
19 Sub ta ntia perforata anterior 31 Colliculu inferior
20 ucleus gyri diagonalis 32 Griseum centrale mesencephali
21 Ansa peduncularis + fibrac amygdalofugales ventrales 33 ucle u raphes dorsalis
22 ucleus cenlra lis amygdalae 34 ucleus interpedunculari
23 ucleus basaLis amygdalae 35 Cortex cerebelli
24 Gyrus dentatus 36 Locus coeruleus
25 Cornu ammonis 37 Area subcoeruIea
26 Subiculum 38 Pedun culus cerebellaris uperio r
27 Gyrus parahippocampa lis 39 uclei centra les cere belli
Fig. 158. The ascending reticular system. Ascending noradrenergic projections from the locus coeruleus
complex
culus complex [344, 793, 1363]. Rostrally, this the mesencephalic and rhombencephalic re-
pathway continues into the diencephalic peri- ticular formation, the contralateral locus
ventricular fibre plexus. Experimental studies coeruleus, the vestibular nuclear complex, the
[1206, 1207] have shown that within the hy- deep cerebellar nuclei, the A1, A2, A5, Cl
pothalamus only the parvocellular portion of and C2 cell groups, the nucleus of the solitary
the paraventricular nucleus receives a sub- tract, the spinal trigeminal nucleus and the
stantial projection from the locus coeruleus marginal zone of the spinal dorsal horn [56,
complex. Other hypothalamic centres receive 94, 225, 241, 254, 320, 487, 721, 949, 1088].
their noradrenergic innervation chiefly from Interestingly, many of these connections reci-
the dorsomedial medullary A2 and lateral procate efferent projections of the locus coer-
tegmental A1 cell groups (see below). uleus. To what extent the fibres of these affer-
The noradrenergic fibres passing to the ent systems are distributed over the entire
cerebellum follow the superior cerebellar pe- complex or specifically address neuronal sub-
duncle and terminate in the central nuclei as populations within the complex is unknown
well as in the cortex. These fibres originate at present.
mainly from the locus coeruleus [1026,1402], Physiological experiments have shown
but some have been reported to arise from that locus coeruleus cells are activated by
the subcoeruleus area [1058]. stressful and particularly by noxious stimuli
A discussion of the internal organisation [230, 1111, 1113, 1114]. However other, non-
of the centres that constitute the locus coeru- threatening stimuli may also influence the ac-
leus complex is beyond the scope of the pres- tivity of locus coeruleus neurons. Thus, it has
ent work. However, in several earlier publica- been demonstrated [354] that in rats locus
tions evidence was presented suggesting that coeruleus cells can be activated by the presen-
only the locus coeruleus as a whole, but also tation of various non-noxious auditory, visu-
its individual neurons innervate widely differ- al or somatosensory stimuli, and that in mon-
ent regions of the central nervous system via keys these cells respond vigorously to com-
collateral branches [1016, 1026, 1375, 1402]. plex arousing stimuli such as preferred food.
More recently, these findings have been sub- Moreover, cardiovascular events such as al-
stantiated by the work of several groups of tered arterial pCO z or pOz levels, and blood
investigators, using mUltiple fluorescent ret- volume or blood pressure variations may
rograde tracers [4, 966, 1147, 1289]. How- markedly affect the locus coeruleus [327,
ever, it has also been experimentally estab- 328]. There is evidence indicating that the ac-
lisl-.ed that, with respect to their efferent pro- tivity of locus coeruleus cells cannot be de-
jections, the locus coeruleus neurons show scribed simply in terms of excitation or inhi-
a considerable degree of regional topograph- bition. Rather, their action selectively en-
ical organisation. Thus, the recent studies of hances or diminishes the effects of the neu-
Loughlin and collaborators [816, 817] have rotransmitters released by other afferents or
shown that the locus coeruleus contains six by intrinsic neurons in their target areas
classes of morphologically different neurons [1416]. On the basis of this and other infor-
and that most of these are clustered within mation [327, 604, 860, 861, 1110, 1111, 1415,
the nucleus and project to particular targets 1417], the generalisation may be madb that
or sets of targets. in the waking state the locus coeruleus exerts
The afferents of the locus coeruleus are an "attention function", i.e. continuously
multifarious and include fibres from the dor- "monitors" the external environment as well
sal prefrontal cortex, the septum, the central as the internal vegetative state for important
amygdaloid nucleus, the bed nucleus of the stimuli/events, and prepares the organism to
stria terminalis, the lateral hypothalamic cope with emergency situations. It has been
area, the periaqueductal grey matter, the ven- suggested that the sole cortical projection to
tral tegmental area, most of the raphe nuclei, the locus coeruleus, which arises from the
prefrontal cortex, may well play an impor- stedt [1402] have been confirmed and ex-
tant role in conveying information about the tended by several more detailed studies.
importance or relevance of complex sensory Fibres arising from these cell groups repor-
events and situations [56]. Finally it may be tedly form a very dense terminal plexus in
mentioned that in several ageing diseases the ventral part of the bed nucleus of the
of the central nervous system, such as stria terminalis and also project to the nucle-
Alzheimer's disease, senile dementia of the us of the diagonal band and the lateral septal
Alzheimer's type and Parkinson's disease, a nucleus [800, 940]. The amygdaloid complex,
significant loss of neurons in the locus coeru- particularly the central nucleus, receives a
leus has been found [847,879,1417]. non-coerulean noradrenergic projection via
Turning now to the ascending efferents of the ventral amygdalofugal pathway [335]. It
the dorsal medullary ( A2) and lateral tegmen- has also been found that the medial preoptic
tal (Ai, A5 and A7) noradrenergic cell groups nucleus, the anterior hypothalamic area, the
(Fig. 159), it should be mentioned first of all supraoptic nucleus and the paraventricular
that, according to the classical mapping nucleus are all densely innervated by the cell
study of Ungerstedt [1402], these cell groups groups of the lower brain stem, but the ven-
collectively give rise to a long, ascending fibre tromedial nucleus is almost devoid of nor-
system, the ventral noradrenergic pathway. adrenergic innervation [942].
According to his description, this pathway Studies using tracer techniques [107, 883,
ascends through the reticular zone of the 1034, 1125, 1181] and particularly analyses
brain stem and continues rostrally, mainly in which the use of retrograde tracers was
within the medial forebrain bundle. Terminal combined with techniques for identification
areas of this pathway have been observed in of labelled catecholaminergic cells [128, 287,
the mesencephalon (the ventrolateral part of 1034, 1206, 1207] have shown that the non-
the substantia grisea centralis and the reticu- coerulean noradrenergic projections to basal
lar formation), the diencephalon (the entire telencephalic and hypothalamic cell masses
hypothalamus, especially the dorsomedial, originate exclusively from the caudal medul-
periventricular, infundibular, supraoptic and lary Al and A2 groups. Thus, it has been
paraventricular nuclei and the internal layer established that the Al group innervates the
of the median eminence), and in the telence- bed nucleus of the stria terminalis and the
phalon (the preoptic area and the bed nucleus medial preoptic area [883], the contralateral
of the stria terminalis). Later studies [624, amygdala [1034], the anterior, lateral and
794, 797, 942, 1333] have shown that the dor- posterior hypothalamic areas [1181], the d~r
sal and ventral noradrenergic bundles as de- sal hypothalamic area, the dorsomedial nu-
scribed by Ungerstedt [1402] cannot be cleus and the median eminence [883] and the
sharply separated; they form one complex, parvocellular and magnocellular portions of
which has been termed the" central tegmen- the paraventricular nucleus as well as the su-
tal tract" [797, 942, 1341] or the "major lon- praoptic nucleus [1206, 1207]. In the magno-
gitudinal catecholamine bundle" [624]. In cellular part of the paraventricular nucleus
this complex, ascending fibres are mixed with and in the supraoptic nucleus, the projections
descending ones, namely the rubro-olivary from the Al group appear to terminate pre-
fibres, to which the name "central tegmental ferentially in areas rich in vasopressinergic
tract" was originally applied. The axons of neurons [1207]. The A2 group has been
the lateral tegmental group feed into it at shown to innervate the medial preoptic area
successive levels through radially coursing, [287, 1034, 1125, 1181], the ipsilateral amyg-
transverse fibres [624]. dala [1034], the anterior, lateral and posterior
As regards the areas of termination of the hypothalamic areas and the dorsomedial nu-
dorsal medullary and lateral tegmental nor- cleus [107, 1181], the nucleus infundibularis
adrenergic cell groups, the findings of Unger- [1125], and the parvocellular portion of the
paraventricular nucleus [82]. Most of the and neuroendocrine responses along various
long, ascending projections from the A1 and routes. Some of these routes may be briefly
A2 cell groups are bilateral with an ipsilateral indicated as follows [112S, 1207]:
predominance.
Apart from the ventral noradrenergic 1. A substantial, though non-catechola-
pathway [1402], or central tegmental tract minergic pathway passes from the solitary
[793, 942, 1341], the dorsal periventricular tract nucleus to the A1 group, and the A1
pathway may also contain long, ascending and A2 groups both project to the AS group.
non-coerulean noradrenergic fibres. It has Loewy and collaborators [806, 808] have
been suggested that in the rat a considerable demonstrated that the AS group projects di-
number of fibres originating from the lower rectly to the sympathetic intermediolateral
medullary cell groups project via this route nucleus in the thoracic cord, and presented
to the periventricular thalamic zone [79S, physiological evidence indicating that the AS
799]. group represents a vasomotor centre.
In addition to long, ascending projections, 2. Alterations in blood pressure and heart
the dorsal medullary and lateral tegmental rate, as well as modifications of respiratory
noradrenergic cell groups give rise to various function, can be induced by stimulation of
propriobulbar projections. The dorsal motor the bed nucleus of the stria terminalis, the
nucleus of the vagus, the facial nucleus, the central amygdaloid nucleus and the medial
motor trigeminal nucleus, the solitary tract preoptic area (see Ricardo and Koh [112S]).
nucleus, the rhombencephalic raphe nuclei All of these telencephalic centres have been
and the parabrachial nuclei are heavily inner- shown to receive a direct projection from the
vated by the non-coerulean noradrenergic A1 and/or A2 groups.
cell groups, whereas the hypoglossal nucleus, 3. The parvocellular portion of the para-
the pontine nuclei, the locus coeruleus and ventricular nucleus may well occupy an anal-
the pontine and medullary reticular forma- ogous position at the diencephalic level. This
tion are less richly innervated from these centre, which is known to influence cardio-
sources. Little is known of the exact localisa- vascular functions, receives direct input from
tion of the cells from which these various the A 1 and A2 groups and projects to various
projections originate. However, there is ex- autonomic centres in the lower brain stem
perimental evidence indicating that the A1 and spinal cord [1207].
group innervates the solitary tract nucleus 4. It has been shown that the peripheral
and other nuclei of the dorsal vagal complex, cardiovascular receptors connected to the va-
the locus coeruleus and the parabrachial nu- gus and glossopharyngeal nerves participate
clei [126, 811, 1207], and that the A2 group in the mechanism of vasopressin (antidiuretic
projects heavily and bilaterally to the solitary hormone) release. The baroreceptors and
tract nucleus [13S7]. atrial stretch receptors inhibit the release
The caudal medullary and lateral tegmen- mechanism, whereas the chemoreceptors
tal noradrenergic cell groups and their effer- probably stimulate vasopressin secretion
ent projections are probably involved in a [112S]. The glossopharyngeal and vagus
wide range of visceral functions, including fibres involved terminate in the solitary tract
cardiovascular control and respiration. The nucleus, and the cells which release vasopres-
nucleus of the solitary tract, in which the A2 sin in the posterior pituitary are located in
group is largely embedded, receives impulses the supraoptic and paraventricular nuclei.
via the vagus and glossopharyngeal nerves, The central route taken by the visceroceptive
from (among others) atrial stretch receptors inputs to the hypothalamus presumably in-
and aortic and carotid baroreceptors and volves: first, the large, non-catecholaminerg-
chemoreceptors. These viscerosensory im- ic pathway which connects the solitary tract
pulses may evoke coordinated autonomic nucleus with the A1 region; and second, the
1 Thalamus, periventricular region

2 ucleus interst.itialis striae terminalis
3 ucleus septi lateralis
4 ucleus para ventricularis, pars magnocellularis
S ucleus para ventricularis, pars parvocellularis 18 Formatio reticularis mesencephali
6 Area lateralis hypothalami 19 Griseum centrale mesencephali
7 Fasciculus telencephalic us medialis 20 Locus coeruleus
8 Fasciculus longitudinalis dorsalis 21 Nuclei parabrachiales
9 ucleus gyri diagonalis 22 Griseum centrale rhombencephali
10 ucleus anterior hypothalami 23 Fasciculus longitudinalis dorsalis
11 ucleus dorsomedialis 24 Longitudinal catecholamine bundle
12 Area caudalis hypothalami 2S Cell group CI
13 ucleus preopticus medialis 26 Cell group C2
14 Nucleus supraopticus 27 Cell group At
15 ucleus infundibularis 28 Cell group A2
t6 Corpus amygdaloideum 29 ucleus dorsalis nervi vagi
17 Eminentia mediana 30 ucleus solitarius
Fig. 159. The ascending reticular system. Ascending projections from the noradrenergic A1 and A2
groups (continuous lines) and the adrenergic C1 and C2 groups (interrupted lines)
direct projection which passes from the Ai lateral pontine tegmentum (Fig. 154C) give
group to the magnocellular parts of the para- rise to an ascending bundle, the dorsal path-
ventricular nucleus and to the supraoptic nu- way of Shute and Lewis [1246]. This fibre
cleus [1207]. system, which projects upon the superior col-
liculus and several thalamic centres, has been
Finally, it should be mentioned that the considered an integral part of the ascending
medullary noradrenergic cell groups have reticular system [905, 907].
been implicated in the control of various an-
terior pituitary hormones, including growth
hormone, luteinizing hormone and ACTH
[942, 1487]. Given the fact that the activity
of the anterior pituitary is regulated by neu-
rosecretory processes in the median emi-
nence, it is noteworthy that noradrenergic
fibres originating from the cell groups men-
tioned are involved in three different projec-
tions leading to that structure, one direct and
two indirect. The direct projection originates
from cell group Ai. The indirect projections
are constituted by fibres arising from the Ai
and A2 groups which terminate in the infun-
dibular nucleus and in the parvocellular divi-
sion of the paraventricular nucleus. Both of
the latter centres project to the median emi-
nence [1207].
As regards the projections ascending from
the adrenergic cell-groups Cl and C2 it has
already been mentioned that fibres originat-
ing from these cell groups join the longitudi-
nal catecholamine bundle, with which they
ascend through the brain stem and enter the
hypothalamus. These fibres give rise to termi-
nal fields in a considerable number of centres,
among which are the griseum centrale of the
rostral rhombencephalon and mesencepha-
lon, the locus coeruleus, some thalamic mid-
line nuclei and the dorsomedial and paraven-
tricular hypothalamic nuclei, the latter of
which shows a high density of terminals [356,
530, 1164]. Little is known about the central
actions of the adrenergic neurons. The dense
innervation of the paraventricular nucleus
suggests that adrenalin might be involved in
oxytocin and vasopressin secretion, and the
innervation of the dorsomedial hypothalamic
nucleus indicates a possible influence on food
intake.
The cholinergic cell-groups Ch5 and Ch6,
which are situated in the rostral part of the
Cerebellum
(Figs. 160-166)
Introduction monosynaptic and multi synaptic nucleo-

olivary pathways (Figs. 163, 164).
Most cerebellar afferents terminate as
Transverse fissures of differing depth divide mossy fibres on granule cells of the cerebellar
the cerebellum into lobes, lobules and folia. cortex, which in turn contact Purkinje cells.
Paramedian sulci, which are deepest in the The axons of the granule cells (the parallel
posterior lobe, separate the vermis from the fibres) course transversely through the super-
hemispheres (Figs. 16-18). The subdivision ficial layer of the cerebellar cortex. Mossy
of the cerebellum has evolved from studies fibres originate from various nuclei in the spi-
of its comparative anatomy and development nal cord and brain stem. The terminations
[141, 762, 763, 764, 1458]. It is covered by of mossy-fibre systems usually include both
a three-layered cortex and contains the deep vermis and hemisphere and are often delim-
cerebellar nuclei within its central white mat- ited by transverse fissures (Fig. 161). This
ter. Afferent systems terminate both in the transverse disposition of the mossy fibre af-
cortex and in the deep nuclei; Purkinje cell ferents is further enhanced by the transverse
axons connect the cortex with the deep cere- orientation of the parallel fibres.
bellar and certain vestibular nuclei; the main Each half of the cerebellar vermis is bila-
efferent pathways take their origin from the terally connected with the vestibular nuclei
deep nuclei. A caudal zone of the cerebellum, and the pontine and bulbar reticular forma-
consisting of the flocculus and the nodule, tion through direct corticovestibular fibres
maintains important afferent and efferent and the medial (fastigial) nucleus with its ef-
connections with the vestibular and oculo- ferent tracts. The cortex of the hemisphere
motor systems. It is known as the vestibulo- projects to the interposed and dentate nuclei
cerebellum (Fig. 162). (Fig. 164). These nuclei which give rise to the
The main neurons of the cerebellar cortex, superior cerebellar peduncle, an important
with their afferent and efferent fibre connec- ascending pathway, which decussates in the
tions, are arranged in two perpendicular mesencephalon and terminates in the red nu-
planes. Purkinje cell dendritic arbors are ori- cleus and thalamus. Through the ventral lat-
ented para sagittally, and the Purkinje cells eral nucleus of the thalamus, the cerebellar
which project to particular deep cerebellar hemisphere is connected with the motor cor-
nuclei are grouped together in parasagittal tex. The influence of the cerebellar hemi-
zones (Fig. 162). Each of these zones, and sphere on motoneurons and interneurons is
the deep cerebellar or vestibular nucleus to therefore mediated through the corticobul-
which it projects, receives a projection from bar, corticospinal, the rubrobulbar and ru-
a subdivision of the contralateral inferior brospinal tracts. Since both these sets of
olive. These olivocerebellar fibres terminate pathways are crossed, lesions of the cerebel-
as climbing fibres on the dendrites of the Pur- lar hemisphere result in ipsilateral loss of mo-
kinje cells. The deep nuclei are reciprocally tor coordination (Figs. 119, 120).
connected with the inferior olive through
Cortex and Central Nuclei the central white matter close to the ventricu-
lar surface. Four nuclei are distinguished
The surface of the cerebellum is covered by (Figs. 103-105): the medial fastigial nucleus,
a cortex that presents a uniform, three- the globose and emboliform nuclei (corre-
layered histological structure throughout its sponding to the posterior and anterior inter-
extent [1037]. These layers are, passing from posed nuclei of lower mammals [1019, 1486]
superficial to deep, the cell-poor molecular and the dentate (lateral cerebellar) nucleus
layer, the monolayered Purkinje layer and [228]. The fastigial nucleus is continuous with
the granular layer. The cerebellar cortex pres- the globose nucleus and the emboliform nu-
ents a lattice structure, with some of its main cleus is connected with the dentate. Purkinje
elements oriented in two perpendicular cell axons pass ventrally between these two
planes [157]. The profusely branching, flat- groups of nuclei, to terminate in the vestibu-
tened dendritic trees of the Purkinje cells ex- lar nuclei.
tend into the molecular layer, where they are
oriented perpendicular to the direction of the
transverse fissures. The deepest layer consists
Types of Afferent and
of numerous small granule cells. The axons
of these cells ascend to the molecular layer Intrinsic Connections
where they dichotomise and run parallel to
the transverse fissures, through the Purkinje Afferent, intrinsic and efferent cerebellar
cell dendritic trees with which they make ex- connections can be distinguished. Cerebellar
citatory synapses. The neurotransmitter of afferents enter the cerebellum from the pons
the parallel fibres is probably glutamate (see via the middle cerebellar peduncle, and from
[997] for a review of the chemical anatomy the spinal cord and the medulla oblongata
of the cerebellar cortex). via the inferior cerebellar peduncle (Fig. 160).
In all layers there are different types of Cerebellar afferents can be subdivided ac-
short axon cells, which close feed-forward or cording to their mode of termination in the
feed-back inhibitory circuits. Golgi cells are cerebellar cortex, into mossy fibres and
located in the granular and Purkinje layers. climbing fibres [200, 1037]. Mossy fibres
Their dendrites extend into the molecular branch profusely and terminate with large
layer and receive excitatory synapses from endings (the mossy fibre rosettes) each of
parallel fibres. Their axonal plexus ramifies which contact a number of granule cell den-
in the granular layer and forms inhibitory, drites. These complex synapses, which also
GABA (gamma-amino-butyric acid)-ergic include the GABA-ergic terminals of the
synapses with the dendrites of the granule Golgi cells, are known as the cerebellar glo-
cells. Basket and stellate cells are found in meruli. The mossy fibres constitute a strongly
the molecular layer. Both are contacted by diverging system. One mossy fibre contacts
parallel fibres and form inhibitory synapses many granule cells; one granule cell synapses
with dendrites of the Purkinje cells. Basket with hundreds of Purkinje cells [962, 1039].
cells are so-called because their axons ramify Mossy fibres originate from different sources
around the primary dendrites and somata of in the pons, the medulla oblongata and the
the Purkinje cells, and terminate in a plume spinal cord. Their neurotransmitters are not
shape around the initial segment of the Pur- known.
kinje-cell axon. GABA has been implicated Climbing fibres are the terminal arborisa-
as the neurotransmitter for both basket and tions of the olivocerebellar fibres [295]. Oli-
stellate cells, but for the stellate cells taurine vocerebellar fibres branch preferentially in
is still a likely candidate [997]. the para sagittal plane [54, 325]. Each olivo-
The deep cerebellar nuclei, to which most cerebellar fibre provides climbing fibres to
of the Purkinje cells project, are located in approximately ten Purkinje cells [926]; each
Cerebellum 223
1 Thalamus
2 ucleu ruber dexter
3 Decussalio pedunculorum cerebellarium
uperiorum
4 Pedunculu cerebellari uperior
5 Vermi cerebelJi
6 Pon
7 Nervu trigeminus
8 Tractus pinocerebellaris anterior
9 Pedunculu cerebellari inferior
10 Pedunculus cerebellari medius
11 Nucleus dentatu
12 Hemispherium cerebelli
13 Nucleus olivaris inferior dexter
14 Tractus olivocerebellari
15 Fibrae arcuatae externae
16 Tractus cuneocerebellaris
17 Tractus spinocerebellaris posterior
Fig. 160. The pedunculi of the cerebellum in a lateral view (3/2 x)

Purkinje cell receives a single climbing fibre. Terminations of Mossy Fibre Systems
The climbing fibres terminate with serial syn-
apses on the smooth, primary-to-tertiary
dendrites of the Purkinje cell. The distal por- Mossy fibres connect motor centres at all lev-
tions of the Purkinje cell dendritic tree, the els of the central nervous system with the
spiny branchlets, receive synaptic contacts cerebellum. Primary afferents project to the
from the parallel fibres. Collaterals of the oli- cerebellum through the posterior spinocere-
vocerebellar fibres have been observed to bellar, central cervical, cuneocerebellar and
contact the inhibitory interneurons in the trigeminocerebellar tracts. Interneurons in
molecular layer and to terminate with mossy the intermediate grey matter of the spinal
fibre rosettes on granule cells [859, 1037], but cord give rise to the anterior and rostral
these contacts have not yet been experimen- spinocerebellar tracts. Subcortical motor
tally verified. Connections of this type may centres such as the vestibular nuclei, the supe-
subserve the long term depression ofPurkinje rior colliculus and the red nucleus are con-
cells which follows the excitation of these ele- nected with the cerebellum, either directly or
ments (the complex spike) by a climbing fibre through mossy fibre systems which relay in
[244]. Studies on the selective uptake and ret- the reticular formation. The pontocerebellar
rograde transport of physiologically inert fibres are the final link in the cerebrocerebel-
D-aspartate strongly suggest that aspartate lar pathway, which also provides the cerebel-
is the neurotransmitter in the climbing fibre lum with an efferent copy of the activity in
pathway [1511]. the corticospinal and corticobulbar tracts.
Other types of afferent, which cannot be Most mossy fibre systems terminate bila-
classified as mossy or climbing fibres, have terally in restricted regions of vermis and
recently been identified in the cerebellar cor- hemispheres, which are sometimes bordered
tex using cytochemical techniques. Seroton- by certain constant interlobular fissures
inergic fibres from the raphe nuclei, and nor- (Fig. 162; [424, 612, 764, 1468]). Root fibres
adrenergic fibres from the locus coeruleus, of the vestibular nerve terminate ipsilaterally
have been found in all layers of the cortex. in the nodule and the adjacent part of the
They appear as a reticulum or as stray fibres uvula. A few terminate in the most ventral
in the granular layer, as a plexus in the Pur- lobules of the anterior lobe and their termi-
kinje layer, and as transversely or radially nation in the flocculus is uncertain [167, 207,
oriented fibres in the molecular layer [5, 121, 706, 713]. Secondary vestibulocerebellar
997]. Fibres with the same appearance and fibres also terminate in the flocculonodular
distribution have recently been shown to ar- and anterior lobes, but their termination is
ise from the hypothalamus [300]. All types bilateral and covers a more extensive area
of afferent fibres provide collaterals to the than the primary vestibulocerebellar projec-
deep nuclei. Intrinsic connections consist of tion [165, 712].
the Purkinje cell axons, which connect the Several spinocerebellar mossy fibre paths
cortex with the deep cerebellar and vestibular have been distinguished (Fig. 161). The pos-
nuclei. Purkinje cell axons give off collaterals terior spinocerebellar and cuneocerebellar
which terminate on neighbouring Purkinje tracts are uncrossed, but the anterior spino-
cells, mainly those lying in the same parasa- cerebellar and rostral spinocerebellar tracts
gittal strip. Purkinje cells are inhibitory, the cross within the spinal cord [1033]. The pos-
majority containing GABA as a neurotrans- terior spinocerebellar tract arises from
mitter and many containing other neuroac- Clarke's thoracic nucleus and from neigh-
tive substances, e.g. the amino acid taurine bouring cells in the dorsal horn and ascends
and the peptide motilin [229, 481, 591, 600, in the superficial part of the posterolateral
853]. funiculus [864,1077,1078, 1266]. It transmits
proprioceptive and exteroceptive signals
Cerebellum 225
from the lower limb. Its forelimb equivalent [5, 324]. Mossy fibres from all subdivisions
is the cuneo cerebellar tract, from the lateral of the spinal trigeminal nucleus terminate in
cuneate nucleus and the adjoining part of the still more dorsal regions [584, 1528, 1541].
medial cuneate nucleus [395,429, 613, 1477]. In the vermis, a mediolateral somatotopic
The anterior spinocerebellar tract arises at pattern prevails. The central cervical spino-
lumbar levels of the cord, from different cell cerebellar projection is mainly represented in
groups in the substantia intermedia and from the most ventral part of the anterior lobe,
the spinal border cells [864, 1077, 1266]. Its where it overlaps with primary and second-
fibres decussate in the ventral commissure ary vestibulocerebellar and (more caudal)
and ascend superficially in the ventrolateral spinocerebellar affere-nt projections [866,
funiculus. According to Lundberg [326, 827], 867].
certain components of the anterior spinocere- Most reticulocerebellar fibres originate
bellar tract originate as collaterals from spi- from the nuclei of the lateral and anterior
nal interneurons. This property distinguishes funiculi, located in the caudal medulla oblon-
the anterior tract from the posterior spino- gata [242]. The nucleus of the lateral funicu-
cerebellar and cuneocerebellar tracts, which lus (lateral reticular nucleus) is located with-
mainly transmit information about external in, and receives terminations from, the as-
events. An equivalent, partially crossed path- cending fibres of the ground bundle of the
way arises from the cervical enlargement lateral funiculus [731] and the descending
[1078, 1514, 1515]. This rostral spinocerebel- fibres of the rubrospinal tract [1441]. The nu-
lar tract should be distinguished from the cleus of the anterior funiculus (paramedian
spinocerebellar fibres which arise from the reticular nucleus) is located adjacent to the
central cervical nucleus located in the sub- medial longitudinal fascicle and the lateral
stantia intermedia of the upper cervical cord vestibulospinal tract, and receives collateral
[272, 866, 867, 1512, 1513]. This nucleus re- terminations from both. Reticulocerebellar
ceives primary input from neck muscles and fibres enter the cerebellum through the resti-
is under a strong vestibular influence. Dor- form body. Those of the nucleus of the ante-
sally located spinocerebellar fibres of the pos- rior funiculus reach the restiform body as ex-
terior tract reach the cerebellum through the ternal arcuate fibres, which pass ventral in
restiform body. The more ventrally located the raphe and through and ventral to the pyr-
spinocerebellar fibres of the anterior, rostral amid [193]. The terminations of the reticulo-
and central cervical tracts pass rostral to the cerebellar fibres overlap with those of the
entrance of the trigeminal nerve and reach spinocerebellar and vestibulocerebellar sys-
the cerebellum via the superior cerebellar pe- tems, but probably spread more widely in the
duncle. hemisphere.
Spinocerebellar and cuneocerebellar fibres Pontocerebellar fibres constitute the larg-
terminate bilaterally in the anterior lobe and est contingent of mossy fibres. They originate
in the pyramis and the adjoining gracile lob- bilaterally from the pontine nuclei [168,
ule of the posterior lobe (Figs. 161,162; [429, 1156], which receive dense, somato-topically
430]). Spinocerebellar projections predomi- organized projections from the entire cere-
nate in the cortex in the deep parts of the bral cortex [122, 173, 409, 1508]. Another
interlobular fissures, the exteroceptive com- contingent of pontocerebellar fibres arises
ponents of the tracts terminating more super- from the pontine tegmental reticular nucleus,
ficially than the proprioceptive components which is located in the ventromedial tegmen-
[323]. A somatotopic pattern of termination tum. It receives projections from the entire
can be discerned in the medial part of the neocortex in addition to a recurrent pathway
hemisphere the pars intermedia [610, 611, from the deep cerebellar nuclei via the de-
612,613]. Here the hindlimb components ter- scending branch of the superior cerebellar pe-
minate ventral to the forelimb components duncle [45, 170, 1380, 1381]. Other afferent
1 Pedunculus cerebri 13 Formatio reticularis myelencephali

2 Nucleus ru ber 14 Tractus cuneocerebellaris
3 Pedunculus cerebellaris superior, ramus descendens 15 Nucleus cuneatus lateralis
4 Tractus ru brospinalis 16 Nucleus pyramidalis
5 Nucleus reticularis tegmenti pontis 17 Tractus spinocerebellaris anterior
6 Nuclei pontis 18 Tractus spinocerebellaris posterior
7 Pedunculus cerebellaris medius 19 Funiculus posterior
8 Pedunculus cerebellaris inferior 20 Cellulae motorae comus anterioris
9 Nuclei centrales cerebelli 21 Substantia intermedia
10 Nervus vestibularis (VIII) 22 Radix dorsalis nervi spinalis
11 Nuclei vestibula res 23 Nucleus thoracicus
12 Lobus Ilocculonodularis 24 "Bordercell " of anterior horn
Fig. 161. The afferent connections of the cerebellum I: distribution of the mossy fibre systems
Cerebellum 227
hemispherium
A
B
...J
...J
W
CD
W
a:
w
u
Fe
"-
UJ
::::> Tu
a:
0
u
Ne
!QOO3~
l~
Li Lingula cere belli 1 Nucleus fastigii
Ce Lobulus centralis 2 Nucleus globosus
Cu Culmen 3 Nucleus emboliformis
De Declive 4 Nucleus dentatus
Fo Folium vermis 5 Nuclei vesti bulares
Tu Tuber vermis
Py Pyramis vermis
Uv Uvula vermis
No Nodulus
Fig. 162. The cerebellar cortex, unfolded in one plane showing the nomenclature of cerebellar lobules
and fissures and the fields of termination of the mossy fibre systems on the left side and the organization
of the corticonuclear and corticovestibular projection on the right side. Pontocerebellar fibres (open
contours), spino- and cuneocerebellar fibres (dotted) and vestibulocerebellar fibres (hatched) . Cortico-
nuclear projection zones and their target nuclei are indicated with the same shadings
projections to the basal and tegmental pon- from the relay cells of all the deep cerebellar
tine nuclei originate from the tectum, the spi- nuclei. This projection is relatively sparse,
nal cord and the dorsal column nuclei. Pon- and it displays roughly the same parasagittal
tocerebellar fibres terminate in all cerebellar organization as the olivocerebellar climbing
lobules, with the possible exception of the fibre projection [425, 876, 1377].
nodule [666]. In the vermis and the pars inter- The termination of the vestibulocerebellar,
media, where their terminations overlap with spinocerebellar and reticulocerebellar mossy
the spinocerebellar projections, they termi- fibres in the granular layer is discontinuous.
nate preferentially in the superficial parts of These mossy fibres terminate in parasagittal-
the lobules. This overlap mainly concerns the ly arranged clusters which alternate with
pontocerebellar projections which transmit empty strips [331, 732, 1210, 1461, 1531]. It
information from the sensorimotor cortex to seems likely that these empty strips are occu-
the "spinal" regions of the cerebellum, and pied by the mossy fibres of other systems
the exteroceptive components of the spino- which terminate in the same region. Ponto-
cerebellar tracts, which also terminate in cerebellar fibres to the hemisphere terminate
more superficial parts of the cortex. The pon- diffusely. Only in the vermis do they show
tocerebellar and spinocerebellar projections a mediolateral periodicity similar to that of
to these lobules display an identical somato- the other mossy fibre systems. Mossy fibre
topic organization [709, 1106, 1107, 1397]. systems issue collaterals to the deep nuclei.
In the vermis, pontocerebellar projections Most of these collaterals seem to come from
preponderate in the declive, folium and afferents arising from the paramedian reticu-
tuber. This part of the vermis corresponds lar nucleus and the pontine tegmental reticu-
to the classical visual projection area of the lar nucleus [329, 393]. In addition, the anteri-
cerebellum [10, 1267]. The pontocerebellar or interposed nucleus receives a collateral
projection to this central part of the vermis projection from the rubrospinal tract [166,
transmits information from the tectum, 575].
which reaches the pontine nuclei through the
tectopontine tract [535] from the striate cor-
tex and from the visual association areas Functional Zones: Organization of the
[122, 398]. In lower mammals dense visual
Olivocerebellar Climbing Fibre System
pontine projections also reach the uvula and
the adjoining para flocculus (the biventral
and the Corticonuclear Projection
lobule of man [185]). Projections to the floc-
culus preferentially originate from the pon- Corticonuclear and corticovestibular fibres
tine tegmental reticular nucleus [393]. In most originate as Purkinje-cell axons from longitu-
of the hemisphere the pontocerebellar projec- dinal zones in the cerebellar cortex. These
tion is the only known afferent system of zones extend perpendicular to the interlobu-
mossy fibres, with the possible exception of lar fissures. The corticonuclear fibres from
the monoaminergic afferents, which have not each of these zones occupy separate compart-
yet been studied in great topographical detail ments in the cerebellar white matter and ter-
[121 ]. minate in a single deep cerebellar or vestibu-
Recently, connections of the limbic system lar nucleus [469, 610, 611, 1457, 1459, 1460,
with the cerebellum have been demonstrated. 1462]. The vermis of the anterior lobe con-
These connections take their origin from the tains the symmetrically disposed A- and B-
hypothalamus and the central grey matter zones. The medial A-zone is connected with
[299, 300, 467]. The fibres ramify and termi- the ipsilateral fastigial nucleus and the medial
nate in all layers of the cortex, mainly in the vestibular nucleus; the lateral B-zone projects
vermal and paravermal areas. Mossy fibres to Deiters' lateral vestibular nucleus. The A-
have also been shown to arise as collaterals zone. extends into the posterior lobe, where
Cerebellum 229
it includes almost the entire vermis. The B- contralateral substantia intermedia and the
zone continues in the lobulus simplex (lobule base of the dorsal horn of the entire spinal
VI of Larsell) immediately behind the prima- cord [53], the dorsal column nuclei and the
ry fissure, but is absent more caudally. The spinal trigeminal nucleus [135, 658, 933, 934].
medial part of the hemisphere, which corre- Purkinje cells of the A-zone (in the medial
sponds to the pars intermedia of Jansen and half of each hemivermis), which project to
Brodal [610, 611, 612], can be subdivided into the fastigial nucleus and the medial vestibular
three C zones, which project to the nucleus nucleus, receive climbing fibres from the cau-
interpositus. The medial (C1) and lateral (C3) dal part of the medial accessory olive. The
zones are connected with the emboliform (or Purkinje cells of the B-zone, which project
anterior interposed) nucleus. The intermedi- to Deiters' lateral vestibular nucleus [40], oc-
ate (C2) zone projects to the globose (or pos- cupy the lateral half of the vermis. They re-
terior interposed) nucleus. The major, lateral ceive their climbing fibres from the caudal
part of the hemisphere contains the two D1- pole of the dorsal accessory olive. Collaterals
and D2-zones, which project to the ventrola- of these olivocerebellar fibres terminate in the
teral, parvocellular and dorsomedial, magno- deep cerebellar or vestibular nucleus which
cellular subdivisions of the dentate nucleus. receives the Purkinje cell axons from their
The C1 and C3 zones are present in the ante- target zone in the cortex, i.e. climbing fibres
rior lobe and the paramedian (gracile) lobule. to the A-zone contribute collaterals to the
The C2-zone and the D1- and D2-zones ex- fastigial nucleus and those to the B-zone to
tend through all the lobules of the cerebellar Deiters' lateral vestibular nucleus.
hemisphere. Longitudinal zones, which are Olivocerebellar fibres to the pars interme-
present in the flocculus and which project to dia take their origin from the rostral halves
the vestibular nuclei and/or the dentate nu- of both accessory olives. Within the pars in-
cleus, replace the D-zones in this part of the termedia they terminate in the three interdigi-
cerebellum. tating C-zones, which in turn project to the
Climbing fibres originate from the contra- anterior and posterior interposed nuclei.
lateral inferior olive and reach the cerebellum These nuclei receive collaterals from the oli-
via the inferior cerebellar peduncle. They vocerebellar fibres that terminate in the pars
show a mediolateral disposition, similar to intermedia. The rostral halves of the dorsal
the intrinsic connections between the cerebel- and medial accessory olives differ in their af-
lar cortex and the deep nuclei (Figs. 163 and ferent connections. The rostral and caudal
164). The corticonuclear and olivocerebellar parts of the dorsal accessory olive share som-
projections will therefore be discussed togeth- esthetic connections from the spinal cord and
er. The terminal fields of the climbing-fibre the brain stem, whereas the rostral half of
paths from subdivisions of the inferior olive the medial accessory olive is dominated by
constitute narrow, parasagittal zones which connections from the rostral central grey
coincide with the corticonuclear projection matter containing the Darkschewitsch' nucle-
zones. The olivocerebellar fibres which termi- us. These connections descend in the medial
nate as climbing fibres in a particular zone tegmental tract, within the mediallongitudi-
also send collaterals to its deep cerebellar tar- nal fascicle [193, 1020].
get nucleus. Climbing fibre zones extend per- Fibres from the principal olive terminate
pendicular to the lobular projection fields of in the D-zones, which occupy the lateral part
the mossy fibre systems [55, 161, 169, 174, of the hemisphere [1155], and in the dentate
175, 261, 322, 394, 452, 453, 1202, 1506]. nucleus, which is the main recipient of the
Fibres from the caudal parts of both accesso- corticonuclear projection from the hemi-
ry olives terminate in the contralateral ver- sphere [394]. The principal olive receives the
mis. These caudal parts of the accessory central tegmental tract from the parvocellu-
olives receive afferent connections from the lar part of the red nucleus [1178, 1304, 1469].
1 Thalamus
2 Nucleus ruber
4 Tractus tegmentalis medialis
5 Tractus tegmental is centralis
6 Decussatio pedundulorum cerebellarium superiorum
8 Tractus spinocerebellaris posterior
9 Nucleus den tatus
11 Nucleus g1obosus
12 Hemisphaerium cerebelli
13 Pedunculus cerebella.ris inferior
14 Tractus olivocerebellaris
16 Amiculum olivae
19 Tractus spino-olivaris
Fig. 163. The olivocerebellar circuits. Position of tracts and nuclei in a dorsal view (5/3 x). The cerebel-
lum was split in the midline and the right half removed
Cerebellum 231
1 Nucleus ruber
4 Pedunculus cerebellaris superior, ramus descendens
5 Tractus tegmentalis medialis
6 Vermis cere belli
7 Hemispherium cerebelli, pars intermedia
8 Hemispherium cerebelli, pars lateralis
11 Nucleus fastigii
12 Nuclei globosus et emboliformis (interpositus)
13 Nucleus dentatus
19 Tractus olivocerebellaris
Nucleus cuneatus lateralis
20 { Nucleus cuneatus medialis
Nucleus gracialis
21 Tractus spino-olivaris
22 {cornu posterius
Substantia intermedia
Fig. 164. The afferent connections of the cerebellum, the climbing fibers and the afferent connections
of the inferior olive
The origin of the projections to the inferior The Vestibulocerebellum

olive from the mesodiencephalic junction is
not restricted to the Darkschewitsch nucleus
and the parvocellular red nucleus [502], but The afferent and efferent connections of the
also includes parts of the central grey matter vestibulocerebellum set this structure apart
and the mesencephalic tegmentum, and visu- from the remainder of the cerebellum. Origi-
al centres such as Cajal's interstitial nucleus nally it was defined as the flocculonodular
of the medial longitudinal fascicle [1031, lobe, the region which receives primary vesti-
1074]. The targets of these descending projec- bular root fibres and projects preferentially
tions to the inferior olive, which also include to the vestibular nuclei [762]. Recently it was
a projection from the superior colliculus [359, shown that the primary vestibular projection
515, 535, 619], also involve those parts of is mostly limited to the nodule [706, 758,
the caudal medial accessory olive which send 759], and that the flocculus is dominated by
climbing fibres to the visual receptive area visuomotor afferents. The common feature
in the vermis of the posterior lobe [10]. Direct of both subdivisions of the vestibulocerebel-
projections from the cerebral cortex to the lum is therefore their efferent projection to
inferior olive are scarce [1176]. Cortical con- the vestibular nuclei, rather than their com-
trol of the olivary complex is mostly effec- mon input. The nodule and flocculus project
tuated through the intermediacy of the red to the vestibular nuclei, but their Purkinje
nucleus and other preolivary centres in the cell axons terminate in different, complemen-
brain stem [37, 38, 39]. Finally it should be tary regions [44, 466, 759, 1456].
pointed out that some parts of the inferior The vestibulocerebellum has been impli-
olive receive a serotoninergic projection from cated in the regulation of compensatory, ves-
the raphe nuclei [689, 1509]. tibulo-ocular, optokinetic and neck reflexes,
The similarity in the topographical organi- all of which pass through the vestibular nu-
zation of the corticonuclear and olivo cere bel- clear complex [600]. The flocculus receives
lar projections is enhanced by the existence information about retinal slip from the nuclei
of reciprocal connections between the deep of the accessory optic system [535, 1254],
cerebellar nuclei and the subdivisions of the through mossy and climbing fibres which
inferior olive from which they receive their originate in the pontine tegmental reticular
collateral projections [228, 440, 658, 1314, nucleus [393] and the dorsal cap of the medial
1378]. Direct, reciprocal nucleo-olivary fibres accessory olive, respectively [18, 392, 758].
originate from a special population of small This makes the flocculus particularly well
GABA-ergic neurons of the deep nuclei [106, equipped to influence eye movements. In-
117, 992]. The projections of the interposed deed, it has been shown to be essential for
and dentate nuclei (to the contralateral red the execution of smooth visual pursuit [1551].
nucleus and the nucleus of Darkschewitsch)
and the central and medial tegmental tracts
form strong feed-back loops which follow the
Efferent Pathways
same topographical pattern.
The main efferent pathways from the cerebel-

lum arise from the deep cerebellar nuclei
(Figs. 165 and 166). The fastigial nucleus
gives rise to the uncinate fascicle, which de-
cussates within the cerebellum, in the cerebel-
lar commissure, and to the uncrossed, direct
fastigiobulbar tract. The uncinate fascicle
passes over the superior cerebellar peduncle
Cerebellum 233
to enter the vestibular nuclei from the lateral which contains the representation of rostral
side. The direct fastigiobulbar fibres enter the body parts [60, 62, 629]. Thalamic projec-
vestibular nuclei from the dorsal side, in the tions from the different cerebellar nuclei ter-
lateral wall of the fourth ventricle. Through minate in alternating sheets or rods, which
these two pathways the fastigial nucleus is show only very little overlap [1369]. Accord-
bilaterally connected with the medial and de- ing to many authors the area of cerebellotha-
scending vestibular nuclei and the medial re- lamic projection also includes rostral parts
ticular formation of the pons and medulla of the ventral lateral nucleus, the adjoining
oblongata [86]. Some fibres of the fastigial ventral anterior nucleus and the ventral me-
nucleus reach the spinal cord [863]. An as- dial nucleus. These differences of interpreta-
cending bundle of fibres, derived from the tion are more apparent than real; they can
uncinate fascicle, courses dorsomedial to the be largely reduced to what Mehler [888] has
superior cerebellar peduncle and subsequent- called "atlas semantics" because the descrip-
ly dorsal to the central tegmental tract, to tions by different authors of the areas to
terminate in the lateral tegmentum of the which the cere bello thalamic fibres are distrib-
mesencephalon and the deep layers of the su- uted are rather similar [43,86, 111,501,659,
perior colliculus. In the diencephalon, its 1280, 1313]. In the caudal part of the ventral
fibres terminate in the intralaminar nuclei lateral nucleus the cerebellar projections
and in the ventral medial and ventral lateral overlap with the terminations of the spino-
nuclei. Some of its fibres recross in the tectal thalamic system [60, 61, 62, 111]. They re-
commissure and in the massa intermedia. main, however, segregated from the palli-
The dentate and interposed nuclei give rise do thalamic and the nigrothalamic projec-
to the superior cerebellar peduncle, which de- tions as well as from and the terminations
cussates in the caudal mesencephalon. Here from the medial lemniscus [587, 629, 659].
fibres of the globose (posterior interposed The cerebellum, especially the fastigial and
nucleus), fastigial and dentate nuclei termi- dentate nuclei, provides one of the main af-
nate in the central grey matter, the deep ferent systems of the intralaminar nuclei.
layers of the superior colliculus and in the Cerebellar fibres terminate in the parafasci-
pretectum [525, 669]. The superior cerebellar cular nucleus and the central lateral nucleus.
peduncle surrounds and traverses the red nu- Through the intralaminar nuclei the cerebel-
cleus. Fibres from the emboli form nucleus lum is connected with the striatum and the
(anterior interposed nucleus) terminate in the cortex of the frontal and parietal lobes. With-
caudal magnocellular part of the red nucleus, in the intralaminar nuclei, the cerebellar af-
fibres from the dentate in its rostral parvocel- ferents overlap with spinothalamic, reticu-
lular part [61, 351]. Beyond the red nucleus, lothalamic, pallidothalamic and nigrotha-
fibres of the superior cerebellar peduncle lamic projections. Connections from the deep
enter the thalamus through the subthalamus, cerebellar nuclei to the oculomotor nuclei
where some of them terminate in the zona come from the cells of the so-called group
incerta. Y [171, 216, 439], an aggregation of cells lo-
Fibres from all of the cerebellar nuclei ter- cated ventral to the dentate nucleus, in the
minate in the thalamus on the caudal part floccular peduncle, which probably belongs
of the ventral lateral nucleus (VLc, [629]), to the superior vestibular nucleus.
which projects to the motor cortex in the Ultimately each hemivermis is bilaterally
frontal lobe. Rostral portions of the cerebel- connected with the spinal cord, through the
lar nuclei project to the lateral part of the fastigial nucleus and the vestibulospinal and
ventral lateral nucleus, which contains the reticulospinal tracts in the medial longitudi-
representation of caudal parts of the body. nal fascicle. Connections between the hemi-
Caudal parts of the cerebellar nuclei project sphere and the cord from the interposed nu-
to medial parts of the ventral lateral nucleus, clei include the superior cerebellar peduncle
ucleus ventralis lateralis

ucleus centralis lateralis } Nuclei
Nucleus centromedianus intra-
ucleus parafascicularis lamina res
ucleus ruber
Fasciculus uncinatus cerebelli,
ramus ascendens
8 Pedunculus cerebellaris superior,
ramus descendens
11 Nucleus fastigii
12 Nucleus dentatus
14 Formatio reticularis myelencephali
Fig. 165. The efferent connections of the cerebellum. Position of tracts and nuclei III a dorsal VieW
(5/3 x). The cerebellum was split in the midline and the right half removed
Cerebellum 235
1 Lobus frontalis
2 Motor cortex (area 4)
3 Lobus parietalis
4 Striatum
6 ucleus ventralis lateralis
7 uclei intralaminares
8 ucleus ruber, pars parvoceUulari
9 Nucleus ruber, pars magnoceUularis
10 Griseum centrale mesencepbali
11 Fasciculus uncinatus cerebeUi, ramus ascendens
14 Pedunculus cerebeUaris superior, ramus descendens
15 Vermis cerebeUi, A-zone
16 Vermis cerebeUi, B-zone
17 Hemispherium cerebeUi, pars intermedia
18 Hemispberium cerebelli,
pars lateraUs
19 ucleus dentatus
20 Nuclei globosus et
emboliformis (interpositus)
21 Nucleus fastigii
23 Nuclei vestibuiares
24 Nodulus
25 Flocculus
26 Nucleus vestibularis latcralis
27 uclei vestibula res medialis,
29
superior el inferior
30 30
28 Formatio reticularis myelencepbali 13
Fig. 166. The efferent connections of the cerebellum

and the rubrospinal tract. The connection, which they terminate. Mossy fibres are there-
starting from the interposed and dentate fore in a position to modulate the activity
nuclei, comprises the superior cerebellar in many different longitudinal cortical zones
peduncle, the ventral lateral nucleus of the and their efferent nuclear channels simulta-
thalamus, the motor cortex and the pyramid. neously.
Because the superior cerebellar peduncle and Climbing fibres, which have a powerful,
the rubrospinal and pyramidal tracts cross, excitatory action on Purkinje cells, but a very
each cerebellar hemisphere is ultimately con- low firing rate, contribute little to the overall
nected with the ipsilateral half of the spinal discharge of the Purkinje cells. The organiza-
cord. tion of the olivocerebellar climbing fibre sys-
tem closely mimics the organization of the
cerebellar efferent system. Each part of the
inferior olive monitors a particular output
Function and Malfunction
channel. Climbing fibres may serve as a level-
of the Cerebellum setting, gating or learning device, which
changes the efficacy of the input to Purkinje
Lesions of the cerebellum result in a loss of cells from mossy and climbing fibres [14, 600,
motor coordination, known as cerebellar 858].
ataxia [403]. There is loss of precision in mus-
cle contraction; movements overshoot or un-
dershoot (dysmetria). Loss of coordination
and dysmetria also affect speech (cerebellar
dysarthria) and ocular movements. Changes
in muscle tone, usually hypotonia, are fre-
quently seen. An explanation of the role the
cerebellum plays in motor coordination is
still lacking. The laterality and localization
of symptoms are usually in accordance with
the known anatomy of the input and output
channels of the cerebellum. Bilateral, axial
ataxia dominates with lesions of the vermis.
The limbs are more affected with damage to
the ipsilateral hemisphere, and vestibular and
oculomotor symptoms result from diseases
that affect the vestibulocerebellum. However,
the correlation between symptoms and the
topography of lesions remains rather crude,
and clinical manifestations of lesions which
damage a single cerebellar zone or a single
deep cerebellar nucleus have not yet been
described.
Mossy fibres provide the main drive for
the coordinating activity of the cerebellum,
but it is not known how an appropriate cere-
bellar output is generated. Mossy fibre sys-
tems generally terminate over wide areas of
the cerebellar cortex, and their mediolateral
spread is further enhanced by the parallel
fibres, the axons of the granular cells on
Thalamocortical and Corticothalamic Connections
(Figs. 167-169)
The Thalamus: Its Parts and Peduncles and along its dorsal surface are termed the
thalamic peduncles. As Fig. 169 shows, these
peduncles are medial to the long corticofugal
The thalamus is a nuclear complex located pathways which descend to the brain stem
in the wall of the diencephalon, caudal to and the spinal cord. The anterior thalamic
the interventricular foramen. During devel- peduncle breaks away from the anterior limb
opment, part of the medial wall of the hemi- of the internal capsule, and its fibres form
sphere adheres to the dorsal thalamic surface a reciprocal connection with the prefrontal
(lamina affixa). Consequently the thalamus and orbitofrontal parts of the cortex and the
becomes located in the floor of the central cingulate gyrus. The superior and posterior
part of the lateral ventricle. Laterally the ex- thalamic peduncles diverge from the posteri-
ternal medullary lamina separates the thala- or limb of the internal capsule, and their
mus from the internal capsule. The reticular fibres form a two-way connection between
nucleus of the thalamus is located between the thalamus and the central parietal and oc-
the external medullary lamina and the inter- cipitotemporal areas. The inferior thalamic
nal capsule. This nucleus is penetrated by peduncle reaches the thalamus at its ventro-
bundles of thalamocortical and corticotha- medial side, medial to the posterior limb of
lamic fibres that become detached from the the internal capsule. It contains fibres con-
internal capsule and enter the thalamus. Ven- necting the thalamus and the orbitofrontal,
tral to the thalamus, the reticular nucleus insular and temporal cortices and the basal
continues into the zona incerta of the subtha- prosencephalon. Amygdalothalamic fibres
lamus. The reticular nucleus and zona incerta enter the inferior thalamic peduncle from the
belong to the ventral thalamus, together with ventral amygdalofugal pathway (cf. Figs.
the ventral nucleus of the lateral geniculate 193, 194 and 201).
body. The main mass of the thalamus is The nomenclature of the thalamic nuclei
known as the dorsal thalamus. The terms in the present work (see also Figs. 24 and
dorsal and ventral thalamus are derived from 25) is derived from Walker [1471], Olszewski
their relative position in the lateral dience- [1028]. Jones' [629] recent monograph on the
phalic wall during development [291, 512]. thalamus contains a complete and authorita-
Fibres from the superior cerebellar peduncle tive review of thalamic morphology and con-
and the globus pallidus (Fig. 173) pass nections. The curved internal medullary la-
through the ventral portion of the external mina divides the thalamus into the medial
medullary lamina between the thalamus and thalamic nucleus and the ventral and lateral
the zona incerta to end in the anterior part groups of thalamic nuclei. Caudally the ven-
of the thalamus. tral group is replaced by the medial genicu-
Corticothalamic and thalamocortical late body, which together with the lateral
fibres that become detached from the corona geniculate body, belongs to the metathala-
radiata and the internal capsule and enter mus. The posterior group of nuclei is located
the thalamus at its rostral and caudal poles in the caudal part of the thalamus, in an area
traversed by the fibres of the medial lemnis- specific nuclei project to the cortex in a more
cus, wedged between the lateral and ventral diffuse manner. They receive afferents from
nuclear groups and the medial geniculate the reticular formation and from many other
body. In its rostral part the internal medul- regions, such as the spinal cord, the sensory
lary lamina encloses the anterior nucleus. relay nuclei, the globus pallidus and the deep
The parataenial and reuniens nuclei are cerebellar nuclei, which project to both non-
small nuclei located adjacent to the medial specific and specific thalamic nuclei. The in-
nucleus in the midline. The lateral nuclear tralaminar nuclei belong to this group, they
group includes the lateral posterior nucleus are characterised by their double projection
and constitutes the massive caudal pole of to the cortex and the striatum. Other thalam-
the thalamus, which is known as the pulvin- ic nuclei with diffuse projections to the cortex
ar. Rostrally it tapers into the lateral dorsal have been identified, such as the magnocellu-
nucleus, which is contained within the split lar part of the medial geniculate body, the
internal medullary lamina, together with the posterior group and, in the rat, the ventral
anterior nucleus. The ventral group of nuclei medial nucleus [507]. It is not known whether
extends more rostrally. It is divided into the the primate thalamus contains an equivalent
ventral anterior, ventral lateral and ventral of the nucleus found in the rodent.
posterior nuclei. The medial, parvocellular Specific and non-specific nuclei also differ
part of the ventral posterior nucleus (Fig. 94) with respect to their laminar projections to
is also known as the ventral medial nucleus. the cerebral cortex [815]. The classical corti-
The posterior nuclear group includes the cell- cal projection of the specific relay nuclei to
dense suprageniculate and limitans nuclei layers III and IV is found only in the primary
and the ill-defined posterior nucleus of the somatosensory, auditory and visual cortices.
thalamus [629]. Most other specific nuclei seem to avoid layer
Apart from myelinated fibres, the internal IV and project to layer III and to the deep
medullary lamina contains the cells of the in- layers V and VI. Thalamocortical projections
tralaminar nuclei. Rostral and caudal groups from specific nuclei are discontinuous, and
of intralaminar nuclei can be distinguished. arranged in patches or strips. The well-
The caudal group includes the prominent known projections of alternate laminae of the
centromedian nucleus, and the parafascicular lateral geniculate to alternate strips in layer
nucleus located medial to it, around the fibre IV of the visual cortex form the centres of
bundles of the habenulo-interpeduncular the ocular dominance columns [566, 570,
tract (the fasciculus retroflexus). The rostral 1507]. Another striking example is the one-
group consists of the central lateral, paracen- to-one relationship maintained in the projec-
tral and central medial nucleus; the latter tion from the representations of the vibrissae
occupies the massa intermedia. in the ventral posterior nucleus of the thala-
mus to the somatosensory cortex in rats and
mice, where the vibrissae are represented by
barrel-like cell condensations in the fourth
Specific and Non-specific Nuclei
cortical layer [1414, 1527].
of the Thalamus Non-specific nuclei project to cortical la-
mina I [627], but the distinction of specific
The nuclei of the thalamus can be subdivided and non-specific nuclei in the thalamus on
into specific and non-specific nuclei. The spe- the basis of their laminar and regional projec-
cific nuclei are relay nuclei which receive well tions is certainly not absolute. More detailed
defined subcortical systems, and often dis- subdivisions of the thalamic nuclei, which ac-
play a high degree of topical organization. count for their diffuse or localised projections
They are reciprocally connected with circum- to single or multiple fields and on the laminar
scribed regions of the cerebral cortex. Non- organization of their thalamocortical projec-
tions, have been proposed for rat [508], cat lar nucleus receives collaterals from both tha-
[832, 833] and primate species [629]. lamocortical and ~orticothalamic fibres [211,
308, 627, 937, 1213]. Many of its cells are
GABA-ergic [558, 1174]. Different sections
of the reticular nucleus are traversed by corti-
Thalamic and Thalamocortical Circuits
cothalamic and thalamocortical fibres from
different thalamic nuclei. This organization
All parts of the cerebral cortex, including the is maintained to some degree in the reciprocal
allocortical fields, are connected with the projection of the reticular nucleus to the spe-
dorsal thalamus. All these connections are cific and non-specific nuclei of the dorsal
strictly reciprocal. The corticothalamic fibres thalamus [1294]. The reticular nucleus, like
arise from the infragranular pyramidal other parts of the ventral thalamus, such as
layers, those terminating in specific nuclei the zona incerta and the ventral nucleus of
originate from layer VI, those terminating in the lateral geniculate body, does not project
non-specific nuclei from layer V [222, 224]. to the cortex. The ventral nucleus of the later-
Intrinsic connections between the nuclei of al geniculate body in primates is represented
the dorsal thalamus [212] have not been con- by the pregeniculate nucleus, which is small
firmed in more recent axonal transport stu- in humans [629].
dies and are probably absent [629].
The subcortical afferents of the dorsal tha-
lamic nuclei terminate on relay cells with
Connections of the Ventral
complex synapses that also contain presyn-
aptic dendrites and axon terminals of pre- Thalamic Nuclei
sumed interneurons. The presence of local-
circuit neurons in the thalamic nuclei has The connections of the ventral nuclei of the
been disputed. Using the Golgi method dorsal thalamus are discussed with the gener-
[1379] and after large inje~tions in the cortex al sensory and taste systems, the cerebellum
with retrograde tracers [1483], the percentage and the extrapyramidal motor system. The
of interneurons in the thalamic nuclei has spinothalamic and lateral trigeminothalamic
been estimated to be 20%-30%. The pres- tracts terminate within the ventral posterior
ence of inhibitory GABA (gamma-amino bu- nucleus and the adjoining part of the ventral
tyric acid)-ergic interneurons in the dorsal lateral nucleus. The projections of the dorsal
thalamus has also been studied, using differ- column nuclei and the principal sensory nu-
ent immunocytochemical or uptake tech- cleus of the trigeminal nerve are restricted
niques (see Jones [629] and Nieuwenhuys to the lateral and the medial parts, respective-
[997] for reviews). The existence of such ele- ly, of the ventral posterior nucleus. They ter-
ments in rat was first denied [558, 1174], but minate in a series of parallel laminae, each
later they were found to be present in the representing a specific region of the body,
rat lateral geniculate body [350], and in the but including different sensory modalities
ventral posterior nucleus of cats and lower along their anteroposterior axis. Within the
primates [1066] using a combination ofretro- ventral posterior nucleus the projections of
grade labelling of the relay cells and an anti- the spinothalamic and lateral trigeminotha-
body against GABA to stain the interneurons larnic tracts are somatotopically organized.
[936]. These inhibitory interneurons are sup- They overlap with the termination of the me-
posedly involved in the postsynaptic inhibi- dial lemniscus [110, 111, 192, 263, 629, 635,
tion which usually follows the excitation of 852, 887]. Uncrossed trigeminothalamic
thalamic relay cells by afferent volleys. How- fibres from the principal sensory nucleus ter-
ever, this effect could also be mediated by minate in the lateral portion of the postero-
cells of the reticular nucleus [629]. The reticu- medial ventral nucleus, medial to the gusta-
tory fibres from the medial parabrachial Connections of the Anterior Nucleus
nucleus [1105, 1538]. and the Lateral Dorsal Nucleus
The ventral posterior nucleus is reciprocal- of the Thalamus
ly connected with areas 3b, 2 and 1 of the
first somatosensory area in the postcentral
gyrus and the second somatosensory area in The anterior nucleus of the thalamus can be
the parietal operculum [362, 629, 635, 790, subdivided into the anteromedial, anterodor-
1094, 1504]. The ipsilateral trigeminocortical sal and anteroventral nuclei. It is enclosed
pathway projects to the ventral part of the within the split internal medullary lamina, to-
first somatosensory area through the medial gether with the lateral dorsal nucleus. The
parvocellular part of the ventral posterome- connections of the anterior nucleus and the
dial nucleus. This part of the nucleus also lateral dorsal nucleus with the limbic cortex
serves as a relay in the gustatory projection are rather similar [629].
to the insular operculum and the orbitofron- The anterior nucleus receives the mamil-
tal cortex [629, 1105]. lothalamic tract. The lateral mamillary nucle-
The posterior part of the primate ventral us projects bilaterally to the anterodorsal nu-
lateral nucleus (VLp, [629]) receives fibres cleus, the medial mamillary nucleus to the
from the central cerebellar nuclei [62]. Along anteromedial and anteroventral nuclei [746,
the border with the ventral posterior nucleus 1433]. There are no projections from the ma-
the cerebellar projections overlap with the milIary body to the lateral dorsal nucleus.
terminations of the spinothalamic and lateral This nucleus receives a projection from the
trigeminothalamic tracts and with the vesti- pretectum [1131, 1134]. The anterior and lat-
bulothalamic projection [757]. Pallidotha- eral dorsal nuclei are reciprocally connected
lamic fibres terminate rostral to the cerebellar with the limbic cortex of the cingulate gyrus,
afferents [297, 480, 501, 587, 737, 972, 1053] the retrosplenial area and the pre- and para-
in the anterior part of the ventral lateral nu- subiculum [655, 1133, 1450, 1452]. Although
cleus (VLa, [629]). A more extensive overlap each of the anterior subnuclei has been relat-
of the pallidal and cerebellar projections has ed to a special field within the limbic cortex
been described [219, 891]. Projections from [1149], their projections appear to be rather
the pars reticulata of the substantia nigra diffuse and overlapping.
[219, 586] occupy the ventromedial part of
the ventral lateral nucleus. The posterior part
of the ventral lateral nucleus projects to corti-
Connections of the Medial Nucleus
cal area 4; the anterior part of the nucleus
of the Thalamus
projects to part of area 6, including the sup-
plementary motor area [629, 1215, 1302].
According to Jones [629], the ventral ante- The medial thalamic nucleus is surrounded
rior nucleus is clearly defined in primates but by the internal medullary lamina with its nu-
difficult to distinguish in lower mammals. Its clei. Medial and ventromedial to it are the
subcortical afferents are still unknown. It is nuclei of the midline (rhomboid nucleus, nu-
reciprocally connected with the frontal eye cleus reuniens and parataenial nucleus). The
field (area 8) and the prefrontal cortex [59, medial nucleus can be subdivided into medial
416,684, 1281]. Some of the corticothalamic magnocellular and lateral parvocellular parts
connections of the prefrontal cortex with the [1028]. The large cells located on the border
ventral anterior nucleus are bilateral [412]. of the lateral division of the nucleus with the
internal medullary lamina (pars paralamel-
laris) belong to the central lateral nucleus
[629]. The medial, magnocellular part of the
medial nucleus is reciprocally connected with
(medial prefrontal and orbitofrontal) olfacto- possibly from other pathways. It maintains
ry-related areas [12, 1537]. It also receives diffuse connections with the auditory fields
fibres from the amygdala [7] and from the and regions beyond. Its thalamocortical
entorhinal and perirhinal cortices and the fibres terminate in layer I, similar to other
cortex of the temporal pole. Direct projec- non-specific nuclei.
tions from the olfactory cortex to the medial
nucleus [718] are few in primates [1169]. The
lateral, parvocellular part of the medial nu-
Connections of the Posterior Nuclei
cleus is connected with the frontal eye field
(area 8) and the prefrontal cortex [12, 416, of the Thalamus
572, 1169, 1215, 1364]. The lateral part of
the nucleus receives afferent connections The posterior nuclei are located between the
from the superior colliculus, substantia nigra, medial geniculate body, the ventral posterior
vestibular nuclei and midbrain tegmental nucleus and the nuclei of the lateral group.
fields (586, 736, 1439]. The medial nucleus The medial part receives terminals of the
receives a projection from the ventral palli- spinothalamic and lateral trigeminothalamic
dum [461, 496, 1169]. Through this pathway, tracts (see Figs. 128 and 132), the lateral part
the ventral striatum (nucleus accumbens) is receives those of the inferior colliculus [727,
connected with the prefrontal cortex. Thala- 886, 1432]. The posterior nucleus is con-
mostriatal projections to the ventral striatum nected with the retroinsular cortex surround-
originate from the parataenial nucleus [448, ing S2 and the adjacent insular cortex [190,
1329]. 630,960].
Connections of the Medial Connections of the Lateral

Geniculate Body Geniculate Body
The medial geniculate body consists of a ven- The dorsal part of the lateral geniculate body
tral and a dorsal subnucleus and a medial is a laminated nucleus. In humans, the nucle-
magnocellular division, located between the us consists of two ventrally placed magnocel-
ventral nucleus and the medial lemniscus. lular laminae (1 and 2) and four parvocellular
The ventral nucleus receives the main path- laminae (3-6; [517]). Laminae 1, 4 and 6 re-
way from the central nucleus of the inferior ceive fibres from the contralateral, laminae
colliculus and projects to the primary audito- 2, 3 and 5 from the ipsilateral eye. The basic
ry cortex. The ventral nucleus and its target primate pattern of the lateral geniculate body
area in the temporal operculum contain a consists of two magnocellular and two parvo-
tonotopic representation. The morphological cellular laminae [653]. Each of these laminae
substrate of this representation is the lamellar contains a full representation of the contra-
organization of incoming fibres and relay lateral visual hemifield. In man and other pri-
cells, first described by Morest [947]. The mates the parvocellular laminae are split, and
dorsal nucleus receives fibres from the peri- interdigitate to constitute four parvocellular
central nucleus of the inferior colliculus [201, layers. As a consequence, the representations
1432]. The dorsal nucleus is connected with of the visual hemifield in the parvocellular
the auditory association cortex in the tempo- laminae are incomplete and complementary
ral plane and the superior temporal gyrus to the representations in the other parvocel-
[190,629,908,1275]. The medial magnocellu- lular laminae carrying information from the
lar division receives collaterals from the me- same eye [250].
dial lemniscus, the spinothalamic tract and
Gyri orbitales
Gyri len">n,"""'~
Gyri oo,;elllle9
1 Gyrus cinguli
2 Corpus striatum 10 Nucleus lateralis posterior
Gyri occipitales
3 Globus paUidus 11 Nucleus centromedianus
4 Nucleus anterior thalami 12 Nucleus parafascicularis
5 Nucleus medialis thalami 13 Pulvinar thalami, pars anterior
6 Nucleus ventralis anterior 14 Pulvinar thalami, pars media lis
7 Nucleus ventralis lateralis 15 Pulvinar thalami, pars lateralis
8 Nucleus ventralis posterior 16 Corpus geniculatum laterale
9 Nucleus ventralis posterior, pars parvocellularis 17 Corpus geniculatum mediale
Fig. 167. Connections between the thalamic nuclei and the cerebral cortex I: diagrammatic horizontal
section. Left: corticothalamic projections; right: thalamocortical projections
15
Fig. 168A, B. Connections between the thalamic nuclei and the cerebral cortex II: cortical projection
areas of the thalamic nuclei . A lateral view ; B medial view. The shading is the same as that used
in Figure 167. For explanation of numbers see also Figure 167
It was pointed out in the discussion of the lobe may therefore lead to an upper quadrant
visual system in the chapter on special senso- anopsia [84]. Within the optic radiation, the
ry systems, that ex-like ganglion cells of the thalamocortical fibres are separated from the
retina, which give rise to the rapidly conduct- more medially located corticothalamic fibres
ing Y -system project preferentially to the [1525].
magnocellular laminae, whereas fJ-cells which
belong to the slowly conducting X-system
project to the parvt>cellular laminae.
Connections of the Lateral
The dorsal part of the lateral geniculate
body is reciprocally connected with the pri- Thalamic Nuclei
mary visual cortex (area 17). Thalamocorti-
cal fibres from the parvocellular laminae ter- The lateral group of thalamic nuclei consists
minate in deep and superficial sub-layers of of the lateral posterior and lateral dorsal nu-
layer IV. The magnocellular laminae project clei and the pulvinar. The connections of the
to an intermediary sublayer within layer IV. lateral dorsal nucleus have been discussed
In addition, magnocellular layers project to with the anterior nucleus. The pulvinar is
layer I and to the border zone of layers V large in primates and can be subdivided into
and VI [499, 570, 654, 1067, 1119, 1507]. In four sub-nuclei: the anterior, medial, lateral
primates, the dorsal part of the lateral geni- and inferior nuclei of the pulvinar [1028]. The
culate body does not project beyond area 17, nuclei of the lateral group are reciprocally
but in lower mammals this nucleus projects connected with the large expanse of associa-
both to area 17 and to the visual association tion cortex in the posterior parietal, occipital
cortex of areas 18 and 19 (see Rodieck [1139] and temporal lobes [59, 102, 190, 205, 274,
for a review). 574, 642, 663, 857, 911, 1062, 1282, 1391,
The projection from one eye, through the 1392, 1399, 1404, 1485, 1542]. The lateral
appropriate laminae to the visual cortex is posterior nucleus and the anterior pulvinar
discontinuous in most primates and orga- are connected with the areas 5 and 7 in the
nized in ocular dominance strips in layer IV. superior and inferior parietal lobules. The
These strips form the basis for the ocular projection of the lateral posterior nucleus is
dominance columns that extend perpendicu- situated rostral to that of the anterior pulvin-
lar to the surface through all cortical layers. ar. The projections from the lateral and infe-
The columns, which have a width of approxi- rior nuclei of the pulvinar include the cortex
mately 0.5 mm, branch off in a roughly par- at the parieto-occipital junction, the circum-
allel manner from the representation of the striate areas 18 and 19 and the primary visual
horizontal meridian. Their width is constant area (area 17). Fibres from the pulvinar ter-
throughout the representations of the central minate in layer I of area 17, and in layers
and peripheral parts of the visual field [565, II and IV of the circumstriate cortex. These
566, 567, 568, 569]. connections are reciprocated by projections
The connection between the lateral geni- from pyramidal cells in layer V of area 17
culate body and the visual cortex is known and layer VI of the circumstriate cortex [104,
as the optic radiation. The ventral part of 205,274,1021,1022,1067,1119,1391,1392,
the optic radiation consists of fibres to the 1485].
portion of the visual cortex located below The projection of the inferior pulvinar ex-
the calcarine sulcus; this portion represents tends into the inferotemporal visual associa-
the upper quadrant of the visual field (see tion cortex. The medial pulvinar projects to
Figs. 144, 145). The fibres curve rostrally in the superior temporal gyrus, the dorsal wall
the lateral wall of the inferior horn of the of the superior temporal sulcus and the tem-
lateral ventricle, before turning towards the poral pole. In addition separate cell popula-
occipital lobe. Deep lesions of the temporal tions in the medial pulvinar project to the
I
1 Gyrus postcentralis (areae 2, 1, 3)
2 Gyrus precentralis (area 4)
3 Gyri frontales (areae 6, 8)
4 Sulcus centralis 13 Putamen
5 Nucleus caudatus 14 Peduncul us thalami inferior
6 Tractus pyramidalis 15 Ansa peduncularis
7 Pedunculus thalami superior 16 Tractus temporopontinus
Capsula interna, (capsula intema, pars su blentiformis)
8 Pedunculus thalami posterior
crus posterius 17 Tractus occipitopontinus
9 Tractus parietopontinus
10 Fibrae corticotegmentales (capsula intema, pars retrolentiformis)
11 Tractus frontopontinus Capsula interna, 18 Radiatio optica
12 Pedunculus thalami anterior crus anterius 19 Stratum sagi ttale
Fig. 169. Lateral view of the thalamic peduncles and the internal capsule (1 /1 x). The proximal part
of the corticopontine and corticospinal fibre bundles has been removed to show the thalamic peduncles,
which are illustrated as compact bundles for reasons of clarity. The origin of the pyramidal tract
in the cerebral cortex has been emphasised (cf. Fig. 170)
frontal eye field [59, 574] and limbic areas The main afferents of the intralaminar nu-
[75]. clei come from the spinothalamic and lateral
The inferior and lateral nuclei of the pul- trigeminothalamic tracts, the cerebellum and
vinar receive important projections from the the globus pallidus. These connections are
superficial layers of the superior colliculus described in other chapters. The projections
and the pretectum. Projections from the from the globus pallidus chiefly involve the
deep, non-visual layers of the superior colli- centromedian and parafascicular nuclei [62,
culus terminate in the medial pulvinar [102, 338, 891, 1054]. The cerebellar and spinotha-
104,572,829,1390]. Direct connections from lamic afferents terminate more rostrally, in
the retina to the inferior pulvinar have been the central, lateral and more rostral intrala-
described [500]. minar rlUclei [188, 263, 852,935]. Fibres from
The extrageniculate pathway from the su- the bulbar reticular formation, which have
perior colliculus via the inferior and lateral been claimed to enforce the diffuse spinotha-
pulvinar to the visual association areas is re- lamic projection to the intralaminar nuclei
sponsible for residual visual discrimination [134], do not appear to be very numerous
after lesions of the striate cortex, as has been [886]. The mesencephalic reticular formation,
demonstrated both in experimental animals including the cuneiform nucleus, gives rise
and in humans [226, 1217, 1490]. to an important projection to the intralamin-
ar nuclei [152, 321].
Connections of the Intralaminar Nuclei
The intralaminar nuclei are characterised by

double projections, a diffuse projection to the
cortex and a projection to the striatum. The
striatal projection was first demonstrated in
human material [1453] and later in experi-
mental animals [313, 314, 990, 1100, 1101].
In cats, both the anterior and posterior group
of intralaminar nuclei project to the entire
striatum in an overlapping fashion [91, 616,
881,1161]. The cortical projections of the in-
tralaminar nuclei are diffuse, but are still con-
centrated in certain areas [508, 637]. A few
cells within the intralaminar nuclei give rise
to branching axons which terminate both in
the cortex and the striatum [835, 1200]. The
thalamocortical connections of the intra-
laminar nuclei are reciprocated by cortico-
thalamic projections. The prefrontal cortex,
cingulate gyrus and premotor area are con-
nected with the anterior group of intralamin-
ar nuclei. The motor and parietal cortices
project to the centromedian and central later-
al nuclei. There are fewer connections with
the occipital and temporal lobes [642, 734,
73~835,881, 1076].
Motor Systems
Long Corticofugal Pathways The majority of the fibres of the pyramidal

tract take their origin from the motor cortex
(area 4; [176]) and caudal premotor cortex
Introduction (area 6), but 20% stem from the somatosen-
sory and parietal cortices [609, 1408, 1409).
Projection fibres from deep pyramidal cell The largest fibres of the pyramidal tract are
layers of the cerebral cortex (cortical layers the axons of the giant pyramidal cells (or
V and VI; [16, 114, 221, 222, 223, 363, 641, Betz's cells) in the motor cortex. In primates
665,667,668,671,672,778,826,1277,1519, they constitute the fastest fibres of the pyra-
1523]) terminate in an orderly manner in the midal tract, which terminate directly on the
striatum, thalamus, brain stem and spinal motoneurons [1085, 1086]. The fibres from
cord. In the depths of the hemisphere these the somatosensory cortex terminate on relay
fibres constitute, together with the thalamo- nuclei in long ascending sensory pathways,
cortical fibres, the corona radiata and the in- including the ventral posterior thalamic nu-
ternal capsule (Fig. 170). Caudal to the optic cleus, the nucleus princeps of the trigeminal
tract they come to lie on the ventral surface nerve, the dorsal funiculus nuclei, the spinal
of the brain stem as the cerebral peduncle. nucleus of the trigeminal nerve and the dorsal
Most of the fibres of the cerebral peduncle horn (see Fig. 134 and the section on general
are thin, but its middle division is distin- sensory systems and taste). The fibres from
guished "by its content of thicker myelinated the medial, frontopontine division of the ce-
fibres. About 10% of these fibres are over rebral peduncle take their origin from the
4 !lm in diameter, the thickest fibres reaching premotor and prefrontal cortex [734, 1508].
a calibre of 20 !lm [761, 1444]. The lateral division of the cerebral peduncle
After having passed the cerebral peduncle contains fibres from the parietal association
the corticofugal fibres enter the pons and cortex, with smaller contributions from the
split in smaller bundles. The medial and later- temporal and occipital lobes [673, 761].
al corticopontine divisions of the cerebral pe- During their course along the brainstem
duncle terminate on the pontine nuclei, and and the cord many fibres detach from the
the middle division of the peduncle con- long corticofugal system, some as collaterals
tinues, caudal to the pons, as the pyramidal of ongoing fibres. Fibres to the pretectum
tract (Fig. 174). The majority of the pyrami- and the superior colliculus leave the internal
dal tract fibres decussates at the bulbospinal capsule, to course through the dorsal thala-
junction to descend in the dorsolateral funi- mus. Other fibres to the tegmentum and the
culus of the cord as the lateral pyramidal tectum mesencephali leave the cerebral pe-
tract. A varying proportion of the fibres of duncle to pass through or along the substan-
the pyramidal tract does not decussate, but tia nigra. An important contingent of fibres
descends in the wall of the anterior fissure detaches from the middle division of the cere-
of the cord as the anterior pyramidal tract bral peduncle and can be followed caudally
to midthoracic levels. through the area of the medial lemniscus
1 Gyrus postcen tralis (areae 2, 1, 3)

2 Sulcus ccntralis
4 Premotor cortex (area 6)
9 Tractus parietopontinus
10 Tractus oecipitopontinus
13 Putamen
14 Globus palJidus
15 Substantia nigra
16 Tractus temporopontinus
17 Nucleus dentatus sinister
19 Pons
Fig. 170. Pictorial survey of the origin of the pyramidal tract in the cerebral cortex and of the long
corticofugal system in a lateral view (1 /1 x). The brain stem and the cerebellum have been cut III
the median plane and the right half has been removed, with the exception of the pyramidal tract
Motor Systems 249
1 ucleus caudatus
2 ucleus ventralis laterali
3 Fasciculus thalamicus
4 Fibrae strionigraIes
5 Putamen
6 Globus pallid us, pa rs lateral i
7 Globus pallid us, pa rs medi alis
8 Fa cicuJu lenticularis
9 ucleu reticularis thalami
10 Zona incerta
11 ucleu ruber
14 Substantia nigra
16 Pons
Fig. 171. The nuclei and fibre bundles of the so-called extrapyramidal system in a lateral view (12/5 x).
Of the fibres originating from the occipital, removed part of the lentiform nucleus, only the ansa
lenticularis is represented
1 Corpus callosum
2 Corona radiata
4 Capsula interna
5 ucleus ventralis lateralis
7 Putamen
9 Globus paUidus, pars medialis
10 Nucleus ruber
11 Capsula interna, pars retrolentiformis 16 Pons
12 Cauda nuclei caudati 17 Pyramis
13 Tractus temporopontinus) 18 Decussalio pyramidum
14 Tractus pyramidalis Pedunculus cerebri 19 Tractus pyramidalis lateralis
15 Tractus frontopontinus 20 Tractus pyramidalis anterior
Fig. 172. The long corticofugal fibre system in a frontal view (6/5 x). The plane of the section shown
in this figure coincides with the long axis of the brain stem
Motor Systems 251
1 Corona radiata 11 Nucleus subthalamicus

2 Corpus nuclei caudati 12 Substantia nigra
3 Putamen 13 Nucleus parafascicularis
4 Fibrae strionigrales 14 Ansa lenticularis
5 Capsula interna, crus posterius 15 Nucleus ruber
6 Nucleus reticularis thalami 16 Tractus opticus
7 Nucleus centromedianus 17 Capsula interna, pars sublentiformis
8 Fasciculus thalamicus 18 Cauda nuclei caudati
9 Zona incerta 19 Pedunculus cerebri
10 Fasciculus lenticularis 20 Pons
Fig. 173. The nuclei and fibres of the so-called extrapyramidal system in a frontal view (12/5 x).
The fibres originating from the removed frontal part of the lentiform nucleus are not illustrated
with the exception of the ansa lenticularis. The plane of the section is the same as in Figure 172
GYRI FRONTALES
'ea 3,
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V)
w
-'
<
cr
o
0..
~
W
fo-
o
....
0:
<3
o
o
o
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~ ~
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1 Nucleus caudatus
2 Nucleus ventralis lateralis
3 Nucleus mediilis thalami 16 ucleus inlerposiluS
4 Nuclei intralaminares thalami 17 ucleus den latus
5 Putamen 18 Formalio relicularis
6 Globus pallid us, pars medialis pontis
7 Colliculus superior a 19 Purkinje-cells
8 Nucleus subthaJamicus 20 Granular cells
9 Nucleus ruber, pars parvoceUularis 21 Tractus rubrospinalis
10 ucleus ruber, pars magnoceUularis 22 Tractus lectospinalis
11 Tractus parietooccipitotemporopontinus 23 Tractus pyramidalis
12 Tractus fronloponlinus lateralis
13 Tractus pyramidalis 24 Substantia intermedia
14 uclei ponlis 25 Cellulae motoriae comus
15 Traclus pyramidalis anlerior anterioris
Fig. 174. The neuronal connections of the pyramidal system. Feedback pathways are in open contours
Motor Systems 253
[761] where it enters the medial reticular for- located in the dorsolateral part of the inter-
mation or rejoins the pyramidal tract [744]. mediate zone of the cord and the lateral part
Corticobulbar fibres detach from the bundles of the lateral (parvocellular) reticular forma-
in the pons and the pyramid. The corticofu- tion of the medulla oblongata and the pons.
gal system presents many species variations The axons of these lateral interneurons are
in the level of its major decussation, in its generally short. In the cord they are located
localization, and in the extent of its descent a few segments rostral to their termination
in the spinal cord [1443]. In humans interindi- in the ipsilateral anterior horn. Interneurons
vidual variations are present with respect to projecting to the ipsilateral ventral part of
the completeness of the pyramidal decussa- the facial nucleus are located in the medulla
tion, which on occassion may have failed to oblongata, with those projecting to the ipsi-
develop [822, 1442] and to the occurrence of lateral hypoglossal nucleus more caudally,
aberrant fascicles, such as the circumolivary and those projecting bilaterally to the motor
bundle, which detaches from the pyramid nucleus of the trigeminal nerve more rostrally
and turns back to terminate on the pontobul- in the lateral reticular formation.
bar body [1319]. Descending fibres of the rUbrobulbar and
rubrospinal tract from the contralateral red
nucleus terminate on interneurons in the lat-
Medial and Lateral Motor Systems eral reticular formation and the dorsolateral
intermediate zone of the spinal cord and di-
The main terminations of the long corticofu- rectly on motoneurons of the nucleus of the
gal system in the brain stem and spinal cord facial nerve and, in the cat, on a small moto-
include the tectum, the basal pontine nuclei, neuronal cell group in the dorsolateral part
the reticular formation and the intermediate of the anterior horn at C8 and T1 [533]. The
zone of the spinal cord, the somatosensory rubrobulbar and rubrospinal tract is somato-
relay nuclei and the motor nuclei. Most of topically organized. Cells projecting to the
these cortical connections are mentioned in facial nerve nucleus occupy the dorsal part
the chapters on the somatosensory system, of the red nucleus, and the cells projecting
the special sensory systems and the cerebel- to the cervical and lumbar enlargements are
lum. The direct and indirect cortical projec- located in its intermediate and ventrolateral
tions through interneurons and brain stem parts respectively [544, 1013, 1092]. Rubro-
pathways to the motor nuclei of the brain spinal fibres which distribute branches to
stem and the spinal cord were extensively both enlargements are few [576, 1239]. The
studied and summarized by Kuypers [744, rubrospinal system has an excitatory influ-
745]. His concept, according to which the ence on contralateral flexor motoneurons
motor system can be subdivided into separate and inhibits contralateral extensors [546,547,
medial and lateral systems is based on the 1198]. Lesions of the rubrospinal tract result
organization of the motoneuronal cell in motor deficits in the execution of indepen-
groups, their interneurons and the descend- dent movements of the limbs, especially of
ing pathways (Fig. 175). their distal parts [768].
The lateral system (Fig. 175A) includes The motoneurons which belong to the me-
the lateral motor column of the spinal cord, dial motor system (Fig. 175 B) innervate
namely the motoneurons innervating distal proximal muscles. They include the medial
muscles of the limbs [364, 1142, 1296] and motor column of the anterior horn, which
several somatomotor cranial nerve nuclei, in innervates the axial muscles and the somato-
particular the ventral (in lower mammals lat- motor nuclei of the Vth, the dorsal (periorbit-
eral) part of the facial nucleus, which inner- al) part of the VIIth and the XIIth cranial
vates the perioral muscles [260, 738]. Inter- nerves, which subserve bilateral movements.
neurons innervating lateral motoneurons are Interneurons belonging to the medial system
A B
Motor Systems 255

3 Frontal eye field (area 8)
4 Tractus corticobulbaris et corticospinalis
5 Nucleus ruber
6 Fibrae hypothalamofugales et amygdalofugales
9 Locus coeruleus
14 Formatio reticularis medialis pontis
15 Formatio reticularis medialis bulbaris
16 Tractus reticulospinalis
17
18
19
Nuclei vestibulares
Nucleus vestibularis lateralis
Tractus vestibulospinal is medialis
,r-'"
\ .
21 Nucleus raphes pallidus \
22 Nucleus nervi facialis, pars dorsalis
23 Nucleus nervi facialis, pars ventralis \
24
25
Nucleus ambiguus
Nucleus nervi facialis
/
26
27
Decussatio pyramidum
Tractus pyramidalis lateralis
/
28 Tractus pyramidalis anterior I
29 Substantia intermedia, pars dorsolateral is
30 Substantia intermedia, pars ventromedialis \
31
32
Cellulae motoriae mediales
Cellulae motoriae laterales \
33
34
Fasciculus longitudinalis medialis
Tractus raphespinalis
\
35 Tractus coeruleospinalis \
\
\
Fig. 17SA-C. The neuronal connections of the lateral motor system (A), the medial motor system
(B) and the motoneuronal connections of the locus coeruleus, the raphe nuclei and the medial reticular
formation (C)
have long axons and often distribute bilater- limb and body movements. Animals with
ally. In the cord they are located in the ven- combined lesions of the pyramidal tracts and
tromedial part of the intermediate zone and the medial longitudinal fascicle failed to right
in the brain stem they occupy the medial part and were unable to stabilize their arm in fine
of the lateral reticular formation. They pro- movements of the hand and the fingers,
ject bilaterally to medially located anterior which they were able to execute normally
horn cells and to the nuclei of the XIIth, [768].
VIIth and Vth cranial nerves [538, 541, 545].
Several descending brain stem pathways,
which terminate on these interneurons, be- Reticulospinal, Rapbespinal
long to the medial motor system. All of these and Coeruleospinal Systems
pathways descend in or near the medial lon-
gitudinal fascicle and the ventral funiculus Until recently the medial reticulospinal fibres
of the cord. The medial reticulospinal tract from the medulla oblongata and the pons
[163, 193, 539, 1015] arises from the medial were considered to belong to a single system
reticular formation of the pons. It descends with its main termination in the medial part
bilaterally in the medial longitudinal fascicle of the intermediate zone [162, 1015]. It was
and the ipsilateral ventral funiculus of the shown, however, that fibres from the medial
cord to terminate in the medial part of the reticular formation of the medulla oblongata
intermediate zone. The medial and lateral below the level of the facial nerve nucleus
vestibulospinal tracts also distribute their distribute more widely, far beyond the limits
fibres to the medial part of the intermediate of the medial motor system, to the entire in-
zone [163, 193, 744, 1015]. The medial ves- termediate zone and the medial and lateral
tibulospinal tract takes its origin from the motoneuronal cell groups of the anterior
medial vestibular nucleus and descends bilat- horn throughout the spinal cord (Fig. 175 C).
erally in the medial longitudinal fascicle to They share this distribution pattern with the
terminate in the cervical cord. The lateral ves- raphe spinal fibres from the nucleus raphe
tibulospinal tract, which arises from Deiters' pallidus, many of which are serotoninergic,
nucleus, is an uncrossed pathway which de- and with the fibres from the locus coeruleus
scends lateral to the medial longitudinal fas- and the nucleus subcoeruleus, many of which
cicle and in the borderzone of the ventral contain noradrenalin as a neurotransmitter
and the lateral funiculus. It terminates [539, 745, 997]. The somatotopical organiza-
throughout the cord and displays a promi- tion of the reticulospinal tracts in their pro-
nent somatotopical organization. Both vesti- jection to the intermediate zone of the spinal
bulospinal tracts contribute fibres to the me- cord is limited and much less pronounced
dial motoneuronal cell groups. The un- than in the rubrospinal and lateral vestibu-
crossed interstitiospinal tract and the crossed lospinal tracts. In this respect they resemble
tectospinal tract also terminate on medially the raphe-spinal projections [576]. The raphe-
located interneurons of the intermediate spinal and coeruleo-spinal pathways, which
zone. Interstitiospinal fibres also establish seem to exert a facilitatory action on moto-
synaptic contacts with neck motoneurons, neurons, and the medial bulbar reticulospinal
whereas tectospinal fibres do not [193, 744, system, which probably contains both inhibi-
1015]. The lateral vestibulospinal tract is an tory and excitatory components, therefore
excitatory pathway for ipsilateral extensor can be considered as parts of a third motor
motoneurons [1085]. The functional proper- system, distinct from the medial and lateral
ties of the pontine reticulospinal pathway are systems considered before. The cerebral cor-
incompletely known. Transsection of the me- tex, the cerebellum and the mesencephalic re-
dial systems at the level of the brain stem ticular formation, with their extensive projec-
results in defects in the steering of integrated tions to the medial bulbar reticular forma-
Motor Systems 257
tion, may use this system to influence muscle fugal pathway from the caudal motor cortex
tone. Moreover the third motor system pro- [52, 399, 533, 539, 544, 576, 1135, 1239,
vides to limbic system with an access to so- 1240].
matomotor control, through the connections The termination of the rubrobulbar and
of the hypothalamus, the amygdala and the rubrospinal tracts includes the contralateral
central grey with the raphe nuclei and the ventral periorbital part of the facial nerve nu-
locus coeruleus [543]. cleus, the lateral reticular formation and the
spinal dorsolateral intermediate zone. At all
these levels it completely overlaps the projec-
Direct and Indirect Cortico-motoneuronal tion from the motor cortex. The size of the
Connections rubrospinal tract seems to be reduced in man
[404, 1440]. It is questionable whether it con-
The cortex projects to the entire pontine and tinues beyond its rubrobulbar trajectory.
bulbar reticular formation, the Vth, VIIth The direct projections of the cortex to the
and XIIth cranial nerve nuclei, the intermedi- medial motoneuronal cell groups of the spi-
ate zone and the anterior horn of the spinal nal cord and their interneurons in the medial
cord. These projections are bilateral to the part of the intermediate zone overlap the in-
lateral reticular formation, the Vth and XIIth direct projections of the cortex through the
cranial nerve nuclei and the ventromedial superior colliculus and the medial pontine re-
parts of the intermediate zone and the anteri- ticular formation. Both the tectospinal tract
or horn. The projections to the facial nerve and the medial reticulospinal tract terminate
nucleus, the dorsolateral part of the interme- on the ventromedial intermediate zone of the
diate zone and the lateral motoneuronal spinal cord [539, 571]. The direct cortico-
groups of the anterior horn are predominant- motoneuronal connections and the projec-
ly crossed [738, 1220]. tions from the cortex to the ventromedial in-
The direct projections to motoneurons intermediate zone are bilaterally organized and
nervating distal muscles of the limbs and the take their origin from a more rostrally locat-
facial muscles take their origin from the cau- ed part of the motor cortex than the projec-
dal part of the motor cortex in the precentral tions to the lateral motor system [744, 747].
gyrus (area 4, [176]). This is in accordance The medial reticular formation receives its
with the somatotopical localization in this cortical input from the caudal part of the pre-
area [1526]. Movements of the hindlimb, the motor area (caudal area 6) and the connec-
forelimb and the face are localized in succes- tions to the superior colliculus include still
sively more lateral and ventral parts of the more rostral parts of the frontal lobe (areas
precentral gyrus. Both limb areas display a 6 and 8 - the frontal eye field - and the ad-
concentric arrangement, with movements joining prefrontal cortex) and parts of the
around distal joints represented centrally and parietal, occipital and temporal cortices [114,
caudally, and movements around more prox- 363,671,672). The connections of area 6 with
imal joints located in the intermediate and the spinal motor apparatus which passes
rostral parts of the motor cortex. Indirect through the medial bulbar reticular forma-
projections to the lateral motor system, tion are not limited to the medial motoneuro-
through the magnocellular part of the red nal cell groups and the ventromedial part of
nucleus and the crossed rubrospinal and ru- the intermediate zone but involve the wide-
brobulbar fibres arise from the same part of spread projections of the reticulospinal com-
the motor cortex [223, 479, 831, 1076]. They ponent of the third motor system to the entire
display the same somatotopical organization intermediate zone and the medial and lateral
in their origin and the focussing of their ter- motoneuronal cell groups.
mination on the appropriate interneuronal The eye muscle nuclei do not receive fibres
and motoneuronal cell groups as the cortico- from the cortex. Cortical control of eye
movements is achieved through the saccade ventral aspect of the medulla oblongata are
generating centres in the mesencephalic teg- difficult to evaluate because in humans these
mentum (rostral interstitial nucleus of the cases are extremely rare [164]. Section of the
medial longitudinal fascicle [196]) and part pyramidal tract in monkeys results in a per-
of the medial (paramedian) pontine reticular manent loss of the ability to execute individ-
formation [194], Cajal's interstitial nucleus of ual finger movements, but is not accompa-
the medial longitudinal fascicle and the supe- nied by hypertonia or increased tendon re-
rior colliculus. The origins of the cortical pro- flexes. The ability to fractionate movements
jections to the saccade generating centres remains the single, most important contribu-
from the frontal eye field [778] overlap those tion of the pyramidal tract to motor control.
to the projections to the superior colliculus. It seems to be dependent upon the direct cor-
It has been found that many of the fibres ticomotoneuronal connections which reach
terminating in the basal pontine nuclei are their highest development in man [164, 744,
collaterals of some of the fibre systems 767]. The hypertonia and the increase of the
mentioned in this section. This holds for the tendon reflexes of the pyramidal syndrome
massive projections to the central part of the may be considered as release phenomena
pontine nuclei from the motor and sensory which appear when the shielding of the moto-
cortices, which take origin as a collateral pro- neurons from their reflex connections during
jection of the pyramidal tract [1397] and for voluntary, cortically induced movements is
the projections to more peripheral parts of abolished and may be dependent on other,
the pontine nuclei through the medial and indirect pathways. Babinski's sign remains a
lateral corticopontine divisions of the cere- reliable indicator of a lesion of the pyramidal
bral peduncle, which in part at least arise tract. The dorsiflexion of the great toe and
as collaterals from the corticotectal projec- the spreading of the toes which occur on
tion[15, 1~68,672]. stimulating the sole of the foot when the pyr-
amidal tract is interrupted should be consid-
ered as part of a flexion synergia, which is
The Pyramidal Tract Syndrome released when the pyramidal control of its
interneurons is abolished and the direct corti-
The pyramidal tract syndrome in man results cal innervation of the motoneurons of the
from an interruption of this tract's fibres any- long extensor of the great toe is lost [1422].
where along their long trajectory from the
cortex to the motoneurons. It consists of a
paralysis of voluntary muscle activity, en-
The So-Called Extrapyramidal
hanced tendon reflexes and muscle tone, dis-
appearence of abdominal reflexes and the Motor System
cremasteric reflex and inversion of the plan-
tar response of the great toe (Babinski's sign). Introduction
The extent and the laterality of the paralysis
depend on the level of the lesion. In most During the first decades of this century, the
cases other, indirect connections of the cortex concept was developed that two independent
with motoneurons are interrupted together systems, the pyramidal and the extrapyrami-
with fibres of the pyramidal tract. The most dal, converge upon the spinal motor appara-
frequent cause of a pyramidal syndrome is tus. In contrast to the direct corticospinal,
a lesion of the internal capsule, where the pyramidal system, the extrapyramidal system
cortico-rubral, -tectal and -reticular fibres lie was thought to be an array of centres which,
intermingled with fibres terminating on mo- together with their emergent fibres, consti-
toneurons and interneurons. The effects of tute a multisynaptic descending system.
isolated lesions of the pyramidal tract at the Striopallidal, pallidorubral, rubrospinal and
Motor Systems 259
reticulospinal pathways were considered to nucleus. The globus pallidus can be subdi-
be the principal links in this system. Its high- vided into an external and an internal seg-
est centre, the striatum, was believed to re- ment. The anterior limb of the internal cap-
ceive its main input from the thalamus. Ex- sule largely separates the caudate nucleus
perimental hodological studies have shown from the putamen, whereas the posterior
that the idea of independently operating py- limb of the internal capsule occupies the
ramidal and extrapyramidal systems has to space between the lentiform nucleus and the
be abandoned. The striatum and the other thalamus.
structures classically known as extrapyrami- Structurally the caudate nucleus and the
dal centres are not interconnected in a unidi- putamen are identical. They have a homoge-
rectional chain-like fashion. Rather, they and neous structure throughout and contain nu-
their emerging fibre systems constitute a merous medium-sized neurons, among which
number of interrelated loops or circuits, from conspicuous large cells are sparsely scattered.
which output systems emerge at several A quantitative analysis [1222] has revealed
points. The following survey will first present that in the human striatum sensu stricto
the structure and positional relations of the about 100 million medium-sized and some
various extrapyramidal centres and their 600000 large cells are present.
fibre connections. This overview will be fol- The globus pallidus, which is of dience-
lowed by some remarks about the internal phalic origin, differs considerably, histologi-
organisation of the striatum. Then, the subdi- cally, from the telencephalically derived stria-
vision of the basal ganglia into dorsal and tal nuclei. It is chiefly composed of rather
ventral districts, as has recently been pro- large, widely spaced, fusiform cells. The total
posed by several authors, will be considered number of pallidal cells is, in humans, about
and the most important connections of the the same as that of the large striatal neurons,
so-called ventral striatum will be discussed. i.e. 600000 [1372]. The globus pallidus is sub-
Finally, attention will be paid to the structure divided into a lateral or external and a medial
and connections of the basal nucleus of or internal segment. In primates these two
Meynert and some associated cell groups. segments are demarcated from each other by
the medial medullary lamina.
The nucleus accumbens is a cell mass
Structural Features which lies closely apposed to the most rostro-
(see also Figs. 61-87; 88-109) ventromedial part of the caudate-putamen
complex. Structurally, this nucleus cannot be
The striatum sensu stricto, i.e. the caudate sharply delimited from the striatum.
nucleus plus the putamen, is by far the largest The subthalamic nucleus is a conspicuous,
subcortical cell mass in the human brain. Its large-celled nucleus that is situated in the
fresh volume is, according to Schroder et al. most caudal part of the diencephalon, dorso-
[1222], approximately 10 cm 3 • Throughout medial to the posterior limb of the internal
its extent, the caudate nucleus is closely relat- capsule. Its medial part overlies the rostral
ed to the telencephalic lateral ventricle portion of the substantia nigra.
(Fig. 28). Its large globular head rests on the The substantia nigra is the largest cell
anterior perforated substance and is anterior- mass of the mesencephalon. Lying between
ly continuous with the putamen. The latter the tegmentum and the cerebral peduncle, its
cell mass is situated medial to the insula and most rostral part extends into the diencepha-
constitutes, with the pallidum or globus palli- lon and closely approaches the globus palli-
dus, the lentriform nucleus, a cone-shaped dus. The substantia nigra can be divided into
complex with the apex directed inwards. The a dorsal, cell-rich, pars compacta and a ven-
putamen constitutes the outer part and the tral, less cellular, pars reticulata. The pars
globus pallidus the inner part of the lentiform compacta is composed of large polygonal
cells which synthesise dopamine. Many of the adjacent ventral striatal sectors will be treat-
dendrites of these elements extend ventrolat- ed separately below.
erally into the pars reticulata. The cells in
the pars reticulata are somewhat smaller than 1. The Principal Striatal Circuit: Cerebral
those in the pars compacta. It is noteworthy Cortex - Striatum - Globus Pallidus - Thala-
that the pars reticulata of the substantia nigra mus - Cerebral Cortex (Figs. 171, 173, 176,
has many features in common with the inter- 177). Contrary to the views of previous
nal segment of the globus pallidus. The dopa- workers it has been established that the whole
minergic neurons in the pars compacta are of the neocortex sends fibres to both the cau-
often designated cell-group A9 [279). date nucleus and the putamen and that all
In addition to the pars compacta of the parts of these two cell masses receive fibres
substantia nigra, there are two other mesen- from the cortex. It was originally held that
cephalic nuclei, the cells of which synthesise the arrangement of this corticostriate projec-
dopamine. These cell groups, neither of tion is on a simple topographical basis in that
which can be sharply delimited from the sub- particular cortical areas project to proximal
stantia nigra, are known as the ventral teg- portions of the striatum [210, 676]. However,
mental area of Tsai (A10) and the nucleus more recent studies [413, 733, 734, 1227,
parabrachialis pigmentosus (A8). The former 1431, 1543] have shown that the terminal
is situated in the rostromedial part of the fields of the corticostriate fibres originating
mesencephalic tegmentum; the latter lies in from circumscribed cortical regions are ar-
the dorsolateral part of the same area and ranged in narrow, longitudinally arranged
forms part of the reticular formation. strips or bands, which often span the entire
Another nucleus which is included in the length of the nucleus. Most cortical areas
circuitry of the basal ganglia is the nucleus project only to the ipsilateral striatum, but
tegmentalis pedunculopontinus, pars com- the premotor (6, 8), motor (4) and somato-
pacta. This rather small cell mass is situated sensory (3, 1, 2) cortical areas have been
in the caudal part of the tegmentum of the shown to be distributed bilaterally [633, 676,
midbrain and lies also embedded into the 733, 734]. The fibres arising from the motor
reticular formation (Fig. 101). cortex project in a somatotopic fashion upon
the putamen (dorsal to ventral: leg, arm,
face) [733]. The corticostriate fibres, which
Fibre Connections are thought to have the excitatory amino acid
glutamate as their neurotransmitter [353], es-
In the ensuing survey of the connections of tablish direct synaptic contacts with the prin-
the striatum and related 'extrapyramidal' cipal striatal efferent neurons, i.e. the so-
centres, we will distinguish a principal striatal called medium-sized spiny cells [1270]. These
circuit and four" accessory" striatal circuits. highly characteristic cells are very common
Following the discussion of these circuits at- in the striatum.
tention will be focussed on some recent addi- Efferent fibres from the striatum converge
tions to our knowledge of the 'extrapyrami- towards the globus pallidus, where they con-
dal' circuitry and on its input and output stitute a massive fibre system, which passes
channels. Because the fibre connections to be radially through both pallidal segments. Dur-
discussed are extensively documented in sev- ing their transit through the globus pallidus
eral review articles [442, 890, 996], references the fibres of this system emit numerous col-
will be mainly confined to the most recent laterals that synapse with pallidal neurons.
literature. It should be emphasised that the The striatofugal bundle then leaves the glo-
description of the connections of the caudate bus pallidus and descends to the substantia
nucleus does not hold for the nucleus accum- nigra, passes through the internal capsule,
bens. The fibre systems related to this and then terminates mainly in the pars reticulata
Motor Systems 261
of that structure. It was previously believed cally organised [297, 687, 737], terminate in
that the striatal projections terminating in the the rostral part of the ventral lateral nucleus
external and internal segments of the globus of the thalamus [1365]. This portion of the
pallidus are collateral branches of striatoni- ventral lateral thalamic nucleus is known to
gral fibres [357,358], but studies using double project to part of cortical area 6, including
retrograde tracing [93, 1018] have shown that the supplementory motor area [1215, 1302,
these three projections come mainly from 1365). The neurotransmitter present in this
separate cell popUlations. In striatofugal pro- thalamocortical projection is not known.
jection neurons several neuroactive princi- The existence of the projections just de-
ples, among which are GABA, substance P, scribed, which are all topically organised,
enkephalin and dynorphin, have been shown strongly suggests that information derived
to be present [438, 443, 997]. Although there from the entire neocortex is processed in the
is evidence suggesting that GABA and enke- striatum, the globus pallidus and the thala-
phalin coexist in many striatal cells, it has mus, respectively, and then is fed back into
been shown that substance P is present only the premotor and supplementory motor ar-
in striatal neurons that do not contain eas of the cerebral cortex.
GABA or enkephalin [93]. Given the facts
that: (a) the external segment of the globus 2 a. The First 'Accessory' Striatal Circuit:
pallidus is particularly densely innervated by Striatum ~ Globus Pallidus ~ Thalamus ~
enkephalinergic striatofugal fibres, but con- Striatum. The striopallidal and pallidal effer-
tains only sparse substance P- and dynor- ent fibres, which form the initial part of this
phin-immunoreactive axones; (b) the internal circuit, have already been dealt with. It has
segment of the globus pallidus and the sub- been shown that a considerable number of
stantia nigra, pars reticulata contain dense fine, presumably collateral fibres leave the
networks of substance P- and dynorphin-pos- fasciculus thalamicus before it reaches the
itive striatal axons, but few that are reactive ventral lateral nucleus. These fibres enter the
for enkephalin; and (c) GABA-ergic striato- internal medullary lamina and terminate in
fugal fibres are numerous in all three target the centromedian and parafascicular nuclei
structures, but form a particularly dense [297, 687, 737, 988]. As has already been
plexus in the substantia nigra [93, 438, 443, mentioned (cf. Section Ascending Reticular
997], it may be concluded that the external System) fibres arising from these intralamin-
segment of the globus pallidus on the one ar nuclei project massively upon the striatum
hand, and the internal segment of the globus [885]. There is a clear differential distribution
pallidus and the substantia nigra on the other of the striatal fibres originating from the cen-
hand are at least in part preferentially inner- tromedian/parafascicular complex, the cen-
vated by different sets of chemically specified tromedian nucleus projecting exclusively to
striatofugal neurons [93]. the putamen and the parafascicular nucleus
The quantitatively most important effer- only to the caudate nucleus [1264]. The cen-
ent system of the striopallidum is the palli- tromedian and parafascicular nuclei do not
do thalamic projection. This projection origi- project only to the striatum, but also to the
nates exclusively from the medial pallidal seg- cerebral cortex. It has been established that
ment. Its fibres, which are thought to contain these two projections arise in separate neu-
GABA, initially constitute two separate bun- rons [1293]. The thalamostriate fibres termi-
dles, the fasciculus lenticularis and the ansa nate, like the corticostriate axons, upon
lenticularis. The fibres merge in Forel's field medium-sized spiny neurons [678] and are
H, after which they ascend as a single bundle, thought to be excitatory [337]. The neuro-
the fasciculus thalamicus, to the rostral part transmitter utilised by the thalamostriate
of the thalamus [988]. The fibres of this palli- fibres is not known.
dothalamic projection, which is topographi-
J Premo tor cortex (area 6)

2 Nucleus caudatus
3 Putamen
4 Nucleus parafascicularis + nucleus centromedianus
5 Nucleus ventraUs lateralis thalami
6 Globus paIJidus, pars laleralis
7 Globus paUidus, pars medialis
8 Nucleus subthaJamicus
JO Substantia nigra , pars reticulata
Fig. 176. The principal striatal circuit and the first, second and third "accessory" striatal circuits,
indicated by framed numbers, as described in the text. Bold lines: principal circuit
Motor Systems 263
1 e rtex prefrontalis
2 Nucleus caudatus
3 Putamen
4 Nucleus ventralis anterior thalami
7 G lobus pallid us, pars lateralis
8 G lobus pallid us, pars medialis
9 ucleus subthalamicus
11 ucleus tegmentalis pedunculopontinus,
pars compacta
Fig. 177. The fourth " accessory" striatal circuit and some ascending connections of the subthalamic
nucleus, the substantia nigra , pars reticulata and the nucleus tegmentalis pedunculopontinus, pars
compacta
Physiological evidence indicates that by in the substantia nigra. A more restricted

way of the thalamostriatal projection poly- contingent of enkephalinergic fibres is also
sensory information is fed into the striatum. present in the striatonigral projection [438,
Krauthamer [714] has expressed the opinion 443,997].
that this information may be important inter The major ascending efferent projection
alia for the execution of appropriate orient- of the substantia nigra is formed by a system
ing responses. He proposes that the loop of extremely fine axons, which originates
between the intralaminar nuclei and the from the dopaminergic cells in the pars com-
striatum enables the latter structure to self- pacta of that structure [30, 279, 1363, 1402].
regulate its polysensory input. This projection is topically organised in three
planes, medial-lateral, rostral-caudal and
2 b. The Second' Accessory' Striatal Circuit: dorsal-ventral [220, 337]. The nigrostriatal
Globus Pallidus - Corpus Subthalamicum - dopaminergic neurons establish direct synap-
Globus Pallidus. The lateral part of the globus tic contacts with striatonigral projection neu-
pallidus projects in a topically organised rons [361], but pharmacological evidence
fashion on the subthalamic nucleus [214, 217, suggests that these fibres exert an additional
872, 988] and the latter centre is known to presynaptic action on the glutamatergic cor-
project massively back to all parts of the glo- ticostriate fibres [705].
bus pallidus [214, 215, 973, 974]. GABA has There is evidence that the dopaminergic
been suggested as the probable neurotrans- nigrostriatal projection participates in the
mitter in both the pallidosubthalamic and the regulation of complex behaviour and plays
subthalamopallidal projection [352, 975, a crucial role in determining the ability of
1160]. Physiological experiments have shown the organism to cope with available extero-
that the predominant response of globus pal- ceptive sensory information in various ways.
lidus neurons to stimulation of the subtha- This specific capacity has been denoted as
lamic nucleus is inhibition [1068]. Clinico- "the ability to arbitrarily switch motor pro-
pathological and experimental evidence indi- grammes" [256, 257, 614].
cate that lesions of the subthalamic nucleus Parkinson's disease is characterised by a
lead to contralateral hemiballism, a vigorous progressive loss of dopaminergic neurons in
involuntary hyperkinesia of the extremities the pars compacta of the substantia nigra,
with a repetitive, stereotyped character [471, with consequent degeneration of their as-
1505]. Most probably interruption of the cending projections and reduction of the
inhibitory, GABA-ergic subthalamopallidal dopamine content in the striatum. In relation
fibres is a salient feature in the neural mecha- to what has been stated above concerning
nism of this subthalamic hyperkinesia [268]. the functions in which the nigrostriatal pro-
jection is involved, it is worthy of note that
2 c. The Third 'Accessory' Striatal Circuit: patients suffering from Parkinson's disease
Striatum - Substantia Ni!{ra - Striatum. It appear to have an impaired ability to switch
has already been mentioned that fibres orig- their behaviour [258].
inating from the caudate nucleus and the pu- Striatonigral fibres synapse directly with
tamen traverse the globus pallidus and subse- nigrostriatal neurons [1271, 1478], and there
quently descend to the substantia nigra. This is evidence indicating that the striatum and
strongly developed projection is topically or- the substantia nigra are interconnected in
ganised [455, 1345]. Its fibres terminate main- point-to-point reciprocity [1264]. Moreover,
ly, but not exclusively, in the external, reticu- it has been shown that striatal substance-P-
lar part of the substantia nigra. The striato- containing neurons which project selectively
nigral connection contains large numbers of to the internal segment of the globus pallidus
GABA-, substance P- and dynorphin-posi- and the substantia nigra are a preferential
tive fibres, all of which form dense networks target for nigrostriatal dopaminergic inner-
Motor Systems 265
vation [92]. It is, however, not the case that 1002, 1192, 1312]. The large neurons consti-
the entire striatonigral projection represents tuting the TPC are thought to be cholinergic
only a 'returnloop' modulating the recipro- [963, 1312]. It will be clear that the connec-
cating nigrostriatal system. The fibres of the tions just mentioned close a considerable
striatonigral system also synapse on the non- number of additional" extrapyramidal" loop
dopaminergic neurons of the pars reticulata, systems.
elements which, as will be discussed below,
form part of another 'accessory striatal cir- 4. Input systems (Fig. 177). The reticular for-
cuit' and also contribute substantially to the mation of the brain stem has a quantitatively
output system of the corpus striatum. very important access to the "extrapyrami-
dal" circuitry. The reticular formation, par-
2 d. The Fourth 'Accessory' Striatal Circuit,' ticularly its mesencephalic portion, is one of
Cortex - Striatum - Substantia Nigra - Thal- the principal sources of afferent fibres to the
amus - Cortex (Fig. 177). A substantial ni- intralaminar nuclei of the thalamus and, as
grothalamic projection originates from the pointed out earlier, the latter projects to both
reticular part of the substantia nigra. Its the caudate nucleus and the putamen. A sec-
fibres are distributed over certain parts of ond important input system to be mentioned
the ventral anterior and medial thalamic nu- here is the meso striatal serotoninergic projec-
clei [219, 586], which project to large areas tion, which originates mainly from the dorsal
of the dorsolateral and orbitofrontal associa- raphe nucleus and terminates throughout the
tion cortex, the frontal eye field, and the sup- striatum, but more, significantly in its ventral
plementory motor area [586]. and medial regions [31, 134, 397, 951, 1264,
1346, 1347].
3. Recently Established Additional Pathways. Before leaving the input systems it should
With the aid of the modern anterograde and be mentioned that our knowledge of the af-
retrograde tracer techniques a considerable ferent connections of one important extra-
number of additional" extrapyramidal" fibre pyramidal centre, the pars compacta of the
connections has been established, only a few substantia nigra, is remarkably scant. Stria-
of which will be mentioned here. It has been tonigral fibres terminate mainly on the den-
shown that the nucleus subthalamicus not drites of the compacta cells which extend into
only sends numerous fibres to the globus pal- the pars reticulata. Several other sources of
lidus, but also projects to the caudate nucleus afferents to the pars compacta have been sug-
and the putamen [90], to the pars reticulata gested, among which are the globus pallidus,
of the substantia nigra [215, 872, 974, 1121] the central amygdaloid nucleus, the lateral
and to the nucleus tegmentalis pedunculo- hypothalamic area and the habenula, but
pontinus, pars compacta (TPC) [319, 601]. none of these connections has been estab-
The latter cell mass, which until recently was lished unequivocally [890]. An afferent pro-
only known ~o receive a projection from the jection to the pars compacta arising from the
globus pallidus [988], also takes up fibres TPC, which has been described by several
from the pars reticulata of the substantia ni- authors [319, 449, 1002, 1192], could not be
gra [94, 95, 319,1053]. The neurons constitut- confirmed by ultrastructural analysis [1312].
ing the latter projection utilise GABA as their
neurotransmitter [237]. The TPC has been 5. Output Channels (Fig. 178). Because the
shown to give rise to a small descending and main outflow from the striatum and the nu-
a much larger ascending efferent projection. clei related to it converges, via the globus
The former will be considered below; the lat- pallidus and the thalamus upon the premotor
ter is distributed mainly to the globus palli- cortex, the fibres originating from this corti-
dus, the intralaminar nuclei of the thalamus cal area constitute the principal output chan-
and the subthalamic nucleus [319, 602, 890, nel of all of these nuclei. Prominent among

3 Nucleus ventralis lateralis thalami
4 Nucleus habenulae lateralis
5 Globus pallidus, pars lateralis
6 Globus pallidus, pars medialis
11 Nucleus tegmentalis pedunculopontinus, pars compacta
12 Fasciculus predorsalis (tractus tectobulbospinalis)
13 Fibrae pedunculopontino-reticulares
Fig. 178. Output channels of the so-called extrapyramidal motor system

Motor Systems 267
the systems emanating from this cortical area TPC, but some descend to the medullary re-
is, of course, the motor part of the pyramidal ticular formation. The TPC, which is situated
tract, but this area also gives rise to a number in the caudolateral midbrain tegmentum,
of other projections, some of which may well roughly corresponds to an area known as the
consist of collaterals of the pyramidal axons. mesencephalic locomotor region. This centre
Such projections terminate in the ventrallat- receives, apart from a projection from the
eral nucleus of the thalamus [742, 890], the substantia nigra pars reticulata, afferents
subthalamic nucleus [478, 1076], the substan- from the motor cortex [748], the globus palli-
tia nigra pars compacta [1405], the TPC [319] dus[214, 297, 687, 988, 1339]andthesubtha-
and in the reticular formation of the brain lamic nucleus [319, 601, 974]. Its efferents in-
stem. Several of these pathways feed back clude ascending fibres which terminate in the
into centres included in one of the striatal subthalamic nucleus, intralaminar nuclei of
circuits, thus closing additional "accessory" the thalamus and the globus pallidus, and
loops. The remaining output pathways of the descending fibres which constitute a pedun-
extrapyramidal motor system include palli- culopontinoreticular projection. The latter
dohabenular, nigrotectal, nigrotegmental has been observed to terminate in the pontine
and pedunculopontinoreticular projections. and medullary parts of the medial reticular
The pallidohabenular fibres originate formation [319,381,382,602].
from the medial pallidal segment and termi- In summary, according to our present
nate in the lateral habenular nucleus [297, state of knowledge, the striatum has four par-
509, 971]. In primates this projection origi- allel conduction lines along which the motor
nates from a peripheral zone of the medial centres in the brain stem and spinal cord may
pallidal segment, which encroaches upon the be influenced. These pass: (1) Via the globus
lateral hypothalamus [1052, 1053]. Nauta pallidus and the thalamus to the premotor
and Domesick [985] have pointed out that cortex and the pyramidal tract; (2) From the
the lateral habenular nucleus projects directly striatum, via the substantia nigra pars reticu-
to the mesencephalic raphe nuclei and is lata, to the superior colliculus; (3) Directly
probably a principal source of afferents to from the substantia nigra to the reticular for-
the serotoninergic cells contained within mation; (4) From the globus pallidus and the
these nuclei. They conjectured that the palli- substantia nigra pars reticulata, to converge
dohabenular connection could form part of upon the TPC, accompanied by descending
a neural circuit: pallidum - lateral habenular efferents from the TPC.
nucleus - dorsal raphe nucleus - striatum -
pallidum.
The nigrotectal and nigroreticular projec- Aspects of the Organisation
tions originate both from the pars reticulata of the Striatum
of the substantia nigra [95, 435, 445, 551,
1129]; both are thought to be inhibitory and The so-called medium-sized spiny cells,
to utilise GABA as their neurotransmitter which are present in enormous numbers in
[233, 237]. The massive nigrotectal projection the caudate nucleus and the putamen, occupy
terminates in a highly regular bandpattern a central position in the circuitry of those
in the middle grey layer of the superior colli- centres. The name of these neurons refers to
culus. The same layer contains the cells from the fact that their dendrites, which radiate
which the predorsal fascicle arises [868]. This and ramify in all directions, are densely cov-
is a large descending bundle which distributes ered with spines [154, 303, 433, 677]. The
fibres to a considerable number of premotor principal afferent systems to the striatum are:
and motor centres in the brain stem and spi- glutamatergic fibres from the cerebral cortex
nal cord (cf. The section Visual System). The [353, 678, 1270], fibres originating from the
nigrotegmental fibres terminate mainly in the intralaminar thalamic nuclei (whose neuro-
transmitter is still unknown) [678], dopamin- histochemical studies have revealed a re-
ergic fibres from the substantia nigra [725] markable heterogeneity within the complex.
and serotonergic fibres from the dorsal raphe The first evidence for this chemoarchitectural
nucleus [1057]. All synapse with the medium- heterogeneity came from studies in which a
size spiny cells. The medium-sized spiny cells staining technique for the enzyme acetylcho-
have long axons which leave the striatum. linesterase (AChE) was applied. These stu-
These axons, which terminate in the globus dies showed that in the caudate-putamen
pallidus and the substantia nigra, constitute complex, 300- to 600-llm-wide zones of low
the principal efferent system of the striatum. AChE activity stand out against an otherwise
Medium-sized spiny cells contain GABA AChE-rich background. Graybiel and Rags-
[349, 1018], substance-P [140], enkephalin dale [441, 442], who first identified these
[302] or dynorphin [1447], and frequently zones, designated them striatal bodies or
GABA as well as enkephalin [57]. striosomes. Graphical reconstructions have
In addition to the extrinsic afferents dis- shown that in most places the striosomes
cussed above, several groups' of intrinsic ele- form part of a complex three-dimensional
ments impinge upon the medium-sized spiny labyrinth. During recent years it has gradual-
cells. Under this heading the medium-sized ly become clear that throughout most of the
spiny cells themselves should be mentioned caudate-putamen complex the striosomes
first because, apart from their long projection and the "matrix" in which they are embed-
axons, these elements give off numerous local ded represent chemoarchitectonically distinct
collateral branches within the striatum, tissue compartments, which are related to the
which have been shown to contact other neu- intrinsic structure of the complex as well as
rons of the same type. Other local-circuit to the organisation of its afferent and efferent
neurons synapsing with medium-sized spiny connections. The following features may be
cells include large elements with aspiny den- mentioned in this context (for details cf. the
drites, containing acetylcholine, GABA-ergic review articles of Graybiel [437, 438]; and
medium-sized aspiny cells and neurons of the Graybie1 and Ragsdale [441, 443]):
same type containing somatostatin. In addi- 1. Apart from a low AChE concentration,
tion, intrinsically organised medium-sized the striosomes show high enkephalin, sub-
aspiny neurons containing VIP [696], CCK stance-P, GABA and neurotensin immuno-
[1352] and neuropeptide Y [1263, 1265] have reactivity [390, 444].
been found. The place of these elements in 2. The complementary matrix compart-
the striatal microcircuit remains to be deter- ment shows, in addition to a high AChE con-
mined. centration, a dense plexus of somatostatin-
Huntington's disease is a rare, dominantly containing fibres [390].
inherited neurodegenerative disorder, char- 3. The striosomes show a remarkably high
acterised by involuntary choreatic move- concentration of opiate receptors [1072].
ments and progressive dementia. The most 4. Neurons expressing high levels of im-
striking neuropathological feature of this dis- munoreactivity to substance P are clustered
ease is severe atrophy of the striatum, with along with dynorphin-positive neurons in the
marked neuronal loss and gliosis. It is worthy striosomes. By contrast enkephalin-positive
of note that the medium-sized aspiny cells neurons, somatostatin-positive neurons and
containing somatostatin are the only class of the large cholinergic interneurons tend to lie
neuron in the striatum to escape degenera- outside these zones [437, 438].
tion during the progression of the disease 5. The striosomes appear to coincide with
[345]. patches of high dopamine density, the so-
Cytoarchitectonically the caudate-puta- called "dopamine islands", which corre-
men complex is a homogeneous structure, spond to a distinct, early-developing contin-
but histochemical and particularly immuno- gent of nigrostriatal fibres [436, 443].
Motor Systems 269
6. Studies with anterograde tracers have tory tubercle should be considered together
revealed that not only the nigrostriatal path- as a ventral portion of the striatum. The ros-
way, but also the thalamostriatal and corti- tral part of the substantia innominata, an-
costriatal projections, terminate in a patchy other basilar forebrain structure, represents
fashion. It has been demonstrated that the a ventral extension of the globus pallidus.
terminal patches of the thalamostriatal fibres The ventral striatum and the ventral palli-
avoid the striosomes [437, 511]. The massive dum are nodal points in a loop system which
corticostriatal projection can be divided into forms a striking pendant of the principal
two fibre contingents, one that avoids the striatal circuit, described above. This loop in-
striosomes and another that projects specifi- cludes: (1) the allocortex, (2) the nucleus ac-
cally to them. Thus, the motor, somatosen- cumbens, i.e. the ventral striatum, (3) the
sory and visual areas preferentially project ventral pallidum, (4) the medial thalamic nu-
to the matrix compartment, but the efferents cleus, (5) the prefrontal, prelimbic and
from the prelimbic cortex are distributed to cingulate regions, (6) cortical area 6, which
the striosomes [311, 390]. receives projections from certain parts of the
7. There is some evidence indicating that cortical regions mentioned under (5). The
the striatal projection neurons are also differ- well-known fact that the nucleus accumbens
entially distributed over the two striatal com- receives a strong dopaminergic input from
partments. Following injections of a retro- the ventral tegmental area, whereas the cau-
grade tracer in the internal pallidal segment date-putamen complex receives a similar pro-
or in the substantia nigra pars reticulata, la- jection from the substantia nigra, further
belled neurons appear to be most densely dis- substantiates the interpretation of the nucle-
tributed in the matrix compartment, whereas us accumbens as a ventral striatum.
after injections centred in the compact part Given the fact that the allocortex forms
of the substantia nigra, labelled neurons are part of the limbic system Heimer et al. [496]
located preferentially in the striosomes [390, conjecture that, whereas the dorsal striato-
391,438]. pallidal system plays a pre-eminent role in
initiating motor activities stemming from
From the foregoing synopsis it may be
cognitive activities, the ventral striatopalli-
concluded that the caudate-putamen complex
dum has a role in initiating movements in
displays an intriguing mosaic-like chemical
response to emotionally or motivationally
heterogeneity and that several structural and
powerful stimuli.
connectional features fit into this mosaic.
A concept related to that of Heimer and
his associates has been recently put forward
by Kelly, Domesick and Nauta [675]. These
Subdivision of the Basal Ganglia
authors found that in the rat a voluminous
into Dorsal and Ventral Sectors
amygdalostriatal projection is present, which
is distributed to all parts of the striatum ex-
With regard to the extent of the striatum and cept the antero-dorso-Iateral quadrant. They
the globus pallidus Heimer and his co- pointed out that this projection widely over-
workers [493, 496, 497] have advanced a laps the striatal projections from the hippo-
view, the essence of which may be summa- campus, the cingulate cortex, the ventral teg-
rised as follows: The caudate nucleus, puta- mental area and the mesencephalic raphe nu-
men and globus pallidus, as usually delineat- clei. Like the amygdalostriatal system, all of
ed, represent only the dorsal part of the stria- these striatal afferents avoid the antero-dor-
tal complex. The nucleus accumbens, which so-lateral striatal sector. Kelley et al. [675]
cytoarchitectonically as well as histochemi- further established that the striatal sector just
cally closely resembles the caudate nucleus mentioned is the main region to which the
and putamen, and certain parts of the olfac- corticostriatal projection is distributed from
the sensorimotor cortex. In view of these the entire ventromedial half of the striatum,
findings they interpreted the large striatal re- but most massively to the ventral striatal
gion receiving a direct projection from the zone that includes the nucleus accumbens [94,
amygdala as the" limbic" and the remainder 1346, 1347]. The dopaminergic innervation
as the "non-limbic" striatal compartment. of the ventral striatum, which originates
Kelley et al. [675] suggested that the amygda- mainly from the ventral tegmental area, is
la, which is known to playa role in the neural very strong and unevenly distributed. This
mechanisms underlying motivation and uneven distribution of dopaminergic termi-
adaptive behaviour, has access via its massive nals is paralleled by the uneven distribution
projection to the striatum, to the initiation of the neuropeptides en kephalin, substance
and patterning of somatomotor behaviour in P and dynorphin and the neurotransmitter
which the latter structure is involved. It is GABA [1463]. It is noteworthy that many
important to note that the nucleus accum- neurons in the ventral tegmental area which
bens is included in the "ventral" striatum project to the ventral striatum contain chole-
as well as in the" limbic" striatum, but that cystokinin [1410, 1547] and that in some of
the former entity extends further dorsally these elements cholecystokinin is co-localised
than the latter. with dopamine [529, 532].
The efferents of the ventral striatum pass
primarily to extrapyramidal centres, like the
Fibre Connections of the Ventral Striatum ventral pallid urn, the ventral tegmental area
(Figs. 179-181) and the pars compacta and pars reticulata
of the substantia nigra, but a number of lim-
The nucleus accumbens and adjacent sectors bic-related structures, such as the septum, the
of the caudate nucleus and the putamen re- bed nucleus of the stria terminalis, the medial
ceive afferents from the cerebral cortex, par- hypothalamus, the central superior nucleus
ticularly from its limbic areas, the amygdala, and the periaqueductal grey matter are also
the midline nuclear complex of the thalamus, innervated [450, 989, 993]. The projection
the ventral tegmental area and the dorsal from the nucleus accumbens to the ventral
raphe nucleus [208, 448, 449, 993]. The corti- pallidum contains a major GABA-ergic com-
cal afferents originate from the medial fron- ponent [930], and substance-P-, enkephalin-
tal cortex, the prelimbic cortex, the posterior and dynorphin-positive fibres have also been
part of the insular cortex, the perirhinal and shown to be present within this projection
entorhinal cortices and the hippocampal su- [462, 463]. The enkephalin- and substance-P-
biculum. There is a topographical relation- positive fibres constitute dense and highly
ship between the subiculum and the nucleus characteristic terminal plexuses within the
accumbens: the rostral (uncal) part of the ventral pallidum. The plexus of enkephalin-
hippocampus projects to the medial nucleus ergic fibres continues dorsally without inter-
accumbens, whereas the caudal (splenial) ruption, into the main mass of the pallidum.
part of the hippocampus is connected with In contrast, the plexus of substance P-posi-
more rostral and lateral parts of this nucleus tive fibres extends for only a very short dis-
[449]. The cortical neurons projecting to the tance into the main pallidal mass, and thus
ventral striatum are thought to be excitatory provides a criterion for determining the ex-
and to utilise either glutamate or aspartate tent of the ventral pallidum [87, 461, 462].
as their neurotransmitter. The amygdalo- It has been demonstrated that the accumbo-
striatal projection arises mainly from the bas- fugal efferents to the ventral pallidum project
al amygdaloid nuclei. Just like the hippocam- monosynaptically to output cells of that
pal efferents, the amygdalostriatal projection structure [1536]. A similar direct, monosyn-
is topographically organised [1170, 1171, aptic relationship has been observed between
1273]. The ventral tegmental area projects to the efferents of the accumbens nucleus which
Motor Systems 271
9 ___ ~
1 Cortex prefrontalis
2 Striatum dorsale
3 Fornix
4 uclei mediani thalami
6 Nucleus habenulae la teralis
7 Cortex insulae
8 Cortex prelimhicus
9 Cortex prefrontalis medialis
10 Striatum ventrale
11 PaUidum ventrale
12 Area tegrnentalis ventralis
13 Fibrae nigrostriatales
15 Substantia nigra , pars compacta
17 ucleus tegmental is pedunculopon tinus,
pars compacta
18 uclei basales amygdalae
19 Subiculum
20 Cortex entorhinalis + perirhinali
21 Fibrae pedunculopontino-reticulares
Fig. 179. Principal connections of the ventral striatum and the ventral pallidum
pass to the pars compacta of the substantia and surrounding structures is paralleled by
nigra and nigrostriatal projection neurons a direct projection from the nucleus accum-
[1271]. In the light of this observation, it is bens to the same area [436, 450]. As has al-
plausible that a strong limbic input from the ready been pointed out, the TPC corresponds
hippocampus and the amygdala, via the ac- roughly to the so-called mesencephalic loco-
cumbens nucleus and subsequently through motor region [381, 932,1259, 1338]. Pharma-
the dopaminergic cells in the substantia ni- cological and physiological evidence suggest
gra, may reach the main, dorsal mass of the that the accumbens nucleus constitutes an in-
striatum [450]. terface between the limbic and the motor sys-
The ventral pallidum receives, in addition tem, subserving a gate function for goal-di-
to a massive projection from the ventral stria- rected behaviours, particularly locomotion
tum, afferents from: (1) the amygdaloid com- [929]. The subicular inputs appear to have
plex, particularly its basal nuclei, (2) several direct access to ventral striatal output neu-
cortical areas, including the orbitofrontal cor- rons, which in their turn project monosyn-
tex, the anterior insular cortex, the antero- aptically to the ventral pallidum. The majori-
ventral part of the temporal iobe and the en- ty of the ventral pallidal neurons are inhib-
torhinal and piriform areas, (3) the midline ited by subicular stimulation. This may be
thalamic nuclear complex, (4) the hypothala- explained by the fact that an excitatory, pre-
mus, particularly the perimamillary region, sumably glutamatergic projection from the
and (5) several centres in the brain stem, in- subiculum to the ventral striatum, activates
cluding the ventral tegmental area, the pars GABA-ergic ventral striatal neurons which
compacta of the substantia nigra, the mesen- project to the ventral pallidum, where they
cephalic raphe nuclei, the parabrachial area exert an inhibitory influence [1536]. Mogen-
and the reticular formation [1168]. It is, how- son and Nielsen [930] provided pharmacolog-
ever, important to note that most of these ical evidence suggesting that the GABA-ergic
five contingents of afferents have been traced projection from the accumbens nucleus to the
to the substantia innominata and the magno- ventral pallidum contributes to locomotor
cellular basal nucleus embedded therein (see activity. It has also been shown that manipu-
below), rather than specifically to the lation of the dopaminergic transmission in
restricted rostral part of the substantia the nucleus accumbens influences locomotor
innominata, which represents the ventral behaviour [1090]. As mentioned, the TPC
pallidum. projects to the pontine and medullary reticu-
The efferents of the ventral pallidum are lar formation. Recently, some of the reticular
distributed to the basal nuclei of the amygda- elements receiving afferents from the TPC
loid complex, the lateral habenula, the medial have been demonstrated to' project to the spi-
thalamic nucleus, the ventral tegmental area nal cord [383], thus completing a long, poly-
and to more-caudal regions of the midbrain synaptic conduction line from the limbic sys-
tegmentum, including the pars compacta of tem to the motor system.
the pedunculopontine tegmental nucleus In summary, the ventral striatum has three
(TPC) [384, 461, 931, 932, 1338). The fibres major output systems: one passing via the
passing to the medial thalamic nucleus are ventral pallidum and the medial thalamic nu-
a link in a projection from the ventral strio- cleus to the prefrontal cortex; a second lead-
pallidum to the medial prefrontal cortex [496, ing via the pars compacta of the substantia
1449]. According to Nauta [984], it is likely nigra and its doparninergic efferents to the
that through this circuit the ventral striopalli- dorsal striatum; and a third by which the
dum can affect the cognitive functions sub- ventral striatum - via the ventral pallidum,
served by the medial prefrontal cortex. The the TPC and the reticular formation - has
conduction line leading from the accumbens access to the somatomotor system.
nucleus via the ventral pallidum to the TPC
Motor Systems 273
The Magnocellular Basomedial terolateral (Ch4 al), intermedioventral (Ch4

Telencephalic Complex iv), intermediodorsal (Ch4 id) and posterior
(Ch p).
The basal forebrain contains a population of
The cell groups Ch1-Ch4 give rise to the
large cholinergic neurons, extending from the
following cholinergic projections:
septal region rostrally to the level of the sub-
thalamic nucleus caudally. Mesulam and col- 1. Fibres originating from Ch1, Ch2 and
laborators [904, 905, 907] subdivided this Ch3 pass via the stria medullaris and the ha-
population in the monkey into four groups, benulointerpeduncular tract to the base of
Ch1-Ch4 (for a map of these cell masses in the midbrain, where they terminate in the
the human brain, see Perry et al. [1069], Hed- interpeduncular nucleus and in the ventral
reen et al. [488] and Saper and Chelimsky tegmental area [346]. Cholinergic fibers
[1189]) : contained in the stria medullaris also project
heavily to the medial habenular nucleus, but
1. The Ch1 group is found in the medial whether the same or different populations of
septal nucleus; about 10% of the neurons in cholinergic neurons innervate the habenula
this centre are cholinergic. and the ventromedial mesencephalon has not
2. The Ch2 group corresponds to the verti- yet been determined.
cal limb of the nucleus of the diagonal band 2. Ch1 and, particularly, Ch2 provide a
of Broca; at least 70% of its neurons are substantial cholinergic projection to the hip-
cholinergic. pocampus, as has already been suggested by
3. The Ch3 group is embedded within the Lewis and Shute [785a]. The fibres pass
horizontal limb of the nucleus of the diagonal through the fornix and terminate in the cornu
band of Broca; only 1 % of the neurons in ammonis as well as in the fascia dentata
this nucleus are cholinergic. [696].
4. The Ch4 group, which is very extensive 3. Ch2 also sends fibres to the lateral
in the human brain, corresponds to the basal hypothalamic area.
nucleus of Meynert [319, 704, 912]. This nu- 4. Cholinergic neurons situated mainly in
cleus consists of smaller or larger groups of Ch3 pass to the olfactory bulb, to terminate
neurons, which are embedded within the sub- in the external layers of that structure.
stantia innominata [488, 1189]. The cell 5. Cholinergic neurons located in the an-
groups are surrounded by islands of acetyl- terolateral portion of Ch4 project, via the
cholinesterase-positive neuropil [1069]. The ventral amygdalofugal pathway, to the
substantia innominata is a vaguely defined amygdaloid complex, mainly to the basal
area situated ventral to the globus pallidus nucleus [696].
(Figs. 90-92). In rodents many of the large 6. The remaining parts of Ch4 provide a
cholinergic neurons are situated within the major cholinergic projection to the entire
portion of the substantia innominata defined neocortex [118, 305, 623, 683, 771, 904, 906,
as the ventral pallidum, but in primates the 910, 925, 1063, 1246]. There is a specific but
neuronal populations of the ventral pallidum overlapping topography in the organisation
and the basal nucleus appear to be less in- of this projection. Ch4 am is the major source
termingled. At least 90% of the neurons in of projections to the medial surface of the
the basal nucleus are cholinergic. In the hu- hemispheres, including the cingulate cortex.
man brain, cell-group Ch4 contains approxi- Ch4al projects to the frontoparietal cortex,
mately 200000 neurons in each hemisphere Ch4i to the prefrontal, lateral peristriate,
[51]. According to Mesulam and his asso- middle temporal and inferotemporal regions,
ciates [905, 906] the cholinergic neurons and Ch4p to the superior temporal gyrus and
which constitute Ch4 can be separated into the temporal pole region [904, 906]. The
five subgroups: anteroventral (Ch4 av), an- routes along which the efferent of the basal
1 eocortex
2 Cingulum
3 Gyrus cinguli
4 Fornix
6 ucleus habenulae medialis
7 ucleus septi medialis eCho 1)
8 Nucleus gyri diagonalis, pars dorsalis eCho 2)
9 Bulbus olfactorius
10 Nucleus gyri diagonalis, pars ventralis eCho 3)
11 Nucleus basalis of Meynert eCho 4)
12 Area tegrnentaJis ventralis
15 Fascia dentata
16 Cornu ammonis
t7 Nuclei basales amygdalae
Fig. 180. Efferent connections of the magnocellular basomedial telencephalic complex

Motor Systems 275
1 Cortex insulae
2 Nuclei septi
3 Hypothalamus
4 Nucleus gyri diagonalis, pars dorsalis
5 Nucleus basalis of Meynert (Ch.4) -
7 Nucleus gyri diagonalis, pars ventralis
8 Cortex orbitofrontalis
9 Cortex prepiriformis
10 Nuclei basales amygdalae
11 Cortex temporalis polaris
12 Cortex entorhinalis
16 Locus coeruleus
17 Nuclei parabrachiales
Fig. 181. Afferents of the nucleus basalis

nucleus reach the cortex have been studied Russchen, Amaral and Price [1168] pointed
in rat [1189], cat [1389] and monkey [696]. out that although the structures which pro-
ject to the basal nucleus are remarkably dis-
Without going into details, fibres forming parate, virtually all of them are integrative
a medial pathway enter the cingulum bundle regions or regions of polysensory conver-
and innervate the cingulate gyrus as well as gence. This holds in particular for the amyg-
dorsal and medial aspects of the cortex, dala, which is itself organised to sample in-
whereas fibres assembling in a lateral path- puts from a variety of cortical, limbic and
way enter the external capsule, fan out within subcortical structures. The amygdala, which
the corona radiata and are distributed to lat- projects massively to the basal nucleus, is be-
eral parts of the parietal, occipital and tem- lieved to playa role in evaluating whether
poral cortices. The medial and lateral path- incoming sensory information is of relevance
ways have also been observed in the human to the individual, in relation to previous ex-
brain [1189]. The cholinergic innervation of perience or to other inputs from the external
the cerebral cortex shows considerable re- or internal environment. Russchen and col-
gional variations in laminar distribution and leagues [1168] consider it likely that the inte-
intensity. This innervation is particularly grated sensory input which the amygdala and
prominent in the caudal orbitofrontal cortex, other areas of the brain provide to the basal
the cingulate gyrus, the insula, the parahip- nucleus leads to activation of the cholinergic
pocampal region and the temporal pole [909]. inputs to the cerebral cortex, and that this
The basal nucleus has been suggested to activation modulates cortical activity such
represent a telencephalic extension of the re- that relevant sensory inputs lead to integrat-
ticular formation of the brain stem [904, 910]. ed motor or emotional responses, or to learn-
It receives afferents from a variety of cortical ing and memory.
and subcortical sources [475, 594, 631, 776, The basal nucleus and its cholinergic corti-
902,904,1004, 1103, 1168, 1196]. The cortical projections have attracted considerable
cal input arises from the prepiriform cortex, interest recently because these structures have
the orbitofrontal cortex, the anterior insula, been implicated in the pathogenesis of Alz-
the rostroventral temporal cortex and the en- heimer's disease (AD) and its variant in the
to rhinal cortex. Thus, in contrast to its wide- elderly, senile dementia of the Alzheimer's
spread projections to all parts of the neocor- type (SDAT). In this disease the amounts of
tex, the basal nucleus receives reciprocal pro- choline acetyltransferase and acetylcholines-
jections from only a relatively small number terase, i.e. the enzymes which are responsible
of cortical areas. The subcortical afferents of for the synthesis and the inactivation of
the basal nucleus originate from: the septal acetylcholine, are considerably reduced in the
nuclei; the nuclei of the diagonal band; the cerebral cortex [284] and the cholinergic
accumbens nucleus - ventral pallidal com- neurons in the basal nucleus undergo a pro-
plex; the amygdala, particularly the basal nu- found and selective degeneration [333, 879,
clei; the hypothalamus; the peri peduncular 880, 1499, 1500]. Whether this loss of cells
nucleus; the pars compacta of the substantia is due to a primary lesion of the perikarya
nigra; the mesencephalic raphe nuclei; the or rather represents a retrograde phenome-
parabrachial nuclei; and the locus coeruleus. non, following degeneration of the wide-
Considering its afferents, Mesulam and co- spread cortical projections and terminals of
workers [904, 905] considered it likely that the basal nucleus neurons, has not yet been
the Ch4 complex acts as a cholinergic relay determined [50, 51,901].
between limbic plus paralimbic areas and the It is worthy of note that in AD/SDAT
entire neocortex in a fashion that may influ- some other centres which project directly to
ence complex behaviour according to the pre- the cortex, namely the noradrenergic locus
vailing emotional and motivational states. coeruleus [142, 847, 856], and the largely se-
Motor Systems 277
lr. tectos pin. tr. peduncu lo· tr. pyram id.

pontinospin.
I
I
I
I
, I ,
. ,, / ')
"' ...:::.:-..::.-:-..:::-..:.:.-:-r -.=.~.
tr. pedunculo-
B pontino retlcul.
Fig. I82A, B. Summary of the so-called extrapyramidal motor system. A the circuitry related to the
dorsal striatum; B the circuitry related to the ventral striatum. In A the bold arrows represent the
main extrapyramidal circuit and the numbers indicate the four accessory circuits described in the
text
rotonergic dorsal raphe nucleus [275, 1498, cortical projection system, is seen not only
1533] are also affected, while at the same time in AD and SDAT, but also in other disorders
such cholinergic cell groups as the large stria- with similar deterioration of memory and
tal local-circuit neurons and the somatic and cognitive functions, like Creutzfeldt-lakob's
visceral motoneurons in the brain stem and disease [50 a], Korsakoffs disease [49], many
spinal cord are spared. Finally, it should be cases of Parkinson's disease [49, 968, 1141,
mentioned that a considerable loss of neu- 1498], and the remarkable Parkinsonism-
rons in the basal nucleus of Meynert, accom- dementia complex of Guam [967].
panied by a deficiency of the cholinergic
Descending Reticular Systems
Descending Projections and the central superior nucleus send fibres

from the Raphe Nuclei to the dorsal tegmental nucleus, the pontine
(Fig. 183) central grey matter, the rhombencephalic
raphe nuclei and the pontine and medullary
reticular formation [133, 134, 1544]. Similar-
The pathways which descend from the raphe ly, the pontine raphe nucleus and nucleus
nuclei may be categorised as: raphes magnus project to the reticular forma-
tion [134].
1. Projections terminating in rhombence-
The raphespinal projection is constituted
phalic centres
by fibres that arise principally from the
2. Raphespinal projections.
raphes magnus, raphes pallidus and raphes
The former have been visualised mainly obscurus nuclei [148, 1402]. In the spinal cord
on the basis of the serotoninergic character most of these fibres descend in the superficial
of their constituent fibres and will be desig- zone of the anterior and lateral funiculi [149,
nated accordingly. The raphespinal projec- 280], with distinct concentrations in the dor-
tions contain many non-serotoninergic fibres sal part of the lateral funiculus and in the
[1258]. medial part of the anterior funiculus [501].
Serotoninergic Projections to Rhombence- The raphespinal fibres terminate preferential-
phalic Centres. The detailed mapping study ly in the superficial layers of the dorsal horn,
of Steinbusch [1288] has shown that in the the intermediolateral cell column and in the
rat a great number of rhombencephalic regions containing somatomotor neurons
centres receive serotoninergic innervation. [280, 1288].
Substantial numbers of serotoninergic fibres According to Skagerberg and Bjorklund
and terminals have been observed in the fol- [1258] the raphe spinal projection comprises
lowing cell masses: the principal sensory nu- three pathways, dorsal, intermediate and
cleus and spinal nucleus of the trigeminal ventral. The dorsal pathway, which contains
nerve, the solitary tract nucleus, the locus a large non-serotoninergic component, arises
coeruleus, the pontine central grey matter, principally from the nucleus raphes magnus,
the parabrachial nuclei, the dorsal tegmental descends through the dorsal part of the later-
nucleus, the nucleus prepositus hypoglossi, al funiculus and terminates mainly in the su-
the pontine and medullary reticular forma- perficial dorsal horn at all spinal cord levels.
tion, the raphe nuclei and most of the somat- The intermediate pathway is largely seroton-
ic and visceral motor nuclei. Little is known inergic, with its cell bodies located in the
of the specific sites at which the serotoninerg- raphes obscurus and raphes pallidus nuclei,
ic projections to all of these centres originate. particularly in the superficial ventrolateral
However, it has been established that the lo- extension of the latter, and terminates mainly
cus coeruleus receives a large projection from in areas of preganglionic autonomic moto-
the dorsal raphe nucleus [134, 225, 252, 949, neurons at thoracolumbar and upper sacral
1180,1350] and that the dorsal raphe nucleus levels. The ventral pathway consists of sero-
1 ucleus rjlphes dorsalis

2 ucleus tegmentalis dorsalis
3 ucleus centralis superior
4 Locus coeruleus
5 Nucleus raphes pontis
6 Nucleus raphes roagnus
7 Formatio reticularis metencephali
A B
10 Nucleus raphes pallid us t9 Zona incerta
11 Nucleus raphes obscurus 20 Area hypothalamica dorsali posterior
12 Formatio reticularis myelencephali 21 Area tegmentalis ventralis
13 Nucleus arcuatus 22 Cortex prefrontalis
14 Ventral raphespinal projection 23 Griseum centrale mesencephali
15 Intermediate raphespinal projection 24 Formatio reticularis mesencephali
16 Dorsal raphespinal projection 25 ucleus spinalis nervi trigemini
17 Substantia gelatinosa 26 uclei funiculi dorsalis
18 Nucleus intermediolateralis 27 Medulla spinalis
Fig. 183A, B. The descending reticular system. A descending projections from the raphe nuclei; B affer-
ent connections to the pontine and medullary raphe nuclei
Descending Reticular Systems 281
toninergic and non-serotoninergic axons de- complex and the spinal cord. A detailed dis-
scending in the anterior funiculus to termi- cussion of the descending pain-control sys-
nate in anterior horn areas along the entire tem is beyond the scope of the present text.
length of the cord. These axons originate Three points should be emphasised, however:
mainly from the nuclei raphes pallidus and
1. Neither the PAG nor the nucleus raphes
obscurus, and from the adjacent reticular for-
magnus is exclusively involved in pain con-
mation.
trol.
As regards the afferents to the caudal
2. All of the pathways involved are to be
group of raphe nuclei, the nucleus raphes
characterised as multitransmitter channels,
magnus is innervated by fibres from the pre-
i.e. they are composed of sub-populations of
frontal cortex, the zona incerta, the periaque-
fibres each containing a different neurotrans-
ductal grey matter, the nucleus cuneiformis,
mitter or combination of neurotransmitters
the mesencephalic raphe nuclei, the tegmen-
[cf.997].
tal dorsal nucleus, the dorsal funicular nuclei
3. The mode in which the descending pain-
and the spinal trigeminal nucleus [172, 209,
control system influences the inflow and pro-
320, 379, 1283]. In addition, the nucleus
cessing of nociceptive impulses in the spinal
raphes magnus receives a substantial projec-
dorsal horn is complex and involves, presum-
tion from the spinal cord [172]. Little is
ably, (a) presynaptic inhibitory action on pri-
known about the afferents to the nuclei
mary afferent fibres, (b) postsynaptic inhibi-
raphes pallidus and obscurus. Inputs to these
tion of nociceptive projection neurons, and
centres originating from the dorsal posterior
(c) excitation as well as inhibition of different
hypothalamic area [553], the periaqueductal
classes of local circuit neurons [83, 406, 754,
grey matter, the ventral tegmental area and
913].
the dorsal tegmental nucleus have been re-
ported [379]. As regards the intermediate raphespinal
The fibres of the dorsal raphespinal projec- projection, there is evidence that the medul-
tion terminate preferentially in laminae I, II lary serotoninergic neurons which project to
and V of the spinal grey matter. There is am- the sympathetic intermediolateral column in
ple morphological and physiological evidence the thoracic spinal cord exert an inhibitory
that these fibres constitute the final link in influence on the neurons in that centre, and
the so-called descending pain-control system that this sympathico-inhibitory pathway par-
[82, 83]. This descending pain-control system, ticipates in the central control of cardiovas-
or 'endogenous analgesia' system, consists of cular function [809].
a series of interrelated centres or areas situ- The fibres of the ventral raphespinal pro-
ated at three different levels of the neuraxis: jection terminate in close contact with moto-
the mesencephalon, the rhombencephalon neurons in the ventral horn, particularly in
and the spinal dorsal horn. It is in the latter primates [703]. Activation of the serotonin-
area that the system inhibits noxiously ergic neurons that project to the ventral horn
evoked activity. The mesencephalic complex facilitates all motoneurons, extensors as well
involved in pain control consists of the cen- as flexors, and (X- as well as y-motoneurons
tral mesencephalic griseum or periaqueductal [703, 964]. It is conceivable that during the
grey matter (PAG), the dorsal raphe nucleus, execution of motor actions that are of vital
which is partly embedded in the central grey priority, such as fight and flight, the dorsal
matter, and the adjacent reticular formation, and ventral raphespinal projections act in
i.e. part of the cuneiform nucleus. At the concert, by suppressing pain transmission on
rhombencephalic level, the nucleus raphes the one hand, and by increasing the respon-
magnus and adjacent portions of the medial siveness of motoneurons on the other hand
reticular formation constitute an important [745, 1464].
relay between the mesencephalic pain-control
Descending Projections project mainly through the posterolateral

from the Medial Reticular Zone funiculi. They extend throughout the length
(Fig. 184) of the spinal cord, terminating bilaterally in
laminae VII, VIII and X, and ipsilaterally
in IV, V and VI [1552]. A large proportion
For the sake of simplicity, we will gather of the reticulospinal fibres give off collateral
under this heading the reticulospinal tracts branches to more than one level of the spinal
and certain descending pathways that im- cord, indicating that the reticular formation
pinge upon the cells from which these tracts is involved in the coordination of activities
originate. It should be appreciated that the at different spinal levels [1075].
reticular neurons that project to the spinal According to Kuypers [741, 745] the me-
cord are influenced by many other atTerents dial reticular formation is involved in (among
(see p. 211) and that the descending parts of other activities) guiding movements of the
the reticular formation cannot be sharply trunk and the proximal parts of the limbs.
separated from the ascending parts. Within As he pointed out, the reticulospinal fibres
the neural network of the reticular formation terminate mainly in the ventromedial area of
there is widespread interaction between as- the spinal intermediate zone, where the inter-
cending and descending impulses [162, 164, neurons that impinge on the motoneurons to
626]. The reticulospinal tracts comprise a the axial and proximal limb muscles are
ponto spinal and a bUlbospinal system [1014]. located. The same function is subserved not
The former consists almost exclusively of un- only by the reticular formation, but also by
crossed fibres, the latter of both crossed and a number of other brain stem centres, such
uncrossed fibres. as the interstitial nucleus of Cajal and the
The pontospinal or medial reticulospinal vestibular nuclei. Kuypers categorised the de-
tract originates from the nucleus reticularis scending projections arising from these
pontis caudalis and from the caudal part of centres as the medial system of brain stem
the nucleus reticularis pontis oralis. It de- pathways. A lateral system of brain stem
scends throughout the length of the spinal pathways projects to the interneurons that
cord, chiefly in the medial part of the anterior impinge on the motoneurons of the lateral
funiculus. Its fibres terminate mainly in la- anterior horns, at the levels of the cervical
minae VII and VIII of the spinal grey matter, and lumbar enlargements. This system,
but also extend into the portion of lamina which preferentially guides distal-extremity
IX occupied by the motoneurons which in- movements, was regarded by Kuypers as
nervate the trunk musculature [539, 626, comprising mainly the crossed rubrospinal
1014]. tract.
The bulbospinal or lateral reticulospinal In addition to control of spinal motor
tract can be subdivided into two components, function, the reticulospinal projections may
a dorsal one originating mainly from the ven- also be involved in transmission of sensory
tral part of the medullary reticular formation impulses. There is evidence suggesting that
and a ventral one arising mainly from the reticulospinal fibres terminating in laminae
dorsal part of the same structure [626, 1552]. IV, V, VI and VII may modulate transmis-
Both components are bilateral, but crossed sion of nociceptive input [1552].
fibres are less numerous than uncrossed ones. Before discussing the main descending
The ventral component, which is confined to pathways that impinge on the cells from
the rostral half of the cord, descends in the which the reticulospinal tracts originate, it
most ventral parts of the anterior and antero- should be noted that the medial reticular for-
lateral funiculi. Its fibres terminate mainly in mation densely innervates most branchiomo-
laminae VII and VIII, and to a lesser extent tor and somatomotor cranial nerve nuclei
in IX. The fibres of the dorsal component [626].
3 Nucleus preoplicus
4 Nucleus anterior hypothalami
5 Hypothalamus
6 Fasciculus telencephalicus medialis
7 Formatio reticularis mesencephali
8 ucleus reticularis pontis oralis
9 Fasciculus uncinatus cere belli
10 ucleus fastigii
11 ucleus reticularis pontis caudalis
13 Nucleus medullae oblongatae centralis
14 Tractus pontospioalis
15 Tractus bulbospinalis
Fig. 184. Descending fibre systems of the medial reticular formation. For the ascending fibres see
Figure 156
Most afferent fibres to the reticulospinal Among the many other fibre systems that
neurons originate from the cerebral cortex. converge on the lower part of the medial re-
These fibres, which travel by way of the corti- ticular formation (cf. p.211 and Fig. 156),
cotegmental part of the pyramidal tract, are fibres originating from the cerebellum should
distributed bilaterally in the reticular forma- be emphasised. These fibre systems originate
tion. Kuypers and colleagues [221, 740, 743] in the fastigial nucleus and pass with the
have shown that the cortical afferents to the crossed and direct fastigiobulbar bundles to
medial reticular formation originate mainly the reticular formation, where they terminate
form the premotor area. In addition to the primarily in the gigantocellular nucleus [85].
medial reticular formation of the lower brain
stem, the mesencephalic reticular formation
also receives many corticofugal fibres. The
Descending Projections from the
mesencephalic reticular formation does not
Lateral Reticular Zone
give rise to direct reticulospinal fibres, but
(Figs. 185-189)
it contains numerous neurons that project to
the pontine and medullary parts of the me-
dial reticular formation [320, 1435], thus Under this heading we will discuss the de-
forming links in an indirect reticulospinal scending projections of the parvocellular re-
pathway. The mesencephalic reticular forma- ticular area, the superficial ventrolateral re-
tion also receives several fibre systems de- ticular area, the lateral pontine tegmentum,
scending from limbic forebrain structures. the noradrenergic cell groups Al-A7 and the
These fibre systems include descending con- adrenergic cell-groups C1-C2. The cells in the
stituents of the medial forebrain bundle, the parvocellular reticular area give rise to rela-
mamillotegmental tract and the medullary tively short ascending and descending axons
stria-habenulointerpeduncular tract (Fig. which together form a proprio bulbar system
198B) [976, 977, 1195, 1295]. (Fig. 185) distributed to the branchiomotor
The pedunculopontine tegmental nucleus, nuclei and the nucleus of the hypoglossal
in the caudolateral part of the mesencephalic nerve [538, 541]. The fibres and parent cells
reticular formation, receives projections de- of this system interconnect the afferent com-
scending from a number of extrapyramidal ponents of certain cranial nerves with the ef-
centres, including the globus pallidus or dor- ferent system of the lower brain stem, thus
sal pallidum [988], the portion of the substan- constituting links in the various bulbar reflex
tia innominata known as the ventral palli- pathways. However, fibres originating from
dum [1338], the subthalamic nucleus [319, higher levels of the brain also terminate on
974] and the pars reticulata of the substantia the cells of the lateral reticular formation.
nigra [94, 319]. As has already been men- Prominent among these are cortical fibres,
tioned, the pedunculopontine nucleus rough- arising from the contralateral motor area (fi-
ly corresponds to the physiologically defined brae corticotegmentales), and mesencephalic
"mesencephalic locomotor region". It gives fibres, arising from the contralateral red nu-
rise to both ascending and descending projec- cleus. According to Kuypers [743] these two
tions. The ascending projections feed back fibre contingents, together with the lateral re-
to various forebrain centres integrated in ex- ticular interneurons and the bulbar motoneu-
trapyramidal circuitry, but the descending rons, constitute the supraspinal part of a
projections pass to the rhombencephalic re- functional system implicated in the control
ticular formation, particularly to those areas of fine, differentiated movements. Some cor-
which give rise to reticulospinal fibres [381, ticofugal fibres of this system bypass the re-
1283]. Thus, the medial reticular formation ticular interneurons and act directly on mo-
appears to be included in several truly extra- toneurons.
pyramidal descending conduction channels.
3 Nucleus ruber, pars magnocellularis
4 Decussatio tegmentajis ventralis
6 Forrnatio reticularis lateralis
7 ucleus motorius nervi trigemini
9 Nucleus nervi hypoglossi
Fig. 185. Descending fibre systems of the lateral reticular formation

It is worthy of note that the parvocellular spiratory group [805, 812]. The ventral
reticular area is traversed by a major aggre- group, which contains inspiratory as well as
gation of catecholaminergic fibres, the longi- expiratory bulbospinal neurons, forms part
tudinal catecholamine bundle of Jones and of the ARSVL. The inspiratory neurons pro-
Friedman [624]. Many of the fibres in this ject to both the phrenic and external intercos-
bundle originate from the locus coeruleus tal motoneurons, whereas the expiratory neu-
complex [626]. It has been suggested that this rons excite spinal motoneurons that inner-
conduit contains not only noradrenergic and vate the internal intercostal and abdominal
adrenergic fibres, but also several peptidergic muscles [812]. The ARSVL also contains
fibre contingents, and in fact represents an neurons which are important in cardiovascu-
"open multineuromediator channel", i.e. a lar control. At least some of these neurons
fibre system which influences surrounding project directly to the preganglionic sympa-
neurons throughout its extent [997]. thetic neurons of the thoracic intermedio-
It has already been mentioned that the su- lateral column [28, 819].
perficial ventrolateral reticular area (ARSVL; The question as to whether the ARSVL
Fig. 186) represents an aggregation of centres can be subdivided into separate functional
involved in visceral control (cf. p. 203). This areas has been addressed by a considerable
area receives a substantial projection from number of investigators (cf. Feldberg [339]
the solitary tract nucleus, which represents for review). The recent physiological map-
the anatomical substrate of baroreceptor, ping studies of McAllen [869, 870] have re-
chemoreceptor and respiratory reflexes [97, vealed the following functional pools of neu-
288, 805, 1125, 1159, 1207]. Efferents from rons: a vasoconstrictor and cardioaccelerator
the ARSVL descend to the spinal cord, where pool, a depressor pool, two separate respira-
they terminate bilaterally in the thoracic in- tory stimulation pools and a sudomotor
termediolateral cell column [28, 127, 498, pool.
807, 921], in lamina IX at the C4-C6 level Electrical stimulation of a number of
and, mainly contralaterally, in laminae VIII brain areas, including the hypothalamus, the
and IX in the thoracic cord [340]. Afferents periaqueductal grey matter and the adjacent
descending to the ARSVL from higher levels tegmentum of the midbrain may elicit a pat-
of the brain include fibres from the Kolliker- tern of somatic and visceral responses char-
Fuse nucleus [367, 1191] the parabrachial nu- acteristic of defensive behaviour. The auto-
clei [367, 1191], the caudal periaqueductal nomic component of this behavioural pattern
grey matter and adjacent tegmentum [818], has been designated the" visceral alerting re-
the cuneiform nucleus [320], the zona incerta sponse". This complex response includes
[41] and the hypothalamus, particularly the powerful respiratory activation, an increase
paraventricular nucleus and the lateral hypo- in arterial blood pressure resulting from an
thalamic area [41,534,553,818,1190]. increased heart rate and cardiac output, and
The Kolliker-Fuse nucleus, a pneumotaxic vasoconstriction in the renal and splanchnic
centre, projects to two lower respiratory beds, accompanied by active vasodilatation
centres known as the dorsal and ventral re- in skeletal muscle. Physiological evidence
spiratory groups. The dorsal group is in the suggests that the ARSVL receives direct in-
ventrolateral part of the solitary tract nucle- put from the hypothalamic and mesencephal-
us. It consists of inspiratory bulbospinal neu- ic "defence areas" and is essential for effec-
rons which project to the spinal cord and tuation of the visceral alerting response [288,
send collaterals to the ventral respiratory 523,818].
group. Inspiratory spinal motoneurons, both The structure and connections of the later-
phrenic (in C4-C6) and intercostal (in the al pontine tegmentum are complex (Fig. 187).
thoracic cord), are monosynaptically excited This area encompasses the relatively distinct
by the bulbospinal neurons of the dorsal re- lateral and medial parabrachial nuclei, which
may be interpreted as viscerosensory relay and descends via the medial forebrain bundle
centres. The Kolliker-Fuse nucleus is a ven- and the central tegmental field. It originates
tral extension of the medial parabrachial nu- from the central amygdaloid nucleus, a car-
cleus. The pneumotaxic centre is a pool of diovascular and respiratory centre, and ter-
cells corresponding to the Kolliker-Fuse minates in the parabrachial region. Just like
nucleus, but encroaching upon the medial the projection from the cortex, this amygda-
parabrachial nucleus. The lateral pontine teg- loparabrachial connection is reciprocated by
mentum also contains two separate regions a corresponding ascending pathway. It is also
involved in the regulation of micturition. One notable that the central amygdaloid nucleus
of these, the M-region of Holstege and colla- also projects to the solitary tract nucleus and
borators [537], is in the dorsomedial part of the dorsal vagal nucleus [1103, 1223].
the lateral pontine tegmentum. It corre- The third contingent of fibres descending
sponds in position to the micturition reflex to the lateral pontine tegmentum originates
centre first described by Barrington [80a]. from the preoptic region and the hypothala-
The other micturition centre is in the ventro- mus [108, 339, 534, 1195]. Although these
lateral pontine tegmentum, and was desig- fibres and their terminal fields are usually
nated by Holstege and colleagues [537] as the considered to be widely and diffusely distrib-
L-region. Although neither of these two re- uted, several more specific connections have
gions is a distinct cytoarchitectonic entity, been recognised within this continuum. The
both give rise to discrete spinal projections median preoptic nucleus and the paraventri-
[537]. The lateral pontine tegmentum receives cular nucleus project preferentially to the lat-
projections descending from the cerebral cor- eral parabrachial nucleus [367, 1359] and the
tex, the amygdala and the hypothalamus and anterior hypothalamic area sends numerous
gives rise to two separate pathways to lower fibres to the M-region [534].
bulbar centres and at least five different pro- As mentioned, the descending efferents of
jections to the spinal cord. These fibre con- the lateral pontine tegmentum can be subdi-
nections will now be briefly discussed. (For vided into projections to the lower medulla
a survey of the ascending projections to and oblongata and projections to the spinal cord.
from the lateral pontine tegmentum, the The bulbar projections terminate in the soli-
reader is referred to pp. 211-217). tary tract nucleus and in the ARSVL [367,
The cortical projection to the lateral pon- 1158]. Many of these fibres originate from
tine area originates from an insular cortical the Kolliker-Fuse nucleus and are involved
area which, on account of its general and in the regulation of respiration.
special (gustatory) visceral afferents, may be Somewhat schematically, the fibres de-
designated the visceral cortex [367]. This in- scending from the lateral pontine tegmentum
sular cortical projection terminates in the to the spinal cord may be divided into five
parabrachial region, mainly in the medial contingents [537, 539, 540]: the subcoeru-
parabrachial nucleus [1186, 1187, 1244, leospinal pathway, the crossed ponto spinal
1540], thus reciprocating a corresponding as- pathway and spinal projections originating
cending projection [1187, 1244]. The visceral from the Kolliker-Fuse nucleus, the M-re-
cortex projects not only to the parabrachial gion and the L-region. Their features are:
region, but also to several other central auto-
nomic structures, among which the solitary 1. The subcoeruleospinal pathway origi-
tract nucleus and the dorsal vagal nucleus nates from the medial part of the lateral pon-
may be especially mentioned [1187, 1243, tine tegmentum. Its fibres pass through the
1413,1540]. lateral zone of the bulbar tegmentum and
The projection to the lateral pontine area continue throughout the length of the spinal
which originates from the amygdala forms cord. They descend in the superficial part of
part of the ventral amygdalofugal pathway the lateral funiculus and are distributed,
---~
1 Nucleus paraventricularis, pars parvocellularis

2 Area hypothalamica lateralis
3 Zona incerta
4 Formatio reticularis mesencephali (nucleus cuneiformis)
6 Nucleus of Kolliker-Fuse
9 Area reticularis superficialis ventrolateralis
10 Nucleus solitarius, pars cardiorespiratoria
11 Fibrae bulbospinales
Fig. 186. The descending reticular system. Descending projections related to the area reticularis superfi-
cialis ventrolateralis
1 Operculum frontoparietale ("cortex visceraJis " )

2 Nucleus para ventricularis
3 Nucleus preopticus median us
4 Nucleus anterior hypothalami
5 Nucleus centralis amygdalae
6 AIeasubcoenliea
7 M-region of Hoistege et a1. (537)
8 L-region of Holstege et al. (537)
9 ucleus parabrachia.lis medialis
10 Nucleus parabrachialjs lateralis
11 Tegmentum pontis ventrolaterale
l2 Nucleus of Kolliker-Fuse
13 Fibrae pontospinales
Fig. 187. The descending reticular system. Descending connections related to the tegmentum pontis
laterale
mainly ipsilaterally, to all parts of the spinal al reticular zone. The projections descending
grey matter [539, 540, 1496]. from these cell groups will now be dealt with.
2. The crossed ponto spinal pathway arises It is, however, important to note that these
from the ventrolateral part of the pontine projections show a considerable overlap with
tegmentum, decussates at rostral pontine lev- the various "non-transmitter-specified"
els and descends via the lateral bulbar teg- pathways discussed above.
mentum to the contralateral spinal dorsolat- The descending efferents of the locus coer-
eral funiculus. From this pathway fibres are uleus complex (Fig. 188) follow two different
distributed to spinal laminae I and II and routes, which may be designated the caudal
to the lateral parts of laminae V and VI of limbs of the dorsal periventricular pathway
the intermediate zone throughout the length and of the dorsal noradrenergic bundle.
of the spinal cord [193, 539, 540]. The caudal limb of the dorsal periventricu-
3. Fibres originating from the Kolliker- lar pathway, like the rostral limb of the same
Fuse nucleus descend via the ARSVL to the system, is part of the longitudinal dorsal fas-
spinal cord [367, 749, 1191, 1376]. They pro- ciculus complex [344]. The contribution of
ject to the phrenic nucleus in segments C4-- the locus coeruleus to this pathway in the
C6 and to the T1-T3 autonomic preganglion- rat is confined to a limited number of fibres
ic neurons in the intermediolateral column which arise from the ventral locus coeruleus
[537, 539]. and terminate in the solitary tract nucleus
4. The M-region in the dorsomedial part [1357]. However, in the rhesus monkey a con-
of the lateral pontine tegmentum corre- siderable number of fibres originating from
sponds to the micturition reflex centre of the locus coeruleus has been traced to the
Barrington [80a] and gives rise to fibres dorsal motor vagal nucleus [1496].
which descend via the ARSVL and the super- The caudal limb of the dorsal noradrenergic
ficial part of the lateral funiculus to the sacral bundle is part of the major longitudinal cate-
cord, where they project strongly and bilater- cholamine bundle, described by Jones and
ally to the intermediomedial and intermedio- Friedman [624]. It descends through the
lateral cell groups [537, 810]. The sacral inter- rhombencephalic lateral reticular zone and
mediolateral cell group contains parasympa- continues caudally into the lateral funiculus
thetic neurons which form the motor supply of the spinal cord [624, 942]. In primates and
to the detrusor muscle of the bladder [965]. man this bundle follows the trajectory of the
5. The L-region in the ventrolateral part central tegmental tract for a certain distance
of the lateral pontine tegmentum also gives [1061]. Noradrenergic fibres originating from
rise to fibres which, following a route similar the locus coeruleus complex supply, partly
to those arising from the M-region, descend directly and partly by way of the bundle just
to sacral levels of the spinal-cord. However, discussed, the following rhombencephalic
these fibres are not destributed to autonomic centres [784]: the parvocellular reticular area,
cell groups, but to a somatomotor centre, the the nuclei of the lateral lemniscus, the princi-
nucleus of Onuf [537]. This centre, which was pal sensory trigeminal nucleus, the cochlear
first described by Onufrowicz [1032], is in the nuclei [723, 724], the pontine nuclei, the spi-
central part of the anterior horn of segments nal trigeminal nucleus and the entire rhom-
S1-S2. It innervates the striated muscles of bencephalic reticular formation. The latter
the pelvic floor, including the striated mus- three centres receive, in addition to fibres
cles of the anal and urethral sphincters [751, from the locus coeruleus, input from other
1201,1534]. noradrenergic cell groups.
The locus coeruleus complex innervates all
All of the central noradrenergic cell groups segments of the spinal cord [1016,1496]. This
and most of the central adrenergic cell groups innervation is bilateral, the decussation oc-
are located within the rhombencephalic later- curring at spinal levels [246, 662]. The coeru-
Fig. 188. Fig. 189.
1 Area subcoerulea 1 Cell group A7

2 Locus coeruleus 2 Cell group AS
3 Nuclei lemnisci lateralis 3 Cell group CI
4 Nuclei ponlis 4 Cell group C2
5 Nucleus sensorius principalis nervi trigemini 5 Nucleus dorsalis nervi vagi
6 Formatio reticularis metencephali 6 Nucleus solitarius
7 Nucleus reticularis parvocellularis 7 Longi tudinal catecholamine bundle
8 Nucleus cochlearis ventralis 8 Fibrae reticulospinales
10 Longitudinal catecholamine bundle
12 Formatio reticularis myelencephali
16 Fibrae reticulospinalcs
Fig. 188. The descending reticular system. Descending projections from the locus coeruleus complex
Fig. 189. The descending reticular system. Descending projections from the noradrenergic AS and
A7 groups (continuous lines) and from the adrenergic C1 and C2 groups (interrupted lines)
leospinal fibres originate in the ventral part give rise to which terminal networks is lack-
of the locus coeruleus and in the subcoeruleus ing. However, Hokfelt and colleagues [531]
area [460, 473, 862, 1205, 1495], descend in have presented experimental evidence indi-
the lateral funiculus of the cord [1496], and cating that the projections to the spinal cord
terminate in the ventral parts of the dorsal originate mainly from cell group C1
horn (laminae IV, V, VI), the intermediate (Fig. 154A). The experiments of Ross et al.
grey matter and the central horn [794, 1496]. [1157] have also shown that adrenalin-con-
Onufs nucleus receives a particularly dense taining neurons in the C1 group project to
noradrenergic innervation [701, 702, 922], the thoracic cord.
but the projection descending to this nucleus Physiological experiments [281, 282, 869,
from the pontine L-region also contains non- 870] have shown that the ventrolateral por-
noradrenergic fibres [537]. The locus coeru- tion of the medulla oblongata contains a cir-
leus complex does not innervate the sympa- cumscribed and highly sensitive vasopressor
thetic intermediolateral column in the thorac- area, and evidence has been presented sug-
ic cord [247,1016,1496]; however, in the rhe- gesting that this centre corresponds to the
sus monkey the parasympathetic pregang- adrenergic C1 group [420]. Thus it seems like-
lionic neurons in the sacral cord have been ly the noradrenergic A5 group and the adren-
shown to receive a major noradrenergic input ergic C1 group both project to the sympa-
from the locus coeruleus [1496]. The projec- thetic preganglionic neurons in the thoracic
tion to these neurons from the pontine M- spinal cord and that both provide a portion
region, on the other hand, is non-noradrener- of the anatomic substrate of vasomotor con-
gic [810]. According to the earlier studies of trol.
Dahlstrom and Fuxe [280] the non-coerulean
noradrenergic fibres descending to the spinal
cord originate mainly from the lower medul-
lary A1 and A2 groups. However, more re-
cent experimental investigations have re-
vealed that this projection arises largely, if
not exclusively, from the pontine groups A5
and A7 [127, 795, 808,1494,1495] (Fig. 189).
This ponto spinal projection presumably de-
scends in the longitudinal catecholamine
bundle and enters the deeper part of the later-
al funiculus of the spinal cord [808, 1205].
Its fibres terminate in the superficial layers
of the dorsal horn, in the area around the
central canal and, with great density, in the
thoracic sympathetic intermediolateral col-
umn [230, 271,808, 1495). The fibres project-
ing to the sympathetic preganglionic neurons
originate mainly from the A5 group, which
presumably represents a vasomotor centre
[808,1524].
The nucleus of the solitary tract and the
dorsal vagal nucleus, both in the caudal
rhombencephalon, and the intermediolateral
nucleus in the spinal cord all show a high
density of adrenergic terminals [530]. Exact
knowledge of which adrenergic cell groups
Olfactory and Limbic Systems
Introduction will also be given to the olfactory system,

which is intimately related to some compo-
Certain territories of the diencephalon, the nents of the limbic system. Indeed, an earlier
telencephalon and the mesencephalon are generation of neuroanatomists considered
structurally and functionally so closely inter- most of the structures referred here to as
related that they may be considered a single limbic, as forming parts of the rhinencepha-
functional complex, which has been desig- lon. The intricacy of the relationship to be
nated as the limbic system. In most general dealt with require, as far as the illustrations
terms, this complex may be characterised as are concerned, a three-step procedure. Fig-
being directly involved in processes aimed at ure 190 presents the various moieties of the
the survival of the individual (organism) and limbic system and their fibre connections in
of the species. As such, the mammalian lim- an extremely schematised fashion. This figure
bic system plays a key role in [825, 842, 843, will serve as an introductory key diagram.
844,977,982,986,997,1337]: In Fig. 191 the major limbic conduction
channels are shown semidiagrammatically.
1. The maintenance of homeostasis, a com-
This figure is intended as a link between the
plex task accomplished by
key diagram Fig. 190 and Figs. 193-213,
(a) Activation of visceral effector mecha-
which show the limbic structures and connec-
nisms
tions in greater detail. Before commencing
(b) Modulation of hormone release from
a concise appraisal of the limbic system, two
the anterior and posterior pituitary
general remarks should be made. First, the
(c) The initiation of feeding and drinking
relevant literature has grown so overwhelm-
and the related foraging behaviours.
ingly in the past 10 years that any attempt
2. Agonistic (i.e. defense and attack) behav-
at completeness has been necessarily aban-
iour.
doned. Second, as in previous sections, with
3. Sexual and reproductive behaviour.
regard to the fibre connections we have relied
At the telencephalic level, the limbic sys- heavily on the published results of experi-
tem is represented by the septal and preoptic mental work on laboratory animals. It is by
regions, the hippocampus and some adjacent no means unlikely that several or even many
cortical areas, the amygdala and the bed nu- of the hodological data presented do not ap-
cleus of the stria terminalis. The diencephalic ply to the human brain, the more so since
components include the epithalamus, the hy- (a) most experimental studies of limbic path-
pothalamus and the subthalamic zona incer- ways have been carried out using rats rather
tao In the midbrain, the limbic system com- than primates and (b) certain limbic struc-
prises a number of cell masses, all of which tures show considerable interspecies differ-
lie in or close to the median plane. In the ences [1330, 1529].
present section, an attempt will be made to
survey the structure and fibre connections of
all of these territories. Brief consideration
Fig. 190. Summary of the limbicohypothalamic complex. Subdivision of the area into central units
and limbic rings, H hypothalamus; LMA limbic midbrain area; PO preoptic region; S septum
Olfactory and Limbic Systems 295
17
t Cingulum t1 Tractus mamillotegmentalis

2 Fornix 12 Tractus habenulointerpeduncularis
3 Stria terminalis 13 Fasciculus telencephalicus medialis
4 Stria medullaris thalami 14 Pedunculus corporis mamillaris
5 Nucleus anterior thalami 15 Corpus mamillare
6 Nucleus medialis thalami 16 Ansa peduncularis
7 Nuclei habenulae 17 Bulbus olfactorius
8 Tractus mamillothalamicus 18 Stria olfactoria lateralis
9 Fasciculus longitudinalis dorsalis 19 Corpus amygdaloideum
10 Commissura anterior 20 Hippocampus
Fig. 191. The major pathways of the limbic system and the rhinencephalon
The Central Limbic Continuum: more cellular than in women [1318]. Compa-
Sections and Centres rable differences have been found in the rat
(Figs. 192-195; see also Part IV, preoptic region [423, 524, 1251].
Figs. 88-109) The epithalamus comprises, apart from
the pineal gland, the habenular ganglion.
The septal region (septum verum or pre- Within this cell mass, separate medial and
commissural septum) forms part of the me- lateral nuclei can be distinguished. The two
dial wall of the cerebral hemispheres. It is habenular ganglia are interconnected by the
situated directly rostral to the lamina termi- habenular commissure.
nalis within the paraterminal gyrus (Fig. 11). The subthalamic zona incerta is situated
It is bordered dorsally by the corpus callo- in the caudal part of the diencephalon, just
sum, rostrally by the precommissural portion ventral to the thalamus. Its rostral part occu-
of the hippocampus, and caudally by the an- pies a position dorsal to the subthalamic nu-
terior commissure and the preoptic region. cleus (corpus subthalamicum), whereas its
Ventrolaterally it borders on the nucleus ac- caudal part is ventrally bounded by the cere-
cumbens, a large cell mass that in location bral peduncle. The most caudal part consists
and function occupies a position intermedi- of a cell mass known as the peripeduncular
ate between the limbic and striatal or "extra- nucleus [1130], which receives a multifarious
pyramidal" systems. Contrary to the prevail- input from many different parts of the neu-
ing opinion, the septal region is well devel- raxis. According to Roger and Cadusseau
oped in the human brain [42]. It contains a [1140] the zona incerta can be subdivided into
number of rather poorly individualised cell a ventral and a dorsal sub-zone, the former
groups, among which the medial and lateral receiving afferents predominantly from som-
septal nuclei may be mentioned. The ventro- esthetic centres, the latter being the target of
medial part of the septum is occupied by an limbic projections.
aggregation of large cells which forms the The hypothalamus encompasses the most
dorsal or septal limb of what is known as ventral part of the diencephalon, where it
the nucleus of the diagonal band of Broca. forms the floor and contributes to the lateral
The ventral or tubercular limb of this nucleus walls of the third ventricle. Its upper bound-
extends caudolaterally along the ventral sur- ary is marked on the ventricular side by a
face of the hemisphere and marks the caudal shallow groove, the hypothalamic sulcus.
boundary of the olfactory tubercle. As its Caudally the hypothalamus passes gradually
name implies this nucleus is embedded in a over into the periventricular and tegmental
fibre bundle, the diagonal band of Broca. grey matter of the mesencephalon. It is, how-
The preoptic region, which flanks the ros- ever, customary to define the posterior mar-
tral end of the third ventricle, is a rather nar- gin of the hypothalamus as a vertical plane
row vertical strip of tissue that extends from passing just caudal to the mamillary bodies.
the anterior commissure to the rostral edge The latter are paired small eminences located
of the optic chiasm. Although this region is on the basal aspect of the brain. The rostral
of telencephalic origin, it is closely related boundary of the hypothalamus coincides
structurally to the hypothalamus. The three with a vertical plane passing from the fora-
rather poorly differentiated cell masses pres- men of Monro to the middle of the optic
ent in it - the preoptic periventricular nucle- chiasm. The infundibular stalk, which is situ-
us, the medial preoptic nucleus and the later- ated directly posterior to the optic chiasm,
al preoptic nucleus - may be considered ros- connects the funnel-shaped rostroventral
tral extensions of the three hypothalamic cell part of the hypothalamus with the pituitary
zones. Notably, the human preoptic region gland.
contains a sexually dimorphic nucleus. In Within the hypothalamus three medio-
men, this nucleus is considerably larger and laterally arranged zones can be recognised
1 Columna fornicis 13 Nucleus supraopticus

2 Nucleus paraventricularis 14 Nucleus suprachiasmaticus
3 Area lateralis hypothalami 15 Nucleus infundibularis
4 Nucleus posterior hypothalami 16 Arteria hypophyseos superior dextra
5 Area tegmentalis ventralis 17 Infundibulum
6 Nucleus preopticus medialis 18 Pars infundibularis } L b "
7 Nucleus anterior hypothalami 19 " t I"
P ars d IS 0 us antenor
a IS h h
8 Nucleus dorsomedialis 20 Pars intermedia ypop yseos
9 Nucleus ventromedialis 21 Lobus posterior hypophyseos
10 Fasciculus mamillaris princeps 22 Sinus intercavernosus posterior
11 Corpus mamillare 23 Sinus intercavernosus anterior
12 Nucleus preopticus lateralis 24 Arteria hypophyseos inferior sinistra
25 Arteria hypophyseos inferior dextra
Fig. 192. The hypothalamic nuclei and the relationship between the hypothalamus and the pituitary
gland (4/1 x)
1 Gyrus cinguli 13 Crus fornicis

2 Indusium griseum 14 Gyrus fasciolaris
3 Stria terminalis 15 Fasciola cinerea
4 Nucleus medialis thalami 16 Fissura choroidea
5 Nuclei habenulae 17 Gyrus dentatus
6 Nucleus ruber 18 Subiculum
7 Fasciculus telencephalicus medialis 19 Cornu ammonis
8 Corpus mamillare 20 Site of limbus Giacomini
9 Septum verum 21 Nucleus corticalis amygdalae
10 Area subcallosa 22 Nucleus anterior amygdalae
11 Gyrus diagonalis 23 Nuclei basalis + lateralis amygdalae
12 Fibrae amygdalofugales ventrales 24 Cortex prepiriformis
Fig. 193. The structures of the limbic and olfactory systems and some input-output pathways as seen
in a medial view (3/2 x). Some displacement of structures serves to bring other structures in view.
The walls of the third ventricle and the brain stem have been omitted almost completely; of the
thalamus only the anterior, medial and habenular nuclei are illustrated
1 Stria terminalis . { stria terminalis

14 Precommlssural stria medullaris thalami
2 Fornix components of fornix
3 Commissura fornicis
15 Stria terminalis postcommissuralis
16 Septum verum
17 Lamina terminal is
6 Tela choroidea ventriculi tertii
20 Corpus mamillare
9 Nuclei habenulae
21 Nucleus ruber
10 Commissura habenulae
22 Tractus mamillotegmentalis
Fig. 194. The central part of the limbic area; medial view of nuclei and tracts (5/2 x).
I Stria longitudinalis medialis

3 Indusium griseum
4 Fornix precommissuralis
6 Columna fornicis
10 Subiculum
II Ventriculus lateralis, cornu
inferius
12 Cornu ammonis (gyrus
uncinatus)
13 Limbus Giacomini
14 Cornu ammonis (gyrus
intralimbicus)
IS Cornu ammonis (digitationes
hippocampi)
16 Corpus fornicis
18 Gyrus dentatus
19 Crus fornicis
20 Commissura fornicis
21 Site of corpus callosum
22 Gyrus fasciolaris
Fig. 195. The limbic structures isolated from most of their surroundings, seen from above (2/1 x).
3 Stria olfactoria medialis
6 Tuberculum olfactorium
7 Limen insulae
8 Bandeletta diagonalis
10 Fasciculus telencephalicus
medialis
11 Fibrae amygdalofugales
ventrales
12 Nucleus corticalis
13 Nucleus anterior
14 Nucleus lateralis Corpus
15 Nucleus cen tralis amygdalo-
16 Nucleus medialis ideum
17 Nucleus basalis
(accessorius)
21 Stria terminalis
22 Commissura habenulae
23 Nuclei habenulae
Fig. 196. The rhinencephalic structures isolated from most of their surroundings, seen from above
(2/1 x). The interrupted line indicates the limen insulae and its continuation around the "hilus" of
the temporal lobe
[267], a periventricular, a medial and a lateral is rostrally continuous with the lateral hypo-
zone. The periventricular zone consists of a thalamic area (Fig. 97) - and the interpedun-
few layers of small cells arranged parallel to cular nucleus. The latter is an unpaired medi-
the ventricular surface. These layers alternate an structure which lies in the caudobasal re-
with thin sheets of fine, mostly unmyelinated gion of the midbrain. The second group is
fibres. The nucleus infundibularis, a cell mass composed of the central grey matter, the dor-
in the basal hypothalamus, may be consid- sal raphe nucleus, the superior central nucle-
ered a derivative of the periventricular zone. us (of Bechterew) and the dorsal tegmental
The medial zone, which is relatively cellular, nucleus (of Gudden). The mesencephalic cen-
contains a number of variably discrete cell tral grey matter is formed by a mantle of
masses. The latter are usually placed in three densely packed, mostly small cells surround-
groups, an anterior, an intermediate and a ing the cerebral aqueduct. Rostrally it is di-
posterior group. The anterior group is rectly continuous with the subependymal
formed by the rather ill-defined anterior nu- grey matter of the third ventricle and caudal-
cleus [1194], the distinct, magnocellular su- ly it blends into the central grey matter of
praoptic and paraventricular nuclei, and the the pontine tegmentum. In humans, the dor-
small suprachiasmatic nucleus. The interme- sal tegmental nucleus is a small, diffuse cell
diate group consists of the dorsomedial and mass, which flanks the caudal part of the dor-
ventromedial nuclei, which are poorly delim- sal raphe nucleus at the level of the trochlear
ited from each other. The posterior group nucleus. The two raphe nuclei which form
includes the medial, intermediate and lateral part of the limbic midbrain area, i.e. the dor-
mamillary nuclei and the diffuse posterior sal raphe nucleus and the superior central nu-
nucleus. The latter nucleus, which contains cleus, have already been discussed in the sec-
numerous scattered large cells, is caudally tion concerning the ascending reticular sys-
continuous with the peri ventricular and teg- tem.
mental grey matter of the midbrain. The lat-
eral hypothalamic zone is partly separated
from the medial zone by the postcommissural The Central Limbic Continuum:
fornix, a large bundle connecting the hippo- Circuitry and Major Conduction Channels
campal formation with the mamillary body. (Figs. 197, 198)
The medial surface of the internal capsule
and of the subthalamic region form the later-
al boundary of the lateral hypothalamic Experimental hodological studies have
zone. Caudally this zone passes imperceptibly shown that all of the structural entities in-
over into the mesencephalic ventral tegmen- volved in the limbic system are intercon-
tal area; rostrally it is continuous with the nected by short and long, ascending and de-
lateral preoptic area. Although the lateral hy- scending fibres. Taken together, these fibres
pothalamic zone contains a few local conden- constitute one large functional system, which
sations of cells, most of its territory is occu- Nauta [980, 986] has designated as the 'limbic
pied by a diffuse neuronal matrix known as system-midbrain circuit'. Schematising some-
the lateral hypothalamic area (LHA). This what, it may be stated that the septal,
area constitutes, together with the lateral pre- preoptic and anterior hypothalamic areas
optic area, the bed nucleus of the medial fore- form the rostral pole of this circuit, whereas
brain bundle (see below). the paramedian midbrain area represents its
The so-called limbic midbrain area [977] caudal pole. The hypothalamus may be char-
encompasses two groups of nuclei, both of acterised as a nodal way station interposed
which occupy a paramedian position. The between these rostral and caudal poles. Two
first group consists of the ventral tegmental large telencephalic parts of the limbic system,
area of Tsai - a poorly defined cell mass that namely the amygdala and the hippocampal
formation, as well as the olfactory system, After this introductory survey of the cen-
are reciprocally connected with the rostral tral circuitry of the limbic system (Fig. 190),
pole of the circuit (Figs. 190, 191). These brief consideration will be given to some of
structures and their fibre connections will be the major conduction channels involved in
discussed below. The caudal pole of the cir- this circuitry.
cuit may be considered as a paramedian sub- The fasciculus medialis telencephali or me-
division of the brain stem reticular formation dial forebrain bundle (MFB) (Fig. 191) may
[321, 986]. The centres forming this pole are be considered the central longitudinal path-
largely integrated into both ascending and way of the limbic forebrain-midbrain contin-
descending pathways. The ascending path- uum. It is an assemblage of loosely arranged,
ways connect the lower parts of the reticular mostly thin fibres, which extends from the
formation, and the visceral sensory centres septal area to the tegmentum of the mid-
situated in the caudal part of the medulla brain. It traverses the lateral preopticohy-
oblongata, with the hypothalamus. The de- pothalamic area, the scattered neurons of
scending pathways convey impulses from the which are collectively designated as the bed
hypothalamus to the visceral and somatic nucleus of the medial forebrain bundle. The
motor centres in the brain stem and spinal bundle is highly complex, comprising a vari-
cord. ety of short and long ascending and descend-
The mamillary body, which is situated in ing links [998, 1438, 1445, 1446]. In the tran-
the caudobasal part of the hypothalamus, re- sitional area of the diencephalon and the
ceives a large projection from the hippocam- mesencephalon, the MFB fibres are rear-
pal formation and sends most of its efferents ranged into a smaller medial and a larger
to the anterior nucleus of the thalamus. lateral stream [534, 557, 983]. The medial
These two connections form part of a closed fibre stream roughly maintains the sagittal
hippocampo-mamillo-thalamo-cingulo-hip- orientation of the hypothalamic trajectory of
pocampal system known as the circuit of the bundle. It passes through the medial parts
Papez (Fig. 206; [1051]). of the mesencephalic and rhombencephalic
The effector mechanism of the hypothala- tegmental areas, just next to the raphe nuclei.
mus include, apart from fibre systems de- The medial stream is composed of descending
scending to the brain stem and spinal cord, fibres, by which several hypothalamic centres
two hypothalamohypophyseal pathways. project to the raphe nuclei and to the adja-
One of these, the partly neural and partly cent parts of the medial reticular formation
humoral tuberoinfundibulohypophyseal sys- [534, 983], and also comprises fibres which
tem, is the means by which the hypothalamus ascend from the raphe nuclei to the lateral
controls the production of the various anteri- hypothalamus, from where they pass to a va-
or pituitary hormones. Most fibres that inter- riety of diencephalic and telencephalic
connect the rostral and caudal poles of the centres (cf. the section on ascending reticular
limbic forebrain-midbrain continuum tra- systems). The lateral stream of fibres extend-
verse the hypothalamus. The descending limb ing from the MFB to the brain stem sweeps
of this transhypothalamic projection is, laterally and caudally over the dorsal border
however, supplemented by a notable dorsal of the substantia nigra into a ventrolateral
conducting route that bypasses the hypo- tegmental position immediately caudal to the
thalamus. This route, which is synaptically substantia nigra; at this level it curves dorso-
interrupted in the epithalamic habenular medially and then descends through the me-
nuclei, is formed by the habenulopetal stria sencephalic central tegmental area to the lat-
medullaris and the habenulofugal habenulo- eral tegmental field of the pons and the me-
interpeduncular tract (Fig. 155 B). dulla oblongata. This fibre stream contains
fibres descending from the central nucleus of
the amygdala [550, 1103, 1412], the bed nu-
1 Fibres ascending to medial parts of th.e cerebral cortex

2 Fibres ascending to lateral parts of the cerebral cortex
3 Fornix
4 Nucleus septi lateralis
5 Nucleus interstitialis striae termina1is
6 Nucleus habenulae lateralis A
7 Nucleus septi medialis 35 Cortex infralimbicus (area 25)
8 Nucleus para ventricularis 36 Cortex prelimbicus (area 32)
9 Area lateralis hypothalami 37 Medial and orbital regions of cortex prefrontalis
10 Nucleus posterior hypothalami 38 Nucleus accumbens
11 Zona incerta 39 Thberculum olfactorium
12 Nucleus dorsomedialis 40 Stria medullaris thalami
13 Nucleus ventromedialis 41 Nucleus habenulae medialis
14 Nucleus anterior hypothalami 42 Nucleus peripeduncularis
15 Nucleus preopticus medialis 43 Fasciculus telencephalicus medialis
16 Nucleus gyri diagonalis 44 Nucleus interpeduncularis
17 Nucleus infundibularis 45 Nucleus tegmentalis pedunculopontinus
18 Corpus mamillare 46 Area subcoerulea
19 Area tegmentalis ventralis 47 Nucleus rapbes magnus
20 Griseum centrale mesencephali 48 Formatio reticularis metencephali
21 Nucleus raphes dorsalis 49 Formatio reticularis myelencephli
22 Nucleus centralis superior 50 Adrenergic cell group C2
23 Nucleus tegmentalis dorsalis 51 Nucleus reticularis parvocellularis
24 Nuclei parabrachiales 52 Nucleus rapbes paUidus
25 Locus coeruleus 53 Adrenergic cell group Ct
26 Griseum centrale pontis 54 Area reticularis superficialis ventrolateralis
27 Fasciculus longitudinalis dorsalis 55 Nucleus raphes obscurus
28 Nucleus solitarius 56 Nucleus dorsalis nervi vagi
29 Noradrenergic cell group At 57 Fibrae hypothalamospinales
30 Noradrenergic cell group A2 58 Organum subfornica1e
31 Area postrema 59 Organum vasculosum laminae terminalis
32 Fibres passing to the nucleus medialis and 60 Nucleus supraopticus
nucleus centralis amygdalae 61 Nucleus suprachiasmaticus
33 Fibres originating from the nucleus-medialis 62 Fibrae retinohypothalamicae
and nucleus centralis amygdalae 63 Eminentia mediana
34 Nuclei mediani thalami Ii Foramen ventriculare
c
Fig. 197 A-C. Fibre connections of the telencephalic and diencephalic portions of the central limbic
continuum; A ascending projections; B descending projections; C short connections
36
1 Cortex prelimbicus (area 32)

2 Gyrus cinguli (area 24)
3 Cortex prefrontalis
4 Nucleus interstitialis striae terrninalis A
5 Nuclei mediani thalami
7 Nucleus habenulae lateralis
10 Nucleus septi lateralis 26 Area tegmentalis ventralis
11 Nucleus septi medialis 27 Formatio reticularis mesencephali
12 Nucleus caudatus + putamen 28 Griseum centrale mesencephali
13 Nucleus accumbens 29 Nucleus raphes dorsalis
14 Nucleus gyri diagonaiis, pars dorsalis 30 Nucleus centralis superior
15 Thberculum olfactorium 31 Nuclei parabrachiales
16 Nucleus gyri diagonalis, pars ventralis 32 Locus coeruleus
17 Nucleus preopticus medialis 33 Noradrenergic cell group A7
18 Nucleus paraventricularis 34 Cortex entorhinalis (area 28)
19 Nucleus anterior hypothalami 35 Formatio reticularis metencephali
20 Nucleus centralis amygdalae 36 Nuclei centrales cere belli
21 Nucleus ventromedialis 37 Noradrenergic cell group A5
22 Nucleus posterior hypothalami 38 Formatio reticularis myelencephali
23 Fasciculus medialis telencephali 39 Nucleus solitarius
24 Zona incerta 40 Noradrenergic cell group Al
25 Colliculus superior 41 Fasciculus anterolateralis
42
I
I
I
I
I
2 I
I
42 Cortex frontalis (areae 6 + 8)

43 Cortex prefrontalis (areae 9, 10, 13)
44 Cortex insulae
46 Corpus mamillare
47 Substantia nigra, pars compacta B
49 ucleus interpeduncularis
50 ucleus medialis amygdalae
51 uclei basalis + lateralis amygdalae
52 Subiculum
53 ucleus tegmentalis dorsalis
54 Area subcoerulea
57 Nucleus preposi tus hypoglossi
S8 Nucleus raphes pallidus
S9 Nucleus reticularis parvocellularis
61 Area reticularis superficialis ven trolateralis
62 Fibrae mesencephalospinales
Fig. 198A, B. Fibre connections of the bed nucleus of the stria terminalis and of limbic midbrain
centres; A ascending projections ; B descending projections
cleus of the stria terminalis [542, 1412] and nates in the dorsal tegmental nucleus and in
several hypothalamic areas [534, 557, 825, the nucleus reticularis tegmenti pontis of
1184]. These descending fibres terminate in Bechterew [269, 1123].
a variety of brain stem centres, among which The mamillary peduncle receives fibres
the substantia nigra (pars compacta) para- from the dorsal tegmental nucleus. It passes
brachial nuclei, locus coeruleus, nucleus sub- ventrally and then ascends along the ventral
coeruleus, the noradrenergic Al, A2 and A5 surface of the midbrain to the mamillary
groups, the superficial ventrolateral reticular body, where most of its fibres terminate.
area and the dorsal vagal complex may be Some of its fibres join the medial forebrain
especially mentioned. Most of these descend- bundle and spread to the lateral preopticohy-
ing MFB projections are reciprocated by cor- pothalamic zone and the medial septal nucle-
responding ascending projections [1445, us [946, 987].
1446]. Indeed, the MFB is a major descend- It has already been mentioned that the
ing and ascending link between the forebrain stria medullar is thalami and the habenuloin-
and the brain stem. terpeduncular tract together constitute a dor-
The dorsal longitudinal fasciculus of SchUtz sal diencephalic conducting system that is
is, much like the medial forebrain bundle, synaptically interrupted in the habenular nu-
a composite system consisting of thin ascend- clei [1317]. The fibres constituting the stria
ing and descending fibres. This fascicle ex- medullaris assemble in the area behind the
tends from the posterior part of the hypotha- anterior commissure and pass along the dor-
lamus to the caudal medulla oblongata and somedial border of the thalamus to the ha-
occupies a periventricular position over its benular nuclei (Fig. 194). The habenulointer-
entire length. Rostrally its fibres become in- peduncular tract descends from the habenu-
corporated into the hypothalamic periventri- lar nuclei to the basal region of the midbrain,
cular system (Fig. 191). In the older literature where part of its fibres terminate in the inter-
[266, 986], it has been reported that most of peduncular nucleus. The two epithalamic cell
the ascending and descending projections masses, i.e. the medial and lateral habenular
contained within the dorsal longitudinal fas- nuclei, represent mutually isolated processing
cicle are synaptically interrupted in either the stations in the dorsal diencephalic route.
central grey matter of the midbrain or in the Their connections will be considered in the
dorsal tegmental nucleus of Gudden. More next section.
recently it has, however, been established that Hypothalamohypophyseal pathways are
substantial fibre contingents passing directly shown in Figs. 192 and 197C. The magnocel-
from the forebrain to the autonomic centres lular nuclei of the anterior hypothalamus, i.e.
of the lower medulla oblongata and vice ver- the supraoptic and paraventricular nuclei,
sa are also present [472, 1124, 1190). give rise to axons that descend through the
The mamillotegmental tract, containing infundibular stalk to the posterior lobe of
the efferents of the mamillary nuclei, consti- the pituitary. These axons, which together
tutes a large, compact bundle known as the form the supraopticoparaventriculohypo-
principal mamillary bundle (fasciculus ma- physeal tract, transport colloid droplets con-
millaris princeps). This bundle passes dorsal- taining the hormones oxytocin and vasopres-
ly for a short distance and then splits up into sin to the posterior pituitary lobe or neuro-
two components, the larger mamillothalamic hypophysis, where they are released into the
and the smaller mamillotegmental tract blood. The cells of the nucleus infundibularis
(Fig. 191). The mamillothalamic tract, which are involved in controlling the secretion of
passes to the anterior thalamic nucleus, the anterior pituitary hormones. They exert
forms part of the so-called Papez circuit. The this control by means of regulating hormones
mamillotegmental tract curves caudally into that stimulate or inhibit the liberation of the
the tegmentum of the midbrain and termi- hormones produced in the pituitary. Each of
the latter has a corresponding regulating hor- The CVOs are primarily looked upon as
mone. The regulating hormones pass from chemosensitive zones, which monitor the
the infundibular nucleus, along the axons of changing levels of circulating hormones and
its constituent cells, to the median eminence, other substances. The efferent fibres of the
where they are released from the axon termi- CVO neurons pass to other neural centres
nals into the capillaries of the hypophyseal and function as initial links in central regula-
portal system. The latter system forms a vas- tory circuits [159, 1123, 1253, 1336). Three
cular link between the infundibulum and the important circumventricular organs, the sub-
adenohypophysis. It is noteworthy that the fornical organ, the organum vasculosum la-
median eminence, a conspicuous neurohemal minae terminalis and the area postrema, will
organ situated in the anterior wall of the in- now be briefly discussed. The subfornical or-
fundibular stalk, receives (apart from the gan (SFO) and the organum vasculosum of
neurosecretory axons just mentioned) several the lamina terminalis (OVLT) are both un-
other afferent fibre systems that are thought paired structures situated in the wall of the
to be concerned with the release of the regu- small nonevaginated portion of the telence-
lating hormones. phalon, i.e. the telencephalon impar (cf.
Fig. 2), the former rostrodorsally, just be-
tween the two interventricular foramina, the
latter rostroventrally, immediately dorsal to
The Central Limbic Continuum:
the optic chiasm.
Functional Connections
The SFO is highly vascular and contains
of Individual Centres numerous receptors for angiotensin II. It re-
ceives neural afferents from the preoptic re-
In the present section, a concise survey will gion and the anterior hypothalamic area
be presented of the functional connections [792]. Its efferents include fibres projecting
of the centres which together constitute the to the paraventricular and supraoptic nuclei
central limbic forebrain-midbrain contin- [916], the infralimbic prefrontal cortex, the
uum. This survey will be prefaced by some bed nucleus of the stria terminalis, the medial
notes on the circumventricular organs, re- hypothalamic area, the substantia innomina-
markable structures which are all either di- ta, the zona incerta and the lateral hypo-
rectly or indirectly related to the central limb- thalamic area [1336]. The SFO plays a
ic continuum. prominent role in body fluid balance [621].
Circumventricular organs (CVOs) are Activation of its neurons by blood-borne
small loci of specialised tissue which, as their angiotensin II is known to lead to the release
name implies, are situated in the immediate of vasopressin from the posterior pituitary, to
vicinity of the ventricular system. Most of stimulate thirst and to induce drinking behav-
these loci contain typical neurons, and all of iour. Swanson and Lind [1336] have recently
them lie in regions of the central nervous sys- indicated how the different efferent projec-
tem devoid of a blood-brain barrier. The neu- tions of the SFO might be involved in the
rons in the CVO's may be influenced by: various components of drinking behaviour.
Their suggestions included the possibility
1. Neural afferents entering the organs from that the efferents to the substantia innomina-
other parts of the central nervous system ta, zona incerta and lateral hypothalamic
or from the periphery. area may be involved in behavioural arousal
2. Substances carried by the cerebrospinal as well as in the modulation of somatomotor
fluid. control systems and that the efferents to the
3. Plasma-borne substances. The latter are infralimbic cortical areas and the bed nucleus
generally considered to playa prominent of the stria terminalis may playa role in cog-
role. nitive and/or affective components of thirst.
The OVLT and the immediately adjacent The septal region contains the lateral sep-
peri ventricular zone of the preoptic region tal nucleus and the medial septal complex.
are often together designated as the periven- The latter comprises the medial septal nucle-
tricular anteroventral third ventricle (A V3V) us and the medial or dorsal nucleus of the
[621]. Like the SFO the OVLT is a receptor diagonal band of Broaca. Stimulation and le-
site for the central effects of angiotensin II sion experiments suggest that the septal re-
[1083, 1253]. It plays a role in homeostatic gion is involved in behavioural activities in
functions of water-electrolyte balance and many different spheres, such as feeding and
blood-pressure regulation and has also been drinking, micturition and defecation, sexual
implicated in the control of LHRH secretion and reproductive behaviour and aggression
[1091]. The OVLT receives afferents from, [456, 1060, 1248].
among other structures, the SFO, the lateral The lateral septal nucleus receives a major
preoptic and lateral hypothalamic areas, the afferent projection from Ammon's horn and
anterior, dorsomedial and ventromedial hy- the subiculum of the hippocampal formation,
pothalamic nuclei and from the locus coeru- which reaches the septum by way of the pre-
leus [202, 825]. It has been shown to project commissural fornix (Figs. 197 B, 205; [894,
to the dorsomedial, supraoptic and paraven- 1330]). Other afferents to the lateral septal
tricular nuclei [825, 916, 1249]. nucleus originate from the preoptic region
As its name implies, the area postrema is, [823], several hypothalamic centres (includ-
contrary to the SFO and the OVLT, located ing the anterior, paraventricular and ventro-
in the caudalmost part of the brain. It is a medial nuclei and the lateral hypothalamic
small, bilateral, spongy structure that pro- area; [254, 781, 823, 1193]), the noradrener-
trudes into the fourth ventricle on either side gic locus coeruleus and A1 and A2 areas
of the midline, immediately rostral to the [800], the largely serotoninergic raphe nuclei
obex. Physiological experiments suggest that [134, 697, 823], the laterodorsal tegmental
the area postrema is a chemoreceptive trigger nucleus, the parabrachial nuclei, the
zone forming part of the neurocircuitry un- Kolliker-Fuse nucleus and the dorsal vagal
derlying water and energy balance, as well complex [823]. The lateral septal nucleus
as cardiovascular regulation [255, 580, 917]. gives rise to three major efferent projections.
The area is also known to play an important First, it projects massively upon the medial
role in eliciting nausea and vomiting in re- septal and diagonal band complex [1331].
sponse to circulating emetic substances [145, Second, it projects, mainly via the MFB, to_
146]. The neural connectivity of the area pos- the medial and lateral preoptic, anterior and
trema has been recently reviewed by Leslie lateral hypothalamic, supramamillary and
and Gwyn [782]. It receives a significant ventral tegmental regions [107, 1331]. Third,
visceral afferent input, by way of the vagus it projects through the stria medullaris to
nerve, which includes sensory information some thalamic midline nuclei and to the me-
from thoracic and abdominal viscera. Baro- dial habenular nucleus [1331].
receptive information is conveyed to the area As indicated above, the complex formed
postrema via the carotid sinus branch of the by the medial septal and diagonal band nu-
glossopharyngeal nerve. Moreover, the area clei (medial septal complex) receives a mas-
has been reported to receive a direct projec- sive projection from the lateral septal nucle-
tion from a number of scattered hypothalamus. Other afferents to this complex arise from
ic cells, situated dorsolateral to the dorso- the lateral preopticohypothalamic area [1195,
medial hypothalamic nucleus [556]. The two 1320], the medial mamillary nucleus [1433]
principal outputs of the area postrema are and the dorsal tegmental nucleus [946, 987].
directed to the subjacent nucleus of the soli- The medial septal complex projects massively
tary tract and the ventrolateral portions of to the hippocampal formation, including the
the medial and lateral parabrachial nuclei subiculum [1331], and to the en to rhinal cor-
[924, 948, 1273, 1396].
tex [20]. Additional efferents of this complex many forebrain centres that project to this
have been traced to the preoptic region, the nucleus also contain large numbers of ster-
lateral hypothalamic area, the mamillary oid-concentrating cells. These hormone-con-
complex, the supramamillary region, the ven- centrating elements may well compose a neu-
tral tegmental area and the mesencephalic ral network that coordinates the influences
raphe nuclei [1331]. of circulating gonadal steroids on specific
From this brief overview, it appears that brain regions in a number of physiological
the septal nuclei form part of a number of and behavioural responses [1250, 1306].
neuronal loops which also include the hippo- Efferents of the MPN have been traced
campal formation, the preoptic region, the to a considerable number of telencephalic,
hypothalamus and a number of monoamin- diencephalic and brain stem centres [34, 253,
ergic centres in the brain stem as notable 1320]. Telencephalic efferents project to the
nodal points. However, the assertion of nucleus of the diagonal band, the lateral sep-
Swanson and Cowan [1331], made in 1979, tal nucleus, the substantia innominata and
that it is difficult to relate these connectional the lateral preoptic area, and via the stria
data in any meaningful way to the results terminalis to its bed nucleus and to the me-
of stimulation or ablation experiments still dial amygdaloid nucleus. The diencephalic ef-
remains valid. ferents include:
The preoptic region comprises the periven-
tricular, medial and lateral preoptic nuclei. 1. Fibres passing through the stria medullar-
The periventricular nucleus has already been is to the lateral habenula.
discussed along with the OVLT; hence we 2. Fibres projecting to the periventricular
confine ourselves here to the medial and lat- thalamic nuclei via the periventricular
eral preoptic nuclei. fibre system.
The medial preoptic nucleus (MPN) is 3. Subthalamic efferents terminating in the
thought to play an important role in a variety zona incerta [1339].
of physiological, behavioural and endocrine 4. Hypothalamic projections to the anterior
functions, including thermoregulation [147, and lateral hypothalamic areas and to the
470], hypovolaemic thirst [1337], male sexual median eminence.
behaviour [241 a], maternal behaviour [1012]
and modulation of gonadotropin secretion Efferents of the PMN passing to the brain
by the anterior pituitary [660]. It receives neu- stem descend via the periventricular bundle
ral afferents from widely distributed areas in of Schutz to the mesencephalic and pontine
both the forebrain and the brain stem. Major central grey matter, and (by way of the me-
inputs arise from the medial amygdaloid nu- dial forebrain bundle) to the ventral tegmen-
cleus, the subiculum, the bed nucleus of the tal area, the pedunculopontine nucleus and,
stria terminalis and the lateral septal nucleus. sparsely, to other parts of the mesencephalic
Other sources of afferents to the MPN in- reticular formation [34, 1339]. It has been
clude the infralimbic and insular cortical ar- found that numerous gonadal-steroid-con-
eas, the nucleus accumbens, the substantia centrating cells in the MPN project to the
innominata, most hypothalamic centres, the mesencephalic central grey matter and to the
peripeduncular nucleus, the ventral tegmen- ventral tegmental area [334].
tal area, the periaqueductal grey matter, the As regards the possible functional signifi-
raphe nuclei, the locus coeruleus and the A1 cance of the various connections of the
area, and finally the caudal, interoceptive MPN, the following suggestions have been
part of the nucleus of the solitary tract [107, made:
1250]. It is also worth mentioning that the
MPN contains numerous cells that concen- 1. The afferents originating from the me-
trate gonadal steroids [1197, 1311] and that dial amygdala may convey olfactory im-
pulses, whereas higher-order visual, auditory preoptic nucleus, the anterior, paraventricu-
and tactile information may be relayed to the lar, dorsomedial, ventromedial and posterior
MPN via the subiculum. All of these sensory hypothalamic nuclei, the lateral hypothalam-
stimuli have been shown to influence a ic area, the mesencephalic and pontine cen-
number of functions in which the MPN is tral grey matter, the dorsal raphe nucleus,
implicated, including the control of gonado- the parabrachial nuclei and the locus coeru-
tropin release, sexual behaviour and mater- leus. The LPN gives rise to both ascending
nal behaviour [1250]. and descending fibres [1320]. Ascending
2. Modulatory effects of gonadal hor- fibres pass to the medial septal-diagonal
mones may influence MPN neurons directly, band complex. Descending fibres course
but may also exert indirect effects through through the stria medullaris to the lateral ha-
their action on steroid-concentrating cells benula and (via the medial forebrain bundle)
that supply afferents to the nucleus [1250]. to the mamillary body, the supramamillary
3. The direct estrogen-sensitive pathways region, the ventral tegmental area, the pedun-
from the MPN to the periaqueductal grey culopontine nucleus and the periaqueductal
matter and the ventral tegmental area may grey matter [1338, 1339, 1343]. The specific
control behaviour such as male sexual behav- functional significance of the LPN is less un-
iour and maternal behaviour [334]. derstood than that of the MPN. However,
4. There is considerable physiological evi- there is evidence indicating that the LPN, to-
dence indicating that the MPN plays an im- gether with the adjacent substantia innomi-
portant role in mediating thermoregulatory, nata (often referred to collectively as the sub-
ingestive and reproductive behaviours, all of pallidal region) participate in the regulation
which involve foraging behaviour and thus of locomotor activity [1338, 1339, 1343].
locomotion [1337]. In this context it is of par- Turning to the epithalamus, the connec-
ticular interest that the MPN projects directly tions of the medial and lateral habenular nu-
to two functionally defined "locomotion re- clei may now be considered.
gions", the caudal part of the zona incerta, The medial habenular nucleus (MHN) re-
which appears to lie within the" subthalamic ceives, by way of the stria medullaris, fibres
locomotor region ", and the pedunculopon- originating from the medial septal-diagonal
tine nucleus, which lies within the" mesence- band complex [1331]. A relatively less numer-
phalic locomotor region" [1339]. The projec- ous set of afferents reaches the MHN from
tion from the MPN to the ventral tegmental the mesencephalic raphe nuclei and from the
area may also influence locomotor activity, ventral part of the periaqueductal grey mat-
since this area sends a massive dopaminergic ter [1317]. The efferents of the MHN termi-
projection to the nucleus accumbens (which nate exclusively in the interpeduncular nucle-
represents a nodal point for the relay oftelen- us [510, 854]. The efferent fibres of the inter-
cephalic limbic information to diencephalic peduncular nucleus, which project massively
and mesencephalic sites involved in somato- to the dorsal raphe and central superior nu-
motor control [1339]). clei form a further link in the "medial haben-
ular path" (Fig. 155B; [447, 841, 977]).
The lateral preoptic nucleus (LPN) is usu- The lateral habenular nucleus (LHN) re-
ally considered to represent the most rostral ceives its afferents principally from the nuclei
part of the diffusely organised lateral preopti- of the diagonal band, the lateral preopticohy-
co-hypothalamic cell continuum. Afferents to pothalamic area, the substantia innominata
the LPN have been reported [693] to origi- and the medial pallidal segment, whereas the
nate from the central amygdaloid nucleus, ventral tegmental area, the mesencephalic
the bed nucleus of the stria terminalis, the raphe nuclei and the ventral part of the peria-
lateral septal nucleus, the nuclei of the diago- queductal grey matter give rise to ascending
nal band, the nucleus accumbens, the medial input to this nucleus [509, 510, 871, 983,
1055, 1317, 1388]. Fibres originating from the the superior colliculus, the periaqueductal
LHN descend in the habenulointerpeduncu- grey matter, the red nucleus, the mesence-
lar tract, bypass the interpeduncular nucleus phalic and rhombencephalic reticular forma-
and terminate in various mesencephalic tion, the pedunculopontine nucleus, the nu-
centres, including the pars compacta of the cleus raphes magnus, the inferior olivary
substantia nigra, the ventral part of the peri a- complex and the spinal cord [1122, 1143,
queductal grey matter, the dorsal raphe and 1295,1335,1479].
central superior nuclei and the mesencephalic The peripeduncular nucleus may be consid-
reticular formation [8, 509, 510, 983]. ered to represent the most caudal part of the
The functional role of the habenular nuclei zona incerta [1130]. This nucleus is bidirec-
is not clear. Judging from their fibre connectionally connected with the ventromedial hy-
tions, they represent processing stations in pothalamic nucleus and has also been re-
strictly separate medial and lateral habenular ported to project to the amygdala, the medial
paths. The medial habenular path, which is preoptic and the lateral hypothalamic areas
synaptically interrupted in the medial habenu- [107, 692, 693, 889, 1250].
lar and interpeduncular nuclei, conveys in- The zona incerta is usually regarded as
formation from medial septal and ultimately a rostral extension of the brain stem reticular
from hippocampal antecedents. In contrast, formation [1235]. On the basis of anterograde
the lateral preopticohypothalamic area and tracer experiments, Roger and Cadusseau
the globus pallidus provide the LHN with [1140] recently distinguished a ventral and a
the majority of its afferents. The efferents of dorsal sub-zone within the zona incerta. Ac-
the LHN bypass the interpeduncular nucleus, cording to their observations, the ventral
but the medial and lateral habenular paths sub-zone receives its afferents principally
converge upon the mesencephalic raphe nu- from somesthetic centres, such as the soma-
clei. To what extent these two afferent sys- tosensory cortex, the trigeminal complex and
tems interact with each other at the level of the dorsal column nuclei, and from the supe-
the raphe nuclei is unknown at present. The rior colliculus and the cerebellum, whereas
projection from the LHN to the pars com- in the dorsal sub-zone a limbic input from
pacta of the substantia nigra presumably the cingulate cortex and the ventromedial nu-
closes a nigro-striato-pallido-habenulo-nigral cleus prevails. The zona incerta is thought
circuit [510]. to playa crucial role in the neuronal mecha-
The zona incerta, which forms part of the nisms that control ingestive behaviour [1123].
subthalamic region, derives its input from the The anterior hypothalamic nucleus and sur-
medial prefrontal, cingulate and somatosen- rounding areas receive afferents from the
sory cortices [1140, 1235], the central amyg- ventral subiculum, the medial amygdaloid
daloid nucleus [1140], the substantia innomi- nuclei, the lateral septal nucleus, the dorso-
nata [240], the ventromedial hypothalamic medial and ventromedial hypothalamic nu-
nucleus [722], and from a variety of struc- clei, the mesencephalic central grey matter,
tures situated in the brain stem, such as the the mesencephalic raphe nuclei, the lateral
superior colliculus, the periaqueductal grey parabrachial nucleus, the locus coeruleus, the
matter, the mesencephalic reticular forma- noradrenergic areas A1 and A2, and the soli-
tion, the medial and lateral parabrachial nu- tary nucleus [107,825,1261,1298,1338]. The
clei, the deep cerebellar nuclei, the sensory anterior hypothalamic area has primarily
trigeminal nuclear complex and the dorsal short connections to the adjoining medial
column nuclei [1140, 1235, 1434]. Efferents preoptic nucleus, the dorsomedial and ven-
from the zona incerta terminate in a large tromedial hypothalamic nuclei, the infundi-
number of thalamic centres, the posterior bular nucleus and the lateral hypothalamic
and dorsomedial hypothalamic nuclei and area [254, 1194, 1196]. However, it also has
the lateral and dorsal hypothalamic areas, some more distant projections, including as-
cending fibres which are distributed to the The supraoptic and paraventricular hypo-
lateral part of the septum [254, 1194], and thalamic nuclei. It has long been known that
descending fibres which project (via the me- the neurohormones vasopressin and oxytocin
dial forebrain bundle) to the dorsal prema- are produced in the supraoptic (SO) and
millary nuclei, the posterior hypothalamic paraventricular hypothalamic (PVH) nuclei,
area, the periaqueductal grey matter, the ros- two centres situated in the anterior hypotha-
tral raphe nuclei, the ventral tegmental area, . lamus [80]. Large neurons situated in these
the interpeduncular nucleus and to several nuclei synthesise the hormones mentioned
rhombencephalic centres [254, 534, 1194]. and give rise to axons which descend through
According to Holstege [534], the fibres pass- the infundibular stalk to the posterior pitui-
ing to rhombencephalic centres split up in tary, thus forming the supraoptico- and para-
the caudal mesencephalon, into separate me- ventriculohypophyseal tracts (Fig. 192). The
dial and lateral streams. The medial fibre hormones are transported along the axons
stream, which is mainly derived from the me- of these pathways and released into the blood
dial parts of the anterior hypothalamic area, vessels of the neurohypophysis. Vasopressin
descends in the medial part of the rhomb- is responsible for antidiuresis, promoting
encephalic medial tegmentum, distributing reabsorption of water by the kidney; oxyto-
fibres to the raphe nuclei and to the adjacent cin stimulates smooth muscle in the uterus
medial reticular formation. The lateral fibre and mammary glands to contract.
stream, which is mainly derived from the lat- The PVH contains, in addition to the large
eral parts of the anterior hypothalamic area, neurosecretory elements discussed above, nu-
descends in the lateral tegmental field and merous smaller peptidergic elements. The lat-
distributes fibres to the parabrachial nuclei, ter are concentrated in the caudal portion of
the locus coeruleus, the nucleus subcoeruleus, the nucleus and can be subdivided into two
the micturition-coordinating region, the lat- sub-populations. The elements of the first
eral part of the reticular formation (including sub-population constitute a parvocellular
the superficial ventrolateral reticular area) neurosecretory system, the fibres of which
and the dorsal vagal complex. pass to the external lamina of the median
The anterior hypothalamic area has been eminence, where their terminals contact the
implicated in the regulation of temperature hypophyseal portal capillaries [47, 1269].
[1199], respiration [664] and cardiovascular This sub-population comprises, apart from
functions [389]. According to Fuchs and col- vasopressinergic neurons, numerous elements
laborators [365, 366] the feline anterior hypo- containing corticotropin releasing factor
thalamic area integrates affective defense (CRF). There is evidence indicating that
behaviour. These authors pointed out that these CRF -containing cells, via their projec-
the autonomic functions mentioned above tion to the median eminence, constitute the
are all important for the expression of affec- major pathway in the hypothalamic expres-
tive defense behaviour and that the input to sion of motivational drives and the emotional
the anterior hypothalamic area from the state. The CRF, once liberated in the hypo-
amygdala and the septum may well provide physeal portal system, activates ACTH secre-
the basis through which modulation of this tion by the anterior pituitary, which in turn
behaviour is achieved. Moreover, Fuchs and leads to corticosteroid release from the adre-
collaborators [365, 366] provided physiologi- nal glands [825]. The elements of the second
cal evidence indicating that the principal sub-population give rise to a descending pro-
pathway subserving affective defense behav- jection consisting of thin unmyelinated,
iour in the cat goes from the ventromedial mainly oxytocinergic fibres, which innervate
hypothalamic nucleus, via the anterior hypo- primarily a number of autonomic centres in
thalamic area to the periaqueductal grey mat- the brain stem and the spinal cord [182, 554,
ter. 1322, 1553]. The fibres of this descending
PVHprojection follow two different path- nuclei; the lateral hypothalamic area [1195];
ways, a dorsomedial one and a ventrolateral the lateral parabrachial nucleus [107, 1387]
one [534, 824, 1340]. The dorsomedial path- and some catecholaminergic centres in the
way descends through the central grey matter lower brain stem. The latter include the nor-
and innervates that structure, the Edinger- adrenergic locus coeruleus and Ai and A2
Westphal nucleus, the lateral parabrachial cell groups and the adrenergic C1 and C2
nucleus and the locus coeruleus. In the rostral cell groups [107, 1181, 1340] (Fig. 154). It is
rhombencephalon, the dorsomedial pathway important to note that the various afferent
shifts laterally to join the ventrolateral path- projections to the PVH tend to terminate se-
way [824], which descends through the me- lectively on one of the functional neuronal
dial forebrain bundle initially, then moves populations within the nucleus, i.e. the large
laterally in the midbrain, where it covers the vasopressinergic neurons, the large oxytocin-
lateral part of the substantia nigra. In the ergic neurons, the small hypophysiotrophic
pons this pathway moves ventrally and neurons and the small, caudally projecting
reaches the ventral part of the lateral reticu- " autonomic" neurons. Thus, the projection
lar formation, through which it passes cau- from the Ai region provides the route
dally to enter the dorsolateral funiculus of through which the control of vasopressin se-
the spinal cord. Fibres belonging to this path- cretion by visceral afferent stimuli is achieved
way supply the superficial ventrolateral retic- [1207, 1340].
ular formation, including the noradrenergic The SO projects exclusively to the posteri-
Ai and the adrenergic C1 regions. At the or pituitary. Its afferents closely correspond
level of the inferior olive, the pathway issues to those to the PVH [1340, 1386]. The SO
numerous fibres which arch dorsomedially to and PVH are extremely vascular. It has been
innervate the dorsal motor nucleus of the va- frequently suggested that humoral factors, in
gus nerve and the nucleus of the solitary addition to the neural inputs described
tract, including the noradrenergic A2 and the above, may influence the activity of the cells
adrenergic C2 regions [254, 534, 824, 1190, in these nuclei. For one group of hormones,
1322, 1335]. The fibres attaining the spinal the sex steroids, a direct humoral influence
cord project to the marginal zone of the dor- of this type, on the cells in the SO and PVH,
sal horn, the area surrounding the central ca- has been firmly established [1340].
nal (i.e. area X of Rexed), preganglionic cell The primary functions of the PVH, and
groups of both the sympathetic and parasym- the SO are directly related to the magnocellu-
pathetic divisions of the autonomic system lar neurosecretory projections to the posteri-
[472, 554, 824, 1190, 1335, 1393] and, or pituitary. In addition, however, the PVH
remarkably, the somatomotor nucleus of has a second component which, via the medi-
Onuf [534]. The last mentioned centre is situ- an eminence, participates in regulating the
ated in segments S1-S2 and innervates the activity of the anterior pituitary and a third
pelvic floor musculature, including the striat- component that modulates a number of auto-
ed muscle sphincters of the urethra and anus nomic and behavioural processes [1340].
[1201]. Thus, it is known that the parvocellular part
Afferents to the PVH originate from: the of the PVH participates in the regulation of
bed nucleus of the stria terminalis [1331, feeding and drinking behaviour [825, 1337]
1340]; the preoptic region [1320], including and also plays an important role in cardio-
the organum vasculosum laminae terminalis vascular control [1207]. The concentration of
(OVLT) and the subfornical organ [916, the three functionally distinct centres men-
1249]; several nearby hypothalamic regions, tioned above within the PVH should be
including the suprachiasmatic [107, 1327], viewed as the morphological expression of
anterior hypothalamic [1194], ventromedial the role which this nucleus plays in the inte-
hypothalamic [1193] and infundibular [644] gration of behavioural and autonomic func-
tions with their endocrine correlates. Finally, put of the SCH may be best viewed as a two-
it may be mentioned that the projection of stage process in concert with the subparaven-
the PVN to the superficial zone of the spinal tricular zone.
dorsal horn may be involved in pain control
mechanisms [534]. The dorsomedial nucleus of the hypothala-
The small suprachiasmatic nucleus (SCH) , mus (DMH) constitutes, together with the
which is situated in the rostroventral part of ventromedial hypothalamic nucleus (VMH),
the hypothalamus, may be considered as the the nucleus infundibularis and the adjacent
endogenous clock of the brain. It is critically segment of the lateral hypothalamic area
involved in the generation and entrainment (LHA) the intermediate or tuberal region of
of a number of circadian rhythms, including the hypothalamus.
patterns of general activity (sleeping/wak- The rather diffusely organised DMH re-
ing), feeding and drinking behaviour and ceives afferents from the central amygdaloid
hormonal secretion [1123, 1154]. The visual nucleus, the bed nucleus of the stria termina-
system participates in the regulation of these lis, the nucleus accumbens, the lateral septal
behavioural and endocrine rhythms via a di- nucleus, the organum vasculosum of the la-
rect retinohypothalamic projection, which mina terminal is (OVLT), the ventromedial
terminates in the SCH [251, 939, 944, 1373], hypothalamic nucleus, the LHA, the peri-
as well as via a pathway originating from aqueductal grey matter, the lateral parabra-
the ventral part of the lateral geniculate body chial nucleus, the nucleus of the solitary tract
[1332]. Other inputs to the SCH arise from and from the noradrenergic A1 and A2 areas
the subiculum [849] and from the mesence- [692, 825, 1125, 1206, 1334, 1353].
phalic raphe nucleus [67, 134]. The DMH sends dense projections to the
The SCH imposes circadian organisation subfornical organ, the OVL T and the parvo-
on the functional activity in many parts of cellular part of the paraventricular hypotha-
the brain. The efferent channels which may lamic nucleus [825]. Less-dense projections
be implicated in the mediation of these influ- reach the VMH, and LHA and a number
ences include projections to the medial pre- of brain-stem centres, including the ventral
optic and lateral septal nuclei, the dorsome- tegmental area, the periaqueductal grey mat-
dial hypothalamic nucleus, the lateral hypo- ter, the parabrachial nuclei, the rhombence-
thalamic area, the peri ventricular thalamic phalic lateral reticular zone, the ambiguous
nucleus and the ventral lateral geniculate nu- nucleus, the dorsal motor nucleus of the va-
cleus [107, 143, 1290, 1327, 1482]. However, gus and the area postrema [825, 1368].
by far the densest terminal field arising from The DMH is one of the" feeding-related
cells in the SCH ends in a nearby periventri- nuclei" of the hypothalamus. Taking the pat-
cular area extending medially from the ante- tern of its connections into consideration,
rior hypothalamic nucleus to the paraventri- Luiten and collaborators [825] made the fol-
cular nucleus [1482]. This periventricular lowing suggestions concerning the possible
area, which has been designated by Watts functional roles of this centre:
and colleagues [1482] as the subparaventricu-
lar zone, has been found to project to essen- 1. The strong and partially reciprocal con-
tially the same regions as the SCH, but much nections with a number of circumventricular
more densely, and also to send fibres back organs indicate that the DMH may control
to the SCH. In addition, the subparaventri- the access to nervous tissue of blood-borne
cular zone innervates much more of the later- humoral factors.
al septal nucleus than the SCH and also sends 2. The reciprocal connections with the
fibres throughout the length of the periaque- VMH and the LHA may indicate that the
ductal grey matter [1481, 1482]. According DMH fulfills a bridge function between
to Watts and colleagues [1481, 1482], the out- the former two centres, which, according to
the so-called "hunger-satiety dual-centre and collaborators [825] emphasised that the
theory" (see below), represent antagonists in input to the LHA is only small. the descend-
the control of feeding. ing projections that originate from the VMH
3. The strong projection to the parvocellu- follow two main pathways, a large dorsal one
lar part of the paraventricular hypothalamic and a smaller ventral one. The dorsal path-
nucleus suggests direct access of the DMH way follows the periventricular system as far
to the neuroendocrine system. as caudally as the locus coeruleus. This bun-
4. Direct as well as indirect descending dle densely innervates the periaqueductal
pathways connect the DMH with the ambig- grey matter, which is the major target of de-
uous nucleus and with the dorsal motor nu- scending projections from the VMH [825].
cleus of the vagus, groups of parasympathetic Minor projections from the dorsal pathway
motor cells which innervate the endocrine reach the mesencephalic reticular formation,
pancreas and influence insulin secretion. including the pedunculopontine nucleus and
the locus coeruleus [1196]. The second VMH
The VMH receives strong inputs from the pathway descends in the most ventral part
two major limbic territories in the telence- of the hypothalamus and distributes fibres
phalon, i.e. the hippocampal formation and to the ventral tegmental area, the peripedun-
the amygdaloid complex. The hippocampal cular nucleus and, like the dorsal pathway,
input arises from the ventral subiculum, to the mesencephalic reticular formation.
whereas the amygdalar input principally ar- Experimental data suggest that the VMH
ises from the medial amygdaloid nucleus. plays an important role in the initiation of
Both groups of direct projections are presum- agonistic behaviour and in the control of
ably augmented by indirect ones, because the feeding. It has long been known that lesions
hippocampus projects strongly to the lateral in the area of the VMH produce extreme rage
septal nucleus, which in turn supplies the reactions, but this behaviour cannot be sim-
VMH, and the amygdala sends numerous ply explained as resulting from release of in-
fibres to the bed nucleus of the stria termina- hibitory influences, because stimulation of
lis, which also projects to the VMH [107, 692, the VMH, particularly of its ventrolateral
721, 825, 873, 1330]. Major projections to part, may also elicit aggressive behaviour
the VMH also arise from the peri peduncular [801]. It has already been mentioned that ac-
and lateral parabrachial nuclei. Numerous cording to Fuchs and collaborators [365, 366]
other centres provide less extensive inputs to an important pathway subserving defense
the VMH. These include the supraoptic, me- behaviour passes from the VMH, via the an-
dial preoptic, anterior hypothalamic and dor- terior hypothalamic area, to the periaqueduc-
somedial hypothalamic nuclei, the parvocel- tal grey matter. As regards the control of
lular part of the paraventricular hypothalam- feeding, lesions in the VMH produce hyper-
ic nucleus, the periaqueductal grey matter, phagia and obesity, whereas LHA lesions
the mesencephalic raphe nuclei, the locus produce aphagia; electrical stimulation on
coeruleus and the nucleus of the solitary tract the other hand, elicits feeding behaviour
[107, 692, 825, 873]. from the LHA and feeding suppression from
The VMH has rather widespread ascend- the VMH. On the basis of these observations
ing, intrahypothalamic and descending pro- the tuberal part of the LHA has been desig-
jections [27, 722, 825, 1196]. Ascending fibres nated as the" feeding centre" and the VMH
supply the preoptic region, the bed nucleus as the" satiety centre" [486]. Because, firstly
of the stria terminalis, the lateral septal nucle- direct connections between these two antago-
us and the central amygdaloid nucleus. Intra- nistic feeding control centres are only weakly
hypothalamic projections have been traced developed and, secondly both are reciprocal-
to the anterior, dorsomedial, and premamill- ly connected with the DMH, Luiten and col-
ary nuclei and to the LHA, although Luiten laborators [825] conjectured that the latter
may act as a bridge between the VMH and benular nucleus, the LHA, the mesencephalic
the LHA. Finally, it is worthy of mention and pontine central grey matter, the dorsal
that according to Luiten and colleagues [825] tegmental nucleus, the locus coeruleus, the
the large projection from the VMH to the nucleus of the solitary tract and the lateral
periaqueductal grey matter may well repre- part of the medullary reticular formation
sent the first link in a preponderantly ortho- [107, 825]. Experimental data on the efferent
sympathetic pathway which, via synaptic re- projections of the posterior hypothalamic nu-
lays in the ventromedial and ventrolateral cleus per se are to our knowledge not avail-
portions of the medullary reticular formation able. However, Veazey and colleagues [1433]
and in the intermediolateral cell columns in have reported that this nucleus, together with
the thoracolumbar cord, innervates (among the posterior part of the LHA, appears to
other structures) the pancreas and the adre- contribute substantially to the ascending
nal medulla. components of the medial forebrain bundle
The small-celled infundibular nucleus (or and (through its descending projection to the
arcuate nucleus) is located in the most ventral mesencephalic and pontine central grey mat-
part of the tuberal region of the hypothala- ter) to the mesencephalic reticular formation,
mus (Fig. 192). It has already been men- the central superior nucleus of the raphe
tioned that populations of neurons situated complex and the locus coeruleus. It is known
within this cell mass produce a number of that the posterior part of the hypothalamus
neurohormones which regulate the synthesis is involved in the generation of emotional
and release of the various adenohypophyseal stress and the regulation of body tempera-
hormones. The axons of the infundibular ture, particularly the control of heat produc-
neurons convey the regulatory neurohor- tion and conservation. However, there is no
mones via the tuberoinfundibular tract to the evidence that these functions can be specifi-
median eminence, where they are released in cally attributed to the posterior hypothalam-
the capillary loops of the hypothalamohypo- ic nucleus.
physeal portal system (Figs. 192 and 197). By The portion of the medial hypothalamic
the vessels of this local system the regulatory zone situated directly caudal to the posterior
factors are transported to the glandular pitui- hypothalamic nucleus contains two rather
tary. It should be mentioned that the infundi- small cell masses which are known as the dor-
bular nucleus, apart from numerous pepti- sal and ventral premamillary nuclei. The fibre
dergic elements, also contains a substantial connections of these centres are imperfectly
dopaminergic population. The elements of known. The following data on these connec-
this so-called A12 group project to the medi- tions are derived from the recent work of
an eminence and are known to exert, via the Luiten and colleagues [825]. The dorsal pre-
hypothalamohypophyseal portal system, an mamillary nucleus receives input from the
inhibitory action on hormone release from ventromedial hypothalamic nucleus and, par-
prolactin-secreting cells in the anterior pitui- ticularly, from the LHA. The ventral prema-
tary lobe [941]. Little is known about the af- miliary nucleus (PMV) receives a dense and
ferents to the infundibular nucleus. In a re- highly specific projection from the medial
cent study using a retrograde tracer tech- amygdaloid nucleus, and some fibres orig-
nique, Gruber and colleagues [458] reported inating from the dorsomedial hypothalamic
the existence of projections from the subforn- nucleus have also been traced to this centre.
ical organ, the organum vasculosum of the Efferents of the PMV pass to the ventral half
lamina terminalis (0 VL T) and the mesence- of the ventromedial hypothalamic nucleus
phalic raphe nuclei to this centre. and to the parvicellular portion of the para-
The posterior hypothalamic nucleus has ventricular nucleus. Studies of lesions in the
been reported to receive afferents from the medial amygdala and in the PMV indicate
nuclei of the diagonal band, the lateral ha- that these interrelated nuclei participate in
a brain substrate for the control of offensive MFB to traverse the medial septal- diagonal
and defensive behaviours [825]. band complex and enter the fornix and the
The lateral hypothalamic area (LHA) cingulate bundle, from which it is distributed
which, as has already been mentioned, is tra- to the hippocampal formation, particularly
versed throughout its length by the medial to the dentate gyrus, and to medial cortical
forebrain bundle (MFB), receives (mainly via fields, respectively. The lateral pathway runs
that pathway) substantial projections from through the lateral part of the MFB, then
the prelimbic and infralimbic cortices (areas arches laterally to enter the external capsule,
32 and 25, respectively), the central amygda- from which it is distributed to lateral cortical
loid nucleus, the olfactory tubercle, the nucle- fields [1188]. The neurons projecting to the
us accumbens, the lateral septal nucleus, the substantia innominata and the cerebral cor-
peripeduncular nucleus, the periaqueductal tex tend to cluster in the caudal portion of
and pontine central grey matter, the mesence- the LHA [1188], whereas the elements sup-
phalic raphe nuclei, the lateral parabrachial plying the medial septal - diagonal band
nucleus and the locus coeruleus. Other affer- complex and the lateral preoptic area are
ents to the LHA come from the medial and mainly situated in the rostral part of the
orbital regions of the prefrontal cortex, the LHA [1195]. The rostral part of the LHA
subiculum, the bed nucleus of the stria ter- also projects, via the stria medullaris, to the
minalis, the vertical limb of the nucleus of periventricular thalamic nuclei and the lateral
the diagonal band, the medial and lateral pre- habenular nucleus [1184, 1195]. Shen [1237]
optic nuclei, the dorsomedial hypothalamic described fibres passing from the LHA to the
nucleus, the lateral habenular nucleus, the internal layer of the median eminence and
ventral tegmental area and the caudal part the posterior lobe of the pituitary.
of the solitary tract nucleus [107, 555, 693, Large numbers of fibres, originating main-
825, 986, 1125]. ly from the tuberal and caudal portions of
As regards the efferent connections of the the LHA, descend to the brain stem, where
LHA the following general statements can they contribute to the dorsal longitudinal fas-
be made: cicle of Schutz, as well as to the medial and
lateral caudal extensions of the MFB [534,
1. All parts of the LHA contribute ascending
553, 557, 825, 1184, 1195, 1237, 1412]. The
and descending fibres to the MFB [1195].
fibres descending in the periventricular bun-
2. Most of the intrahypothalamic output of
dle of Schutz terminate in the periaqueductal
the LHA is aimed at other parts and re-
and pontine central grey matter. The fibres
gions of the LHA itself [825].
following the medial caudal extension of the
3. The spread of lateral hypothalamic effer-
MFB terminate in the ventral tegmental area,
ents is extraoridinarily wide and extends
the raphe nuclei and in the adjacent parts
from the neocortex to the spinal cord.
of the medial reticular formation. Those
4. Cells at different levels of the LHA may
which descend in the lateral caudal extension
have differential projections [1195].
of the MFB project to a variety of grisea,
Ascending efferents from the LHA enter among which are the mesencephalic central
the MFB and pass to the substantia innomin- tegmental field, including the pedunculopon-
ata [1195], the lateral preoptic area [1195, tine tegmental nucleus, the medial and lateral
1237], the medial septal - diagonal band parabrachial nuclei, the locus coeruleus and
complex [1195,1237], the central amygdaloid the area subcoerulea, the rhombencephalic
nucleus, which is reached via the ventral lateral reticular formation, including the su-
"amygdalofugal" pathway [27], and the cere- perficial ventrolateral reticular area and the
bral cortex [683, 691, 1188]. The fibres pass- catecholaminergic centres embedded therein,
ing to the cortex follow two different path- the ambiguous nucleus and the dorsal vagal
ways: a medial pathway which leaves the complex. About half of the hypothalamic
neurons which project directly to the spinal and sympathetic as well as parasympathetic
cord are contained in the LHA. These fibres cell groups via consecutive relays in the peri-
terminate mainly in the spinal central grey aqueductal grey matter and the lateral part
matter (Rexed's lamina X) and in the thora- of the reticular formation.
columbar intermediolateral column [534,
552, 1335, 1393]. The Ventral Tegmental Area (VT A), or ven-
Interestingly, the LHA has been recently tral tegmental nucleus of Tsai, is situated in
reported to be the main source of a hypotha- the medioventral part of the midbrain and
lamocerebellar projection. The fibres of this may be regarded as a caudal extension of
projection have been found to traverse the the LHA. This rather ill-defined cell mass
periaqueductal grey matter to reach the deep contains a substantial population of dopa-
cerebellar nuclei as well as the cerebellar cor- minergic neurons, which is known as the Al0
tex, particularly the vermis and the flocculus. group [279]. Ascending afferents to the VT A,
The fibres distributed to the cortex terminate originate from the locus coeruleus and other
neither as mossy fibres nor as climbing fibres, noradrenergic cell groups including Al, A5
but display the multilayered mode of termi- and A 7, the dorsal raphe and central superior
nation previously described for monoamin- nuclei, the deep cerebellar nuclei, and the me-
ergic axons [299, 300, 301,468]. dial and lateral parabrachial nuclei. These
With regard to the functions of the LHA fibres probably attain the VTA mainly by
it will be clear that, from a connectional point way of the mamillary peduncle [1084, 1252].
of view, the cells of this area are ideally Descending projections in the MFB appear
placed to collect and process much of the to provide the major source of inputs to the
ascending and descending information in the VT A. These fibres originate mainly from the
MFB [986, 1195]. It has long been held that anterior limbic cortex (areas 24 and 32) [89,
the functional state of the medial hypotha- 1084], the bed nucleus of the stria terminalis
lamic nuclei is mainly regulated by the infor- and the lateral septal nucleus [1331], the me-
mation sampled from this complex longitudi- dial preoptic nucleus [1320], the ventromedial
nal conduction channel through the media- hypothalamic nucleus [825, 1193], the lateral
tion of short transverse fibres interconnecting preopticohypothalamic zone [1195,1320] and
the medial and lateral hypothalamic cell the lateral habenular nucleus [1084, 1252].
zones. The" dual-centre" model for the regu- The efferent connections of the VT A have
lation of food intake, already alluded to, been studied by Beckstead and colleagues
should be placed in the context of this general [94], Simon and collaborators [1252] and
concept. However, it is now known that the Swanson [1325]. According to Swanson, the
medially placed nuclei of the hypothalamus output of the VT A may be divided into three
have a great number of direct long efferent major components. The first and most sub-
and afferent connections with many extra- stantial component ascends through the
hypothalamic centres. Moreover, Luiten and MFB to innervate a variety of telencephalic
collaborators [825] have recently presented regions, including the anterior cingulate (area
evidence which suggests that the LHA parti- 24) and prelimbic (area 32) cortices, the ento-
cipates as an entity with its own characteris- rhinal cortex, the amygdala (particularly the
tics in hypothalamic functions and is only central nucleus), the bed nucleus of the stria
partially dominated by the longitudinal terminalis, the accumbens nucleus and adja-
stream of the MFB. These authors emphas- cent parts of the striatum, the lateral septal
ised the role of the LHA as an output station nucleus and the nuclei of the diagonal band.
for the autonomic nervous system, reaching The second component ascends dorsally
preganglionic parasympathetic motor ele- from the VTA to innervate some cell groups
ments directly by its projections to the am- in the medial part of the thalamus and the
biguous nucleus and the dorsal vagal nucleus, lateral habenular nucleus. The third compo-
nent descends through the midbrain to inner- premamillary nuclei as well as the LHA [99,
vate the periaqueductal grey matter, the 454,649, 722, 825, 849,1193,1194], the supe-
para brachial nuclei, the central superior nu- rior colliculus [649, 849], the mesencephalic
cleus of the raphe and the locus coeruleus. reticular formation [849], the substantia nigra
Swanson [1325] found that each of the pro- pars reticulata [99, 454,551], the dorsal raphe
jections from the VTA to specific regions in and central superior nuclei [99, 855], the locus
the telencephalon, the diencephalon and the coeruleus [99], the medial and lateral para-
brain stem arise from essentially separate brachial nuclei [99, 855], the pontine and
populations of cells, and that the proportion medullary reticular formation [99, 855] and
of dopaminergic cells in each of these popula- the spinal cord [887]. It is noteworthy that
tions is different, and varies from over 80% the PAG contains large numbers of short,
(e.g. in cells projecting to the accumbens nu- longitudinally arranged intrinsic fibres [99,
cleus) to less than 1 % (in cells projecting to 851].
the lateral habenular nucleus and the locus Somewhat schematising, the efferents of
coeruleus). the PAG can be subdivided into the follow-
The VT A has been shown to be important ing four groups: (a) ascending fibres, passing
in the regulation of motor, motivational and rostrally and rostroventrally in a periventri-
affective processes. Swanson and Mogenson cular position, (b) fibres passing to the ros-
[1337] have presented evidence that a multi- tromedial tegmentum, some of which con-
synaptic pathway passing from the medial tinue rostrally in the medial forebrain bundle,
preoptic area to the nucleus accumbens, by (c) fibres fanning out radially in the mid-
way of the VTA, is involved in the regulation brain, which together constitute Weissche-
of thirst. Pijnenburg and colleagues [1090] del's "radiatio grisea tegmenti" [1491], and
demonstrated that activation of the projec- (d) fibres descending to the rhombencepha-
tion from the VT A to the accumbens nucleus lon and spinal cord. The ascending periven-
produces enhanced locomotor activity. Fi- tricular fibres constitute the direct rostral ex-
nally, it should be mentioned that hyperacti- tension of the longitudinal dorsal fasciculus
vity of the VTA and its dopaminergic out- of Schutz (Fig. 191). These fibres terminate
flow to the accumbens nucleus and other te- in the zona incerta, the reticular thalamic nu-
lencephalic limbic structures may be impor- cleus, the midline and intralaminar thalamic
tant in the pathophysiology of schizophrenia nuclei, the medial and lateral preoptic nuclei
[119, 1297). and in various hypothalamic centres, includ-
The periaqueductal grey matter (PAG) or ing the anterior, ventromedial and posterior
central mesencephalic griseum is a conspicu- nuclei [234,317,850]. The PAG fibres pass-
ous midline structure which surrounds the ce- ing to the rostromedial tegmentum terminate
rebral aqueduct from the level of the para- in the ventral tegmental area and, via the
brachial nuclei to the posterior commissure MFB, in the LHA [234]. Eberhart and colla-
(Figs. 97-102). This centre, which consists borators [317] recently reported, in the rat,
mainly of rather densely packed small neu- the presence of a strong projection pathway
rons, receives afferents from a variety of which, after having emanated from the PAG,
structures, including: the premotor (areas 6 ascends through the MFB and distributes
and 8), prefrontal (areas 9, 10 and 13), anteri- fibres to numerous telencephalic regions,
or cingulate (area 24) and insular cortices [77, among which are the medial and lateral sep-
99,156,238,474,649,849,991]; the amygda- tal nuclei, the nuclei of the diagonal band,
la (basal lateral and central nuclei [649, 849]; the accumbens nucleus, the olfactory tuber-
the medial preoptic area [34, 99, 855]; the cle, the bed nucleus of the stria terminalis,
zona incerta [551, 649, 825, 849, 855]; the the central amygdaloid nucleus and the fron-
hypothalamus, particularly the anterior, ven- tal cortex. The radially directed PAG fibres
tromedial (a massive projection!) and dorsal can be traced to the superior colliculus and
to the mesencephalic reticular formation, in- tion of a brain area extending from the me-
cluding the cuneiform nucleus [234, 850, 851]. dial hypothalamus into the mesencephalic
The fibres originating from the PAG which central grey matter. The caudal part of the
descend to the rhombencephalon pass PAG contains a small population of neurons,
through its tegmental field and distribute the excitation of which elicits both behav-
fibres to the locus coeruleus, the lateral para- ioural and cardiovascular components of the
brachial nucleus, the nuclei raphes magnus defense reaction [878]. It has already been
and pallidus, the pontine and medullary me- mentioned that Fuchs and colleagues [365,
dial reticular formation, the ambiguous nu- 366] have adduced morphological and physi-
cleus and the portion of the lateral reticular ological evidence suggesting that an impor-
formation surrounding that nucleus [239, tant circuit subserving feline affective defense
825, 851]. A small number of PAG fibres behaviour includes an ascending component
enters the spinal cord and passes through the from the ventromedial nucleus to the anterior
lateral funiculus to the intermediolateral zone hypothalamus and a descending component
of the upper thoracic segments [825]. The from the anterior hypothalamus to the PAG.
PAG plays an important role in a variety The authors emphasised that the anterior hy-
of functions and behavioural patterns, pothalamus, which receives input from key
among which nociception, defense reactions, limbic structures such as the amygdala and
vocalisation, the lordosis reflex that forms the septal area, may well sub serve an inte-
part of female reproductive behaviour and grating function with respect to affective de-
the regulation of pancreatic and adrenal se- fense behaviour. It is worth mentioning in
cretion may be mentioned. this context that Luiten and collaborators
As regards nociception, the PAG has been [825] have stressed the potential importance
demonstrated to be an essential link in a de- of a somewhat different set of projections
scending pain-control system which ultimate- leading to the PAG as an essential way-sta-
ly inhibits noxiously evoked activity in spinal tion in the output channel to somatic and
dorsal horn neurons (for review see Basbaum autonomic centres involved in agonistic
and Fields [83]). The nucleus raphes magnus, behaviour. In their opinion this circuitry in-
which receives a strong projection from the cludes strong projections from the medial
PAG, serves as a relay between the PAG and amygdaloid nucleus to the ventromedial hy-
the spinal cord neurons. pothalamic and premamillary nuclei, the lat-
Threatening or stressful stimuli may evoke ter centres projecting in turn to the PAG.
a characteristic pattern of behavioural and The PAG also plays an important role in
autonomic responses which is commonly des- the vocal expression of emotion. Electrical
ignated as the affective defense reaction. In stimulation of the PAG readily induces vo-
cats, the somatomotor components of this re- calisation, while lesions in this structure abol-
action include ear retraction, hissing, growl- ish it [648]. Fibres descending from the PAG
ing and striking with the forepaw. The auto- to the ambiguous nucleus, i.e. the centre
nomic response pattern consists of pupillary which innervates the striated muscle fibres
dilatation, piloerection, increase in the rate of the pharynx and the vocal-cord muscles
and amplitude of respiration and a sympa- of the larynx, are considered to convey the
thetic cardiovascular reaction comprising ac- impulses which elicit this vocalisation [649].
celeration of heart rate, elevation of blood The lordosis reflex is elicited in sexually
pressure and vasoconstriction in cutaneous, receptive female rodents by somatosensory
renal and splanchnic beds, but vasodilation stimuli which pass along spinomesencephalic
in skeletal muscles. The classical work of fibres, forming part of the anterolateral sys-
Hess and Brugger [516], Hunsperger [577, tem, to the PAG. The efferent limb of the
578] and others has shown that this defense reflex arc is presumably an indirect projec-
response can be elicited by electrical stimulation which passes from the PAG, via a relay
in the medullary reticular formation, to the afferents from the prefrontal (areas 10--12)
spinal cord. and prelimbic (area 32) cortices [595] and an
The PAG presumably also forms part of anterograde tracer study [451] has revealed
the control mechanisms for feeding and me- that in the rat the dorsal tegmental nucleus
tabolism. Luiten and collaborators [825] re- projects to widespread mesencephalic, dien-
cently adduced experimental neuroanatomi- cephalic and telencephalic centres, among
cal and neurophysiological evidence suggest- which are the superior colliculus, the prema-
ing that a descending sympathetic pathway, millary nuclei, the lateral geniculate body, the
including relay centres in the ventromedial midline, medial, intralaminar and anterior
hypothalamic nucleus, the PAG, the lower thalamic nuclei, the nuclei of the diagonal
medullary reticular formation and the spinal band, the medial and lateral septal nuclei,
intermediolateral column, influences the se- the medial amygdaloid nucleus, the hippo-
cretion of the pancreas and the adrenal me- campal region and the medial frontal cortex.
dulla. The functions of the dorsal tegmental re-
In the preceding paragraphs, the functions gion remain to be explored; however, it may
or possible functional roles of a number of be stated that the recent experimental hodo-
projections descending to and from the PAG logical studies cited have fully confirmed that
has been discussed. The dense projections this region represents a core structure of the
from the P AG to the intralaminar nuclei limbic system.
have been postulated to carry or modify noci- The connections described in this section
ceptive inputs to these parts of the thalamus, are illustrated in a schematic form in Figs.
but on the whole it should be stated that the 197, 198. The main long ascending and de-
role of the projections ascending from the scending pathways of the telencephalic and
PAG is largely unexplored and poorly under- diencephalic portions of the central limbic
stood [317]. continuum are represented in Figs. 197 A and
The most caudal portion of Nauta's limbic 197B, respectively, whereas in Fig. 197C a
midbrain area [977, 982] is the dorsal tegmen- number of shorter connections of the same
tal region, which includes the dorsal and ven- portions are depicted. Figs. 198A, B illus-
tral tegmental nuclei of Gudden. These two trate the principal connections of the centres
cells masses are large and distinct in rodents included in the limbic midbrain area. These
and carnivores, but only poorly developed two figure parts also show the most impor-
in primates. In man, only a relatively diffuse tant connections of the bed nucleus of the
dorsal nucleus can be distinguished [482, stria terminalis, a cell mass which will be dis-
1080]; hence, we will confine ourselves here cussed at the end of the next section.
to a brief overview of the most important
connections of the dorsal tegmental region.
The dorsal tegmental region is reciprocally
connected with the interpeduncular nucleus, The Amygdala
the mamillary body and the habenular nuclei
[447, 451, 483, 595, 802, 1081, 1082]; these Introduction
projections constitute the major afferent and
efferent systems of this region. Reciprocal It has already been mentioned that two large
connections also exist with the posterior hy- telencephalic components of the limbic sys-
pothalamic area [107, 595, 802], the dorsal tem, namely the amygdala and the hippo-
and superior central raphe nuclei [252, 595], campal formation are reciprocally connected
the nucleus praepositus hypoglossi [451, 595] with the rostral part of the central limbic con-
and the rhombencephalic medial reticular tinuum (Fig. 190). These two structures will
formation [802,1081,1082]. Interestingly, the now be discussed. The amygdala, or corpus
dorsal tegmental region also receives direct amygdaloideum, is a large nuclear complex
situated in the dorsomedial portion of the medial aspect of the amygdala, from where
temporal lobe, where it forms part of the ros- it runs a remarkably long curved course
tromedial and rostrodorsal walls of the infe- along the medial border of the caudate nucle-
rior horn of the lateral ventricle (Figs. 195, us to the anterior commissure. Immediately
196). Ontogenetically, the amygdala is a de- dorsocaudal to that commissure it splits up
rivative of the caudal part of the ganglionic into three components:
eminence, an intraventricular protrusion, the
rostral part of which gives rise to the corpus 1. A precommissural or supracommissural
striatum. Although the anlage of the amyg- component whose fibres descend in front
dala is displaced rostroventrally during de- of the anterior commissure.
velopment, it retains direct contact with the 2. A commissural component t~at enters the
striatum. The corpus amygdaloideum is divi- anterior commissure.
sible into a series of nuclei which may be 3. A postcommissural component that de-
separated into two major divisions, a cortico- scends caudal to the anterior commissure.
medial nuclear group and a basolateral nu-
The stria terminalis is composed of both
clear group ([265, 266]; Figs. 90-93). The
amygdalofugal and amygdalopetal fibres.
corticomedial group includes, apart from a
The ventral amygdalofugal pathway is a
number of smaller cell masses, the cortical
large assemblage of rather loosely arranged
amygdaloid nucleus and the medial amygda-
fibres which extends from the amygdaloid
loid nucleus. Together the nuclei of this
complex to the rostral part of the diencepha-
group constitute the dorsomedial part of the
lon. It arises from the dorsomedial part of
complex (Fig. 196). The most prominent sub-
the amygdala, then passes medially and
divisions of the basolateral nuclear group are
somewhat rostrally through the sub lenticular
the lateral amygdaloid nucleus, the basal
regions of the substantia innominata and the
amygdaloid nucleus and the accessory basal
substantia perforata anterior. Some of its
amygdaloid nucleus. A rather small central
fibres pass rostrally to the mediofrontal cor-
nucleus is sometimes included as part of the
tex, others spread in the lateral preoptico-
corticomedial nuclear group, but will be re-
hypothalamic zone, and still others enter the
garded here as a separate entity. This nucleus
inferior thalamic peduncle to terminate in the
is close to the bed nucleus of the stria termi-
medial thalamic nucleus [986]. Contrary to
nalis (which will be considered below) and
what its name suggests, the ventral amygda-
the morphologically most caudal parts of the
lofugal pathway conducts impulses in both
caudate nucleus and the putamen. Like the
directions.
latter two cell masses, the central amygdaloid
nucleus has a high content of dopamine
[1402].
Afferents to the Amygdala (Figs. 199,200)
The afferents of the amygdaloid complex can

Fibre Systems Related to the Amygdala
be grouped into the following five categories:
(Figs. 191-196)
1. Fibres originating from the olfactory bulb
Three large fibre bundles, the lateral olfacto- and from the olfactory part of the cerebral
ry stria, the stria terminalis and the ventral cortex.
amygdalofugal pathway, connect the amyg- 2. Fibres arising from the basomedial telen-
dala with other parts of the brain. The lateral cephalon and from the hypothalamus.
olfactory stria carries secondary olfactory 3. Thalamic afferents.
fibres to the cortical and medial amygdaloid 4. Direct afferents from the brain stem.
nuclei. Its course will be considered below. 5. Projections from various (non-olfactory)
The stria terminalis emerges from the caudo- areas of the cerebral cortex.
These fibre categories, which have been the axons emanating from the hypothalamic
represented diagrammatically in Figs. 199 centres mentioned reach the amygdala via
and 200, will now be briefly discussed. both the stria terminalis and the ventral path-
way.
1. Secondary olfactory fibres originating 3. The amygdala receives input from dor-
from the olfactory bulb pass by way of the sal as well as ventral thalamic regions. The
lateral olfactory tract to the amygdala, where dorsal thalamic centres that have been found
they terminate mainly in the cortical nucleus to distribute fibres to the amygdala include
[1150, 1394]. Apart from this direct olfactory the midline nuclear complex, the parafascicu-
projection, several routes along which the lar nucleus and the medial geniculate body,
amygdala may be indirectly influenced by ol- all of which project predominantly to the
factory impulses have been described. Thus central amygdaloid nucleus [6, 889, 1009,
it has been established that the prepiriform 1036, 1167]. In the ventral thalamus the peri-
cortex distributes fibres to the basolateral peduncular nucleus has been found to project
complex and that the entorhinal cortex pro- to the medial, cortical, central and lateral
jects to the central, basolateral and cortical amygdaloid nuclei [6, 1009, 1034, 1167].
nuclei [1035, 1102, 1166, 1406, 1436]. More- 4. Brain stem neurons located in the peri-
over it has been suggested that the deep aqueductal grey matter (PAG), the substan-
amygdaloid nuclei receive indirect olfactory tia nigra pars compacta, the ventral tegmen-
input via intrinsic connections from the corti- tal area, the dorsal raphe nucleus, the locus
cal part of the amygdala [1035]. coeruleus, the lateral parabrachial nucleus
2. The bed nucleus of the stria terminalis and the nucleus of the solitary tract project
and the nucleus of the horizontal limb of the to the central amygdaloid nucleus [6, 889,
diagonal band project predominantly to the 1005, 1009, 1034, 1167, 1437]. Most of the
medial and central nuclei, whereas the sub- brain stem centres mentioned project to the
stantia innominata distributes fibres to al- amygdala via the ventral amygdalofugal
most the entire expanse of the amygdala [6, pathway. However, the serotoninergic fibres
262, 978, 1167]. Several hypothalamic struc- from the dorsal raphe nucleus and the nor-
tures, including the paraventricular, the ven- adrenergic fibres from the locus coeruleus
tromedial and the infundibular nuclei, as well reach the amygdala via both the stria termina-
as the lateral hypothalamic area (LHA) pro- lis and the ventral pathway. It is noteworthy
ject to the medial part of the amygdala and that the lateral parabrachial nucleus is by far
to the central nucleus [254, 722, 1009, 1117, the most important subdiencephalic source
1167, 1193, 1320, 1437]. In the monkey this of amygdalar afferent connections [889,
hypothalamoamygdalar connection origi- 1034].
nates mainly from the ventromedial nucleus 5. The amygdaloid complex receives a
and from the caudal portion of the LHA [27, powerful input from the cerebral cortex. The
889]. Experimental neuroanatomical studies allocortical and mesocortical regions which
in rat [253] and cat [1167] have revealed that have been shown to distribute fibres to the
fibres originating from the medial preoptic amygdala include the subiculum [1152, 1166],
area pass to the amygdaloid complex, but the anterior cingulate gyrus (area 24) [6,
such fibres have not been observed in the 1035, 1045] and the perirhinal cortex (areas
monkey [889, 1196]. The cells in the bed nu- 35 and 36) [6, 514, 1035, 1166]. Fibres from
cleus of the stria terminalis project to the the anterior cingulate gyrus are distributed
amygdala via the bundle in which they are to the laterobasal nucleus, and fibres from
embedded. The fibres originating from the the perirhinal and subicular cortices have
nucleus of the diagonal band and substantia been traced to the mediobasal nucleus [1427].
innominata pass to the amygdala along the The subiculum also projects to the cortical
ventral "amygdalofugal" pathway, whereas nucleus [1166]. Neocortical afferents to the
1 Stria terminalis
2 ucleus interstitialis striae terminalis
3 uclei mediani thalami
4 Nucleus parafascicularis
8 Nucleus ventromedialis hypothalami
10 Fibrae " amygdalofugales " ventrales
11 ucleus gyri diagonalis, pars ventralis
15 Nuclei corticalis + medialis amygdalae
17 Nuclei basalis + lateralis amygdalae
20 Area tegrnentalis ventralis
23 Locus coeruleus
Fig. 199. Olfactory and subcortical afferent connections of the amygdaloid body
Fig. 200. Cortical afferents to the amygdaloid body. Numbers indicate fields of Brodmann (see Fig. 5)
amygdala ongmate from: the medial pre- These connections will now be discussed:
frontal cortex (areas 11 and 12; [1427]), the
caudal orbitofrontal cortex (areas 13 and 14; la. A large number of fibres originating
[6, 700, 776, 1406]), the insula [6, 661, 961, mainly but not exclusively from the cortico-
1035, 1436], and temporal cortical regions, medial nuclear group passes with the stria
including the temporal polar cortex (area 38), terminalis to the area (directly dorsocaudal
the auditory cortex in the superior temporal to the anterior commissure) in which the
lobe (area 22) and the visual association cor- fibres, as has already been mentioned, split
tex in the inferior temporal lobe (areas 20 up into three components, precommissural,
and 21; [514, 1395, 1502]). All of these neo- commissural and postcommissural. The pre-
cortical areas project principally to the lateral commissural fibres curve caudally around the
and basal nuclei. It has been pointed out by anterior commissure and terminate in the me-
Herzog and van Hoesen [514, 1427] that the dial preoptic and anterior hypothalamic ar-
cortico-amygdaloid projections are topo- eas as well as in the paraventricular and ven-
graphically organised in that allocortical ar- tromedial hypothalamic nuclei [107, 294, 495,
eas project mainly to the medial and cortical 721,873]. The commissural fibres connect the
nuclei, fibres from mesocortical areas project two cortical amygdaloid nuclei [752]. The
mainly to the basal nuclei, and the neocorti- postcommissural fibres spread to the bed nu-
cal areas project only to the lateral portion cleus of the stria terminalis, an elongated cell
of the complex. The central amygdaloid nu- mass that accompanies the stria terminalis
cleus is unusual in that it receives input de- throughout most of its extent, and to the re-
rived from all types of cortex. Thus, it region of the anterior hypothalamic nucleus.
ceives input from: the allocortical prepiri- These postcommissural fibres ongmate
form, periamygdaloid and subicular areas; mainly from the medial, basal and lateral
the mesocortical anterior cingular (area 24), amygdaloid nuclei [781].
infralimbic (area 25) and perirhinal (areas 35 1b. Fibres originating from the basolateral
and 36) regions and the neocortical prelimbic nuclear group and from the cortical areas
(area 32) orbitofrontal (areas 12 and 13), in- surrounding the amygdala have been re-
sular, and temporal (areas 20-22 and 38) reported to pass via the ventral amygdalofugal
gions [661, 978, 1035, 1166, 1427, 1502]. It pathway to a variety of structures, including
has also been reported that those compo- the mediofrontal cortex, the basal septal re-
nents of the amygdaloid complex which region and the lateral preopticohypothalamic
ceive neocortical afferents, such as the lateral zone[721,978,986].
nuclei and the basal complex, project, in turn, 2. A certain proportion of the ventral
upon the cortical nucleus [1103]. amygdalofugal fibres join the inferior tha-
lamic peduncle and pass to the medial nucle-
us of the thalamus [716, 719, 978]. The fibres
Efferents from the Amygdala of this projection originate principally from
(Figs. 201, 202) the basal nuclear group and terminate in the
medial, magnocellular portion of the medial
The efferent projections of the amygdaloid
nucleus [7].
complex are distributed to:
3. The central amygdaloid nucleus gives
1. The septopreopticohypothalamic contin- rise to a large projection, whose constituent
uum. fibres initially travel medially with the ventral
2. The dorsal thalamus. amygdalofugal pathway and then turn cau-
3. A variety of cell groups located m the dally, to descend through the most lateral
brain stem. part of the hypothalamus and the tegmental
4. The corpus striatum. regions of the brain stem [549, 1233, 1412].
5. A number of cortical areas. The most caudal fibres of this so-called
amygdalotegmental projection attain the cer- certain portions of the basal complex project
vical cord [924, 1184]. Numerous fibres from strongly to the entorhinal cortex [720], which
the initial part of the projection are sent to represents the principal source of cortical
the substantia innominata, the nucleus of the input to the hippocampal formation (Figs.
horizontal limb of the diagonal band, the bed 202-204). Additionally, the basal amygdaloid
nucleus of the stria terminalis and the caudal complex has substantial reciprocal intercon-
part of the lateral hypothalamic area (LHA). nections with the subiculum, the major
In the brain stem, fibres are distributed to source of hippocampal output [720, 1427].
numerous centres, including the peripedun- Certain portions of the laterobasal nucleus
cular nucleus, the substantia nigra pars com- project to the prelimbic (area 32), infralimbic
pacta, the ventral tegmental area, the cunei- (area 25), anterior cingulate (area 24) and
form nucleus, the periaqueductal grey mat- perirhinal (areas 35 and 36) cortices as well
ter, the raphe nuclei, the parabrachial nuclei, as to the insula [25, 719, 836, 1097]. The en-
the locus coeruleus and the area subcoerulea, tire frontal cortex, including the motor (area
the rhombencephalic reticular formation, the 4) and premotor (area 6) regions, receives a
dorsal motor nucleus of the vagus and the direct projection from the basal amygdaloid
nucleus of the solitary tract [549, 550, 927, nuclei [66, 606, 803, 804], Particularly heavy
1103, 1223, 1360, 1412]. The medial and lat- projections have been traced to the lateral
eral parabrachial nuclei and the dorsal vagal and medial parts of the orbitofrontal cortex
complex are richly supplied. (areas 12, 13a and 14) [25].
4. Fibres originating mainly from the bas- Amaral and Price [25] recently reported
al amygdaloid nucleus pass via the longi- that all major divisions of the temporal neo-
tudinal association bundle and the stria ter- cortex receive a projection from the amygda-
minalis to the ventral as well as the dorsal loid complex, with the most prominent pro-
striatum. The former includes the accumbens jections ending in the cortex of the temporal
nucleus and certain striatal-like portions of pole and the rostral superior temporal gyrus.
the olfactory tubercle; the latter is composed These authors also found a strong projection
of the nucleus caudatus and the putamen to peristriate regions of the occipital lobe.
[494,496,497]. The fibres entering the dorsal
striatum are topographically organised and
Comments on the Connections
terminate predominantly in the ventral por-
and Functions of the Amygdala
tions of the putamen and ventral and caudal
parts of the caudate nucleus [675, 1170, If we survey the fibre connections of the
1171]. This amygdalostriate projection over- amygdala, it appears that this complex enter-
laps the striatal projections from the tains reciprocal connections with numerous
cingulate cortex, the ventral tegmental area and widespread areas in the brain, extending
and the mesencephalic raphe nuclei [675]. from the cerebral cortex to the nucleus of
Like the amygdalostriatal system all of these the solitary tract. However, a number of con-
striatal afferents avoid the anterodorsolateral nections are strictly unidirectional. Thus, the
striatal sector, which has been shown to be amygdalostriate projection and the amygda-
the main region to which the corticostriatal lothalamic projection terminating in the me-
projection from the sensorimotor cortex is dial thalamic nucleus are not reciprocated by
distributed. As mentioned before, these ob- strio-amygdaloid or thalamo-amygdaloid
servations suggest a division of the striatum projections. The thalamus does send fibres
into "limbic" and "non-limbic" compart- to the amygdala, but these originate from the
ments [675]. midline, parafascicular and medial geniculate
5. Projections arising in specific amygda- nuclei, rather than from the medial nucleus.
loid nuclei terminate in distinct areas of the The central amygdaloid nucleus projects
cerebral cortex. Thus, the lateral nucleus and directly to a considerable number of hypo-
1 Stria terminalis
2 Nucleus medialis thalami, pars medialis
3 Nucleus interstitialis striae terminalis
4 Nucleus caudatus + putamen
5 Commissura anterior 22 Substantia nigra, pars compacta
6 Nucleus paraventricularis 23 Area tegmentalis ventralis
7 Nucleus anterior hypothalami 24 Nucleus peripeduncularis
8 Area lateralis hypothalami 25 Griseum centrale mesencephali
9 Nucleus ventromedialis 26 Nucleus raphes dorsalis
10 Nucleus preopticus 27 Nucleus cuneiformis
11 Nuclei septi + nucleus gyri diagonalis, pars dorsalis 28 Nucleus centralis superior
12 Nucleus accumbens 29 Nucleus subcoeruleus
13 1\Jberculum olfactorium 30 Locus coeruleus
14 Nucleus gyri diagonalis, pars ventralis 31 Nuclei parabrachiales
15 Substantia innominata 32 Formatio reticularis rhombencephali
16 Nucleus centralis amygdalae 33 Nucleus raphes magnus
17 Nuclei corticalis + medialis amygdalae 34 Nucleus raphes pallidus
18 Nuclei basalis + lateralis amygdalae 3S Nucleus raphes obscurus
19 Cortex periamygdaloideus 36 Nucleus dorsalis nervi vagi
20 Subiculum 37 Nucleus solitarius
21 Cortex entorhinalis 38 Fibrae amygdalospinales
Fig. 201. Subcortical and hippocampal efferents of the corpus amygdaloideum

Fig. 202. Efferents of the corpus amygdaloideum to the cerebral cortex. Numbers indicate fields of
Brodmann (see Fig. 5)
thalamic and brain stem centres, and these gested that this projection is implicated in
projections are considered to contribute sig- facial expression and in other forms of motor
nificantly to the organisation of the visceral, behaviour associated with the emotional and/
autonomic, somatosensory and somatomo- or experiential state of the individual [1170].
tor components of affective behaviour, such The amygdala is strongly and reciprocally
as defense or flight. Electrical stimulation of connected with the principal input and out-
the central nucleus has been shown to pro- put centres of the hippocampus, i.e. the ento-
duce somatic, cardiovascular and respiratory rhinal cortex and the subiculum, respectively.
components of the defense response [650, The amygdala and the hippocampus and
1223], and it has been suggested by Kuypers their respective connections with mesocorti-
[745] that certain components of the amygda- cal and neocortical regions are considered to
lotegmental projection become especially ac- form part of an integrated system involved
tive during times of emergency or extreme in the processing of long term memory [918,
stress, and may be instrumental in providing 919]. Clinical and experimental evidence indi-
motivational drive for the execution of move- cates that the medial, magnocellular portion
ments. The same author also postulated that of the medial thalamic nucleus (MDmc) also
other components of the amygdalotegmental has an important mnemonic role [7]. As men-
projection may activate the so-called de- tioned this cell mass receives a substantial
scending pain control system, thus suppress- projection from the amygdala, which arises
ing pain transmission during the execution mainly from the basal nuclear group. This,
of motor actions of vital priority. The various of course, opens the possibility that the prin-
types of affective behaviour are strongly in- cipal projection areas of MDmc, i.e. the
fluenced by extrinsic influences, and physio- medial and orbital prefrontal cortices, may
logical experiments have demonstrated that likewise playa role in memory [7]. Recently,
the amygdaloid complex receives informa- evidence has been presented that the large
tion from all sensory modalities [101, 1185]. cholinergic cells in the substantia innomina-
The pathways through which the amygdala ta, which receive considerable input from the
receives olfactory, gustatory, visual and audi- basal nuclear complex of the amygdala and
tory impulses are well known, and these im- project in a topographically organised fash-
pulses may reach the central nucleus either ion to the cerebral cortex, are also involved
directly, or indirectly, i.e. via intrinsic amyg- in learning and memory [1168].
daloid connections arising from the basal and
lateral nuclei. The gustatory impulses origi-
nate from the insular cortex [961], and it is
Bed Nucleus of the Stria Terminalis
important to note that the visual and audito-
(Figs. 91 and 198)
ry amygdalopetal projections arise exclusive-
ly from association areas and, hence, may
be expected to convey highly processed sen- Before turning to the hippocampus, brief
sory information [1395]. In summary, it is consideration will be given to the interstitial
reasonable to assume that polysynaptic corti- nucleus of the stria terminalis. This cell mass,
co-amygdalotegmental pathways provide the which is also known as the bed nucleus of
anatomical substrate for the initiation and the stria terminalis (BNST) starts from the
integration of somatic and autonomic re- dorsal aspect of the amygdala and extends
sponses associated with affective behaviour. dorsomedially along and among the fibre
Another access route along which the amyg- bundles of the stria terminalis. Its intermedi-
daloid complex participates in the organisa- ate part envelops the anterior commissure
tion of affective and other behaviour may and its dorsal part curves alongside the ros-
well be provided by the large amygdalostriate tral one-third of the thalamus. The interme-
projection [496]. It has recently been sug- diate part of the BNST is rostromedially, me-
dially and caudomedially in direct contact from the BNST to the lateral and basal
with the septal, preoptic and anterior hypo- amygdaloid nuclei have also been described
thalamic areas, respectively [155, 986]. Swan- [253, 254, 1011].
son and Cowan [1331] include the BNST 2. Fibres originating from the BNST have
within the septal complex, but several other been traced to the accumbens nucleus, the
authors [109, 294, 542, 622] have expressed substantia innominata and, via the medial
the opinion that on both positional and con- forebrain bundle, to the preoptic region and
nectional grounds this nucleus and the cen- to most parts of the hypothalamus, including
tral and medial amygdaloid nuclei should be the ventromedial nucleus, the lateral hypo-
considered together as a single anatomical thalamic area, and the premamillary and su-
entity. It is noteworthy that the BNST may pramamillary regions [1331].
show marked sexual dimorphism. It has been 3. The BNST projects to the paraventricu-
reported that in guinea-pigs this nucleus is lar thalamic nucleus (one of the midline nu-
twice as large in males as in females [524]. clei in rodents) and, via the stria medullaris,
The BNST receives afferents from the ce- to the medial portion of the lateral habenular
rebral cortex, the amygdala, the hypothala- nucleus [253, 254, 1320, 1331].
mus and several centres in the brain stem. 4. Numerous efferent fibres of the BNST
The cortical afferents originate from the insu- descend by way of the medial forebrain bun-
la [661] and from the subiculum [1330]. The dle and the mesencephalic central tegmental
BNST is one of the major terminal fields of field to the lateral tegmental field of the pons
the amygdala. Most of these amygdaloid af- and the medulla oblongata [542]. These fibres
ferents stem from the medial nucleus, but the have been observed to project substantially
central nucleus and the basal complex also to the ventral tegmental area, the substantia
contribute to this projection [721, 1492]. Hy- nigra pars compacta, the periaqueductal grey
pothalamic afferents of the BNST include matter and the adjoining cuneiform nucleus,
fibres originating from the paraventricular the dorsal raphe nucleus, the pontine central
[254] and ventromedial nuclei [722, 1193]. In grey matter, the medial and lateral parabra-
the brain stem the periaqueductal grey mat- chial nuclei, the locus coeruleus, the nucleus
ter, the ventral tegmental area, the parabra- subcoeruleus, the nucleus raphes magnus and
chial nuclei and the dorsal vagal complex the adjacent medial reticular formation, the
send fibres to the BNST [1191, 1268]. Ricar- lateral reticular formation, the nucleus of the
doh and Koh [1123, 1124] have emphasized solitary tract and the dorsal vagal nucleus
that the BNST is the recipient of the most [254, 542, 1223, 1268, 1331, 1412]. The two
dense of the pathways that ascend from the latter centres are particularly densely inner-
interoceptive portion of the nucleus of the vated. As Holstege and collaborators [542]
solitary tract to the forebrain. remarked, the brain stem projections of the
The BNST distributes its efferent fibres to BNST are virtually identical to the ones
(1) the amygdala, (2) the mediobasal telence- derived from the central nucleus of the amyg-
phalon and the hypothalamus, (3) the thala- dala.
mus and epithalamus and (4) a considerable
number of grisea located in the brain stem. Physiological experiments indicate that
These connections will now be briefly dis- the BNST, with the amygdala, the septal and
cussed. the preoptic regions, is involved in the regula-
tion of somatic, cardiovascular and respira-
1. A large number of fibres originating tory components of affective behaviour, par-
from the BNST passes, largely with the ven- ticularly flight and defense [825, 1223, 1233].
tral "amygdalofugal" pathway, to the amyg- The descending efferents of the BNST pass-
dala, where they terminate mainly in the me- ing to reticular and autonomic centres in the
dial and central nuclei [1331]. Fibres leading brain stem provide an anatomical substrate
for these regulatory influences. The BNST tion of the retrocommissural hippocampus is
has also been found to play an important recurved dorsally and constitutes a rounded
role in male sexual behaviour [330]. swelling on the medial surface of the hemi-
sphere, known as the uncus. On the ventricu-
lar side this part of the hippocampus is great-
ly enlarged and differentiated into a series
The Hippocampus of separate lobules, the hippocampal digita-
(Figs. 11,23; 93-95; 190, 193, 195) tions. As shown by Figs. 81, 82 and 195, the
most rostral portion of the hippocampal for-
Introduction mation extends for some distance ventral to
the amygdaloid complex.
The hippocampus or hippocampal formation The hippocampal formation constitutes
is a large "C" -shaped structure that forms the archipallial part of the cerebral hemi-
part of the medial wall of the cerebral hemisphere; it contains a relatively simple three-
sphere. This structure can be subdivided layered allocortex throughout its extent. The
morphologically into three parts, precom- retrocommissural hippocampus is clearly dif-
missural, supracommissural and the retro- ferentiated into three longitudinally arranged
commissural. The first two parts are relative- structures: the fascia dentata, the cornu am-
ly small, vestigial structures; the retrocom- monis (Ammons's horn) and the subiculum.
missural hippocampus, on the other hand, The fascia dentata - whose name refers to
is well developed and represents the main the toothed or beaded appearance of its sur-
portion of the hippocampal formation. The face - is the morphologically most medial
names of the three parts, it should be noted, strip of the pallium. It is laterally continuous
refer to their position with respect to the cor- with the cornu ammonis, which in its turn
pus callosum. The precommissural hippo- passes over into the subiculum. Due to the
campus is a narrow vertical structure, situ- infolding of the hippocampus, the fascia den-
ated in the caudal part of the area subcallosa tata is above, and the subiculum below the
just rostral to the septum verum. It continues hippocampal sulcus. The fascia dentata con-
dorsorostrally into the indusium griseum, a tains a granule cell layer of small neurons,
strand of hippocampal tissue which extends whereas large pyramidal elements prevail in
throughout the length of the corpus callo- both the cornu ammonis and the subiculum.
sum. The indusium griseum represents the su- On the medial surface of the hemisphere the
pracommissural hippocampus. Two small subicular cortex is contiguous with the juxtal-
fibre bundles, the medial and laterallongitu- locortex or mesocortex. The latter represents
dinal striae, are embedded in it. These bun- a type of cortex that is transitional between
dles represent a small supracallosal compo- the hippocampal allocortex and the neocor-
nent of the fornix. The principal subcallosal tex. This transitional cortex covers the para-
fornix will be discussed below. Near the hippocampal gyrus and is also found in the
splenium of the corpus callosum, the supra- supracallosal cingulate gyrus.
commissural hippocampus is continuous The cingulate gyrus, which borders on the
with the retrocommissural hippocampus, the vestigial precommissural and supracallosal
expanded morphologically most caudal end parts of the hippocampus, is in the area be-
of the hippocampal formation. During onto- hind the splenium of the corpus callosum di-
genesis, this structure, which is in the medial rectly continuous with the parahippocampal
temporal lobe, rolls in on itself along a longi- gyrus. The hippocampus, the parahippocam-
tudinal groove, the hippocampal sulcus. This pal gyrus and the cingulate gyrus together
infolding causes the retrocommissural hippo- constitute a large arcuate convolution known
campus to protrude into the inferior horn as the limbic lobe (Fig. 4). The allocortical
of the lateral ventricle. The most rostral por- hippocampus forms the" inner ring", where-
as the mesocortical parahippocampal and of fibres decussate to the opposite side, thus
cingulate gyri form the" outer ring" of that constituting the commissure of the fornix.
lobe. Most parts of the hippocampal and Proceeding rostrally over the thalamus, the
parahippocampal cortices in the temporal two crura converge and join to form the cor-
lobe can be further subdivided on both cy- pus of the fornix, which lies immediately be-
toarchitectonical and connectional grounds. neath the corpus callosum. However, at the
Without going into details it may be men- level of the anterior pole of the thalamus the
tioned that within the cornu ammonis there fornical corpus separates again into two bun-
are four different distinguishable fields which dles, the columns of the fornix, which curve
are usually designated as CA1-CA4. CAl ventrally in front of the interventricular fora-
borders on the subiculum, whereas CA4 is men and caudal to the anterior commissure
an aggregation of large cells that, fills the to enter the hypothalamus. Immediately be-
hilus of the curved fascia dentata. Field CAl hind the interventricular foramen a consider-
is remarkably well developed in primates and able number of fibres leave the column and
man [1292]. The subiculum proper is sepa- pass backwards to the anterior nucleus of the
rated by two strips of mesocortex, known as thalamus and to the bed nucleus of the stria
the presubiculum and the parasubiculum, terminalis. Other fibres split off from the for-
from the entorhinal cortex. The latter largely nix just above the anterior commissure and
corresponds to area 28 of Brodmann (Fig. 5) constitute a small precommissural portion of
and occupies a large part of the parahippo- the fornix. The main bundle of the fornix
campal gyrus. It is ventrally flanked by the or postcommissural fornix finally traverses
so-called perirhinal cortex, which encom- the hypothalamus, where most of its fibres
passes areas 35 and 36 of Brodmann. terminate in the mamillary body.
Fibre Systems Related to the Intrinsic Connections of the Hippocampus

Hippocampus (Figs. 203-206) (Fig. 204)
Before focussing on the afferent and efferent A discussion of the complex intrinsic connec-
connections of the hippocampus, two large tions of the hippocampus is beyond the scope
limbic fibre bundles, the cingulum and the of the present work. It is, however, important
fornix, should be briefly commented upon. to note that the main flow of information
The cingulum is a bundle of short and long through this structure is thought to take
association fibres which surrounds the cor- place through a trisynaptic circuit, which be-
pus callosum. Passing through the core of gins with a projection from the entorhinal
the cingulate and parahippocampal gyri, it area to the fascia dentata, is relayed to field
extends from the septal area to the uncus re- CA3 of the cornu ammonis and then pro-
gion in the temporal lobe. The fornix is a ceeds to field CAl. A projection from the
compact fibre bundle connecting the hippo- latter, which terminates in the entorhinal
campus with the hypothalamus and various area, closes this trisynaptic circuit [1323,
other structures. Its fibres first form the al- 1324].
veus, a thin white layer on the ventricular The fibres destined for the fascia dentata
surface of the cornu ammonis, and then con- arise from all parts of the entorhinal area
verge as the fimbria along the medial aspect and constitute the so-called temporo-am-
of the hippocampus. Running posterosuper- monic or perforant path. The fibres of this
iorly, the fibres of the fimbria enter the crus large projection do not terminate exclusively
of the fornix, a flattened structure that arches in the fascia dentata, but also supply the
upwards and medially under the splenium of other parts of the hippocampal formation,
the corpus callosum. In this region a number i.e. the subicular cortices and the various
fields of the cornu ammonis [527, 1330, [1426, 1429]), infralimbic (area 25; [1148,
1550]. The axons of the dentate granule cells, 1426]), insular [1144], cingulate (areas 23 and
which constitute the hippocampal mossy 24; [1046, 1426]) and temporal polar cortices
fibre system, project in a topographically or- (area 38; [1426,1428]). Several of these areas,
dered manner upon the pyramid cells in CA3 particularly the posterior orbitofrontal, ante-
[1342]. Major intrahippocampal outputs of rior cingulate and temporal polar cortex, are
field CA3 are carried by the so-called sites of convergence of multimodal sensory
Schaffer collateral system to field CAl. The information. Pathways leading out of the pri-
direct projection from CAl to the entorhinal mary visual, auditory and somesthetic areas
area (which, as mentioned above, closes the of the neocortex, by way of a variable
trisynaptic principal hippocampal circuit number of intercalated association areas in
[526, 1152, 1153] is supplemented by an indi- the parietal, frontal and temporal lobes, ulti-
rect projection. The latter is synaptically in- mately converge on these areas [640, 1144,
terrupted in the subicular complex, i.e. the 1428, 1429]. Association areas have also been
area where most hippocampal efferents, both found to project directly to the parahippo-
cortical and subcortical, originate [88, 348, campal cortices. Thus, visual [105, 114, 122a,
892, 1152, 1241, 1242, 1273, 1328, 1330, 122b] and auditory [123b] association areas
1430]. Most of the intrinsic connections just project to the perirhinal and entorhinal areas
discussed are transversely arranged with re- [23, 1144, 1426], and fibres originating from
spect to the long axis of the hippocampus. multimodal association areas situated in the
prefrontal cortex (areas 9 and 46; [418, 979]),
inferior parietal lobule (areas 7 and 39;
Afferents to the Hippocampus [1232]), and inferior temporal lobe terminate
(Figs. 203, 204) in the presubiculum.
From the foregoing it may be concluded
Extrinsic inputs, which are presumed to acti- that the parahippocampal cortices and ulti-
vate and modulate the intrinsic hippocampal mately the hippocampus are provided with
circuitry [1324] arise from a variety of a broad spectrum of synthetized sensory-spe-
sources, including (1) various cortical areas, cific as well as multimodal cortical informa-
(2) the amygdaloid complex, (3) the medial tion [1426]. Because the entorhinal area is
septal- diagonal band complex, (4) the thala- also the recipient of olfactory information,
mus, (5) the supramamillary region, and (6) either arising from the olfactory bulb [490,
the mesencephalic raphe nuclei and locus 1105, 1208] or from the prepiriform cortex
coeruleus. (area 51; [464, 719, 720,1146]), it seems likely
that information derived from all sensory
1. Pathways leading out of widespread modalities is fed into the hippocampal for-
cortical areas converge upon the hippocam- mation.
pus. Most of the fibres of this large cortico- Recent experimental neuroanatomical
hippocampal projection terminate in the en- [1144,1145,1146,1425,1522] and physiolog-
torhinal area, i.e. the site of origin of the ical studies [813, 1423, 1424] have shown that
perforant pathway [1144, 1428, 1429], others along its spleniotemporal axis the hippocam-
are synaptically interrupted in the perirhinal pus cannot be considered a homogeneous
as well as in the entorhinal areas [1144, 1430, structure. There is topographical organisa-
1522], and still others bypass the entorhinal tion of the en to rhinal-hippocampal projec-
area to terminate directly in the subicular tion so that a lateromedial axis in the entorhi-
cortices [418, 979, 1232, 1430]. Substantial nal cortex corresponds to the spleniotempor-
contributions to the corticohippocampal pro- al axis of the hippocampus. This means that
jection have been found to emanate from the lateral parts of the entorhinal cortex distrib-
posterior orbitofrontal (areas 12 and 13; ute fibres to more splenial portions of the
hippocampus, whereas progressively more septal nucleus and the nucleus of the diago-
medial parts of the entorhinal cortex project nal band contain large numbers of choliner-
to increasingly more temporal hippocampal gic elements [904, 905, 907], and it may be
portions. The lateral and medial parts of the safely assumed that the numerous cholinergic
entorhinal areas differ considerably with re- fibres present in the hippocampal formation
spect to their afferents. The lateral part re- originate from these neurons [1301, 1466].
ceives predominantly cortical pathways con- The projection from the septum and diagonal
veying information from all sensory modali- band complex to the hippocampal and para-
ties [1144, 1522], whereas the medial parts hippocampal cortices is, however, only par-
are more influenced by subcortical centres tially cholinergic [483] and several other neu-
such as the septum, thalamic midline nuclei rotransmitters, such as gamma-amino butyr-
and amygdala [1145]. These differential in- ic acid (GABA; [698, 1050]) and substance
puts to the medial and lateral parts of the P [1448], may also be contained in this pro-
en to rhinal cortex and the topographical or- jection.
ganisation of the entorhinal-hippocampal The hippocampus shows a characteristic
perforant path discussed above lead to a con- slow rhythmical electrical activity, usually
vergence of exteroceptive sensory informa- designated as the hippocampal theta rhythm.
tion on the splenial portions of the hippo- There is evidence suggesting that both the
campus, whereas information derived from direct septohippocampal and the indirect
limbic-related subcortical centres, presum- septo-entorhinal-hippocampal projections
ably reflecting the intrinsic status of the are important in the generation of the hippo-
organism, is transmitted in particular to campal theta rhythm [813, 920].
the temporal portions of the hippocampal 4. Two parts of the dorsal thalamus, the
formation [1522]. anterior nuclei and the midline complex, pro-
2. The hippocampus and the entorhinal ject to the hippocampal and parahippocam-
cortex receive a substantial input from the pal areas. The fibres originating from the an-
amygdala [22, 88, 717, 719, 720, 1145]. The terior nuclei reach the temporal lobe via the
basolateral amygdaloid nuclei project strong- cingulate bundle and terminate in the presu-
ly to the en to rhinal cortex and also distribute biculum, parasubiculum, and subiculum
numerous fibres to the subiculum [717, 719, proper [17, 309, 1133, 1245]. This projection
720, 1145]. Projections from the cortical and forms part of the so-called circuit of Papez,
medial amygdaloid nuclei to the entorhinal which will be discussed below. The fibres tak-
cortex [88] and to the hippocampal formation ing origin from the midline nuclei (in rat, cat
[1427] have also been described. and monkey most of these fibres arise from
3. A major projection to the hippocampus the nucleus reuniens thalami [22, 88, 296,
and the entorhinal cortex arises from the me- 506,1145,1535]) pass to the genu of the cor-
dial septal nucleus and the nucleus of the di- pus callosum, where they join the cingulate
agonal band [20, 22, 24, 88, 296, 310, 679, bundle [506]. After having curved around the
893, 1059, 1145, 1530]. The fibres constitut- callosal splenium, these fibres innervate the
ing this projection follow three different entorhinal and subicular cortices and the hip-
routes. The majority of fibres pass through pocampal CAl field [506, 1145]. The nucleus
the fornix-fimbria system, whereas smaller reuniens thalami receives input from the
contingents enter the cingulate bundle, or parabrachial nuclei, which in turn receive af-
take a ventral route via the medial forebrain ferents arising from the nucleus of the soli-
bundle and then through the amygdaloid tary tract, a primary viscerosensory centre.
complex [24,1145]. The projection to the hip- Hence, the nucleus reuniens may well help
pocampus terminates principally in the fascia transmit information from the visceral pe-
dentata and the adjacent region of the cornu riphery to limbic structures [506].
ammonis [24, 227, 270, 1128]. The medial
Insula
c am Cornu ammonis
fd Fascia dentata
sub Subiculum
The numbers refer to cortical areas according to Brodmann
Fig. 203. Direct and indirect cortical afferents of the hippocampus

1 Cingulum
2 Fornix
4 Nuclei mediani thalami
6 Nucleus septi medialis
7 Nucleus gyri diagonalis, pars dorsalis
8 Regio supramamillaris
9 Fascia dentata
10 Cornu ammonis
11 Subiculum
12 Nuclei basalis+lateralis amygdalae
13 Tractus temporo-ammonis or "perforant pathway"
Fig. 204. Subcortical afferents and intrinsic connections of the hippocampus

5. The supramamillary region sends, via clusively in the lateral septal nucleus [894,
the fornix, a substantial projection to the hip- 1328, 1330]. These fibres, which are presum-
pocampal region [22, 465, 1059, 1128, 1530]. ably glutamatergic [1550], arise from all cell
Most of these fibres terminate in the fascia fields of the cornu ammonis [715]. Multiple
dentata and in the adjacent CA3 field, but retrograde labelling studies suggest that most
the remaining fields of the cornu Ammonis pyramidal cells in the cornu ammonis which
and the entorhinal cortex are also supplied project to the septum also give rise to intra-
[465, 1145]. hippocampal collaterals [1324]. It is of inter-
6. The hippocampus contains diffusely ar- est that the lateral septal nucleus projects
ranged noradrenergic and serotoninergic massively to the medial septal - diagonal
fibres, which reach this structure via the for- band complex [1108], which is known to pro-
nix as well as via longitudinal striae and the ject back to the hippocampal formation.
cingulate bundle. The noradrenergic fibres 2. The precommissural fornix fibres
arise from the locus coeruleus in the upper originating from the subicular complex are
rhombencephalon, whereas the serotoninerg- distributed to the lateral septal nucleus, the
ic fibres originate from the mesencephalic accumbens nucleus, the anterior olfactory
raphe nuclei, i.e. the dorsal raphe nucleus and nucleus, the precommissural hippocampus,
the central superior nucleus [997]. The raphe the medial part of the frontal cortex and the
projection to the hippocampal region con- gyrus rectus [894, 1152, 1330].
tains a substantial number of non-serotonin- The projection from the subicular complex
ergic fibres [699]. to the accumbens nucleus presumably consti-
tutes an important part of the interface of
the limbic system with central somatomotor
Efferents from the Hippocampus mechanisms [928, 931,1536]. It is known that
(Fig. 205) the nucleus accumbens projects massively to
the ventral pallidum, and that fibres originat-
Turning now to the efferent connections of ing from the latter structure terminate in and
the hippocampus, it should first be stated around the tegmental pedunculopontine nu-
that investigations using so-called tracer cleus, i.e. in an area which has been desig-
techniques have radically changed insight nated as the mesencephalic locomotor region
into the organisation of these connections. [240, 1336]. These connections between the
Contrary to what was believed for almost a nucleus accumbens, the ventral pallidum and
century, the entire postcommissural fornix the mesencephalic locomotor region are im-
and a considerable part of the precommissu- portant in mediating locomotor responses as-
ral fornix originate from the subiculum rath- sociated with exploratory behaviour [928,
er than from the cornu ammonis [894, 1328, 931]. The hippocampus probably affects the
1330]. The contribution of the latter structure programming and execution of these re-
to the fornix has been shown to be only mi- sponses by way of the fibres passing from
nor and to be confined to the precommissural the subicular complex to the nucleus accum-
fornix. In the light of these findings, the hip- bens. In this context it is interesting to note
pocampal efferents may be grouped as fol- that these fibres synapse directly on the neu-
lows: (1) efferents of the cornu ammonis, rons of the accumbens-ventral pallidum pro-
(2) contributions of the subiculum to the pre- jection [1536].
commissural fornix, (3) contributions of the 3. The postcommissural fornix contains,
subiculum to the postcommissural fornix and apart from some hippocampal afferents, only
(4) "non-fornical" efferents. fibres originating from the subicular com-
plex. Most of these fibres terminate in the
1. The precommissural fornix fibres origi- mamillary body; smaller contingents are dis-
nating from the cornu ammonis terminate ex- tributed to the anterior thalamic nucleus, the
bed nucleus of the stria terminalis and the area 41), sensory-specific assocIation areas
cell-free capsular zone surrounding the ven- (visual, areas 19-21; auditory, area 22; soma-
tromedial hypothalamic nucleus [715, 892, tosensory, area 7) as well as multimodal asso-
894, 895, 896, 1152, 1328, 1330]. As regards ciation areas (areas 9, 13, 23, 38 and 46).
the latter connection, it is worth mentioning It follows that the classical view according
that Ricardo [1123], considering that neurons to which the hippocampal formation is an
in the infundibular nucleus send dendrites intermediary station in a system of connec-
into the capsular zone of the ventromedial tions that travel largely from cortical areas
nucleus, suggested that the hippocampus via to subcortical areas has to be abandoned.
this particular connection, may influence the The anatomical evidence clearly indicates
secretion of anterior pituitary hormones. that the key to understanding the function
4. Fibres originating from the subicular of the hippocampus lies in recognising that
complex project massively to the adjacent en- its major connections are with associational
torhinal cortex [88, 593, 1152, 1273, 1426]. cortical areas on the one hand and with sub-
As has already been mentioned, this projec- cortical limbic structures on the other [1324].
tion is reciprocated by a strong entorhinal- Our understanding of the functional sig-
subicular connection. Both projections are nificance of the hippocampal formation is
topographically organised, in that a latero- still fragmentary. However, clinico-patholog-
medial gradient in the entorhinal cortex cor- ical evidence strongly suggests that the hip-
responds to a spleniotemporal gradient along pocampal-parahippocampal complex and its
the longitudinal axis of the subiculum [1425]. bidirectional relationships with the associa-
The entorhinal-hippocampal perforant path- tional cortical areas are prominent in learn-
way shows a similar topographical organisa- ing and memory processes [316, 582, 1426,
tion [1520, 1521]. Other, "non-fornical" 1454]. It is interesting to note in this context
efferents of the subicular complex include that bilateral destruction of the fornix does
projections to the posterior cingulate (area not lead to severe memory disorders [1152].
23) and retrosplenial (areas 29 and 30) cor-
tices, and to the amygdala [593, 896, 1152].
It is important to note that the hippocampus, The Circuit of Papez (Fig. 206)
via the entorhinal (area 28) and perirhinal
(areas 35 and 36) cortices, initiates a series The postcommissural fornix is prominent
of projections to the cerebral cortex that within a system of centres and connections
largely reciprocate the input pathways from that involves both the inner and the outer
the same source (Fig. 203). Thus, the entorhi- ring of the limbic lobe. This system, which
nal cortex is known to project to the pre- has already been briefly discussed, is known
piriform (area 51) prelimbic (area 32), infra- as the "circuit of Papez". It was suggested
limbic (area 25), agranular insular, cingulate by that author [1051] to be an essential part
(areas 23 and 24), retrosplenial (area 29), of the structural basis of emotion. The circuit
perirhinal (areas 35 and 36), posterior orbito- was originally considered to comprise the fol-
frontal (area 13) and temporal (areas 20, 22, lowing structures and connections: hippo-
38 and 41) cortices [418, 708, 1272, 1426, campus - postcommissural fornix - mamil-
1521], whereas the perirhinal cortex projects lary body - mamillothalamic tract - anterior
to the prefrontal (areas 9 and 46), temporal thalamic nucleus - thalamocingulate projec-
(areas 20-22), parietal (area 7) and occipital tion - cingulate gyrus - cingulum bundle -
(area 18) cortices [1426, 1521]. By means of hippocampus.
these diverging projections, hippocampal Experimental neuroanatomical and physi-
output is disseminated over numerous and ological studies have considerably increased
widespread cortical areas, including primary our knowledge of the circuit of Papez. An
sensory areas (olfactory, area 51; auditory, extensive survey of the pertinent literature is
2
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,
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\, \,
,, ''
,
I

2 Cortex retrosplenialis (areae 29, 30)
3 Fornix
6 Hippocampus precommissuralis
7 Cortex frontalis medialis
8 Nuclei septi
9 Gyrus rectus
11 Nucleus olfactorius anterior
12 Nucleus ventromedialis
13 Corpus mamillare
15 Cornu ammonis
16 Subiculum
17 Cortex entorhinalis (area 28)
Fig. 205. Efferents of the hippocampus

1 Area prelimbica (area 32)

3 Cingulum
5 Fornix
7 Tractus mamilJothalamicu
8 Tractus mamilJotegmentalis
9 Corpus mamillare
10 Subiculum
Fig. 206. The circuit of Papez

beyond the scope of the present work. A few The Olfactory System
aspects may be briefly mentioned, however: (Figs. 207, 208)
1. It has been found that hippocampal af-
ferents travelling with the cingulate bundle
terminate in the presubiculum [308, 1241, The central olfactory system or rhinencepha-
1245] and that the latter structure proje.cts lon is entirely confined to the telencephalon,
massively to the entorhinal area [1241,1242]. and comprises bilaterally the olfactory bulb,
Knowing that the entorhinal area projects to the olfactory tract and the basal olfactory
the hippocampal formation by way of the area (Fig. 9). The olfactory bulb represents
perforant pathway, and unaware that the su- the primary centre for reception of olfactory
bicular complex itself gives origin to the post- impulses. It is a small, flattened, ovoid body
commissural fornix, Shipley [1241] suggested that rests on the cribriform plate of the eth-
that projection from the presubiculum to the moid bone. Arising from the posterior pole
entorhinal area" closes" the Papez circuit. of the bulb, the olfactory tract passes back-
2. The indirect hippocampothalamic path- wards over the basal surface of the frontal
way via the mamillary body is supplemented lobe to become attached to the hemisphere.
by a substantial direct pathway [715]. At its site of attachment the tract bifurcates
3. The place of the cingulate gyrus in the into medial and lateral olfactory striae. The
circuit has been questioned. The anterior tha- initial parts of these striae border, together
lamic nucleus supplies all parts of the with the diagonal band of Broca, a territory
cingulate gyrus [592], but this projection is known as the anterior perforated substance.
reportedly not substantial [278]. There is ex- The medial olfactory stria extends towards
perimental evidence indicating that the most the subcallosal area on the medial surface of
rostral (area 32) and the caudal parts (area the frontal lobe; the lateral olfactory stria
23) of the cingulate gyrus project to the ento- passes laterally and then bends sharply
rhinal cortex and to the subicular complex around the limen insulae to enter the rostro-
[89, 1046, 1148]; however, these connections medial part of the temporal lobe (Fig. 196).
too are considered to be only moderately de- A small intermediate olfactory stria con-
veloped [592]. Moreover, the cingulate gyrus tinues the course of the olfactory tract for
appears to entertain primarily widespread re- a short distance and then fans out in the ante-
ciprocal connections with various neocortical rior perforated substance.
areas [74, 1046]. The anterior thalamic nucle- The basal olfactory area is formed by
us, on the other hand, has been found to those grisea that receive direct projections
contribute numerous fibres to the cingUlum from the olfactory bulb and includes the an-
bundle, many of which project directly to the terior olfactory nucleus, the olfactory tuber-
subicular complex [592, 1241, 1245]. In light cle, the prepiriform cortex and the cortical
of these findings it has been proposed that nucleus of the amygdala [1150, 1394]. The
the cingulate cortex - as opposed to the anterior olfactory nucleus consists of some
cingulum bundle - should be considered a small groups of neurons located in the caudal
component of primarily the neocortex rather part of the olfactory bulb, in the olfactory
than the limbic system [278] and that the tract and in the rostral end of the olfactory
"core" of the Papez circuit is constituted by trigone. The olfactory tubercle (so named be-
the subicular complex, the mamillary body, cause its homologue in macrosmatic mam-
the anterior thalamus and their interconnect- mals is marked by a prominent elevation) is
ing fibre systems [592]. Dagi and Poletti [278] represented by a thin sheet of grey matter,
recently even went one step further by not situated in the area of the anterior perforated
including the mamillary body. The circuit of substance. The prepiriform cortex can be di-
Papez and some related connections are dia- vided into two parts, a medial part overlying
grammatically represented in Fig. 206. the lateral olfactory stria as it travels laterally
across the base of the frontal lobe and a later- tal cortex (CPOF), which corresponds ap-
al part extending from the limen insulae over proximately to area 13. The second region
a small anteromedial area of the temporal is the lateroposterior orbitofrontal cortex
lobe. Caudally, the prepiriform cortex is re- (LPOF), which includes the posterior part of
placed by the superficial, cortical nucleus of area 12. The olfactory projection to the
the amygdala. Except for the anterior olfac- CPOF is transthalamic. The medial, magno-
tory nucleus, all grisea that receive direct cellular portion of the medial thalamic nucle-
olfactory afferents exhibit a laminated us (Mmc) receives afferents from the prepiri-
structure [1394]. form cortex and the medial part of the amyg-
The receptive elements of the olfactory ap- dala. The thalamopetal fibres emanating
paratus are slender, bipolar cells found in a from these two destinations of secondary ol-
specialised area of the nasal mucosa. These factory fibres pass medially, enter the medial
elements have extremely fine axons that col- forebrain bundle and ascend via the inferior
lect as small bundles, penetrate the cribriform thalamic peduncle to the thalamus [491, 492].
plate, and enter the olfactory bulb. Within The Mmc projects its fibres to the CPOF
the olfactory bulb, the primary olfactory [981]. Ethophysiological experiments have
fibres synapse with various types of cells, shown that the CPOF plays an essential role
among which the large mitral cells are the in olfactory discrimination [1354].
most conspicuous. For recent reviews on the The olfactory pathways to the LPOF are
microcircuitry of the olfactory bulb, the extrathalamic. Electrophysiological experi-
reader is referred to Macrides and Davis [845] ments suggest that these pathways have their
and Nieuwenhuys, [997]. The axons of these relay neurons primarily in the medial part
olfactory bulb cells pass backwards in the of the amygdala and in the lateral part of
olfactory tract and are distributed to the sec- the substantia innominata. The latter area is
ondary olfactory areas mentioned above. Ex- reported to receive afferents from the medial
perimental studies have conclusively shown amygdala as well as from the prepiriform
that the macroscopic elevation known as the cortex [1355]. In addition, Potter and Nauta
medial olfactory stria does not receive any [1099] have shown that the monkey prorhinal
secondary olfactory fibres [492, 1150, 1394]. cortex, which represents a lateral extension
After having distributed fibres to the anterior of the entorhinal cortex, receives afTerents
olfactory nucleus and the olfactory tubercle, from the prepiriform cortex and projects to
the entire secondary olfactory projection the LPOF.
enters the lateral olfactory stria. Following 2. Neurophysiological evidence indicates
that stria, the fibres reach the prepiriform that olfactory impulses reach the lateral hy-
and entorhinal cortices, and the cortical and pothalamus. However, there is no unanimity
medial amygdaloid nuclei, where they termi- regarding which pathways are involved. The
nate. presence of fibres originating from the olfac-
The most important tertiary olfactory pro- tory tubercle and the primary olfactory cor-
jections and the various routes along which tex which, after having descended with the
the rhinencephalon may discharge impulses medial forebrain bundle, terminate in the lat-
to the neocortex and the reticulomotor appa- eral hypothalamus, has been repeatedly re-
ratus of the brain stem are diagrammatically ported [1102, 1225]. It is, however, important
represented in Fig. 207. These pathways war- to note that Heimer [491, 492] has challenged
rant the following comments: the existence of a substantial olfactory pro-
jection to the lateral hypothalamic area
1. Two regions of the monkey frontal cor- (LHA). According to his observations, the
tex respond to electrical stimulation of the fibres that were supposed to terminate there
olfactory bulb and to odor stimulation [1355]. actually continue to the medial thalamic nu-
One region is the centroposterior orbitofron- cleus. After injecting HRP into the LHA of
1 Lateroposterior orbitofrontal cortex: "LPOF"

2 Centroposterior orbitofrontal cortex: "CPOF"
3 Fornix
4 Stria terminalis
5 Nucleus medialis thalami, pars magnocellularis
6 Nuclei septi
J0 Fila olfactoria
11 Epithelium olfactorium
13 Nucleus olfactorius anterior + tuberculum olfactorium
16 Cortex prepiriformis (area 51)
17 Nuclei corticalis + medialis amygdalae
18 Nuclei basalis + lateralis amygdalae
Fig. 207. Olfactory projections

2 Zona incerta
3 Nucleus preopticus magnocellularis
4 Area hypothalamica lateralis
8 Nucleus olfactorius anterior
9 Tuberculum olfactorium
10 Nucleus gyri diagonalis, pars ventralis
12 Cortex prepiriformis (area 5)
13 Locus coeruleus
Fig. 208. Afferents to the bulbus olfactorius

the monkey, Takagi [1355] found labelled The Limbic System at Large:
neurons in the accumbens nucleus and the A Functional Commentary
septum, but no labelled elements were ob- (Figs. 209-213)
served in either the olfactory tubercle or the
olfactory cortex. He therefore suggested that
olfactory information is probably conveyed In the preceding sections of this chapter de-
from the olfactory bulb through the septum tailed information has been provided on the
and/or the accumbens nucleus to the LHA. various limbic and olfactory centres and their
In several textbooks [151, 213, 266] the sep- fibre connections. In the present, final section
tum is considered to be an important nodal the attempt will be made to place this data
area in the olfactory projection system, re- into an overall functional perspective. To this
ceiving secondary olfactory fibres via a me- end it will be necessary to provide some addi-
dial olfactory tract. As mentioned above, it tional information and to introduce a few
has been established that there is no medial "new" neural entities, viz. the core of the
olfactory tract carrying secondary olfactory neuraxis and the median and lateral para-
fibres; however, this does not exclude the cores. It should be stated at the outset that
possibility of mUltisynaptic connections be- the ensuing synopsis, which is partly based
tween the olfactory bulb and the septum. The on a previous work of one of us (Nieuwen-
same holds true for the accumbens nucleus. huys [997]), focusses more on general mam-
3. The hippocampal formation and the malian conditions, than on specific human
amygdaloid complex represent relay centres aspects, and contains several speculative ele-
in polysynaptic routes by which the rhinence- ments.
phalon is connected with the septum, the hy-
pothalamus and ultimately with the reticulo- 1. Neuromediators. In the preceding sec-
motor apparatus of the brain stem. The suc- tions of this chapter no attempt has been
cessive links in these pathways have already made to indicate all of the neuromediators
been discussed. which have been reported to be present in
Experimental neuroanatomical studies the various limbic centres and pathways so
[160, 285, 286, 293, 950, 1320, 1396, 1549] far. However, recent surveys of the biochemi-
have shown that centrifugal fibres, origina- cal, histochemical and immunohistochemical
ting from many different regions, pass direct- literature [997, 1038] warrant the conclusion
ly to the olfactory bulb. These regions in- that within the confines of the limbic system
clude the anterior olfactory nucleus bilateral- an extraordinarily large number of different
ly, and on the ipsilateral side the olfactory neurotransmitters and neuropeptides are
tubercle, the prepiriform cortex, the nucleus present. It also appears that the limbic system
of the horizontal limb of the diagonal band, shares this particular feature with a number
the substantia innominata, the magnocellular of other central nervous entities, such as the
preoptic area, the LHA, the rostral part of ventral parts of the striatum, the thalamic
the zona incerta, the (noradrenergic) locus midline nuclei, the periaqueductal grey, the
ceruleus and the (largely serotoninergic) dor- parabrachial nuclei, the dorsal vagal com-
sal raphe nucleus. A substantial number of plex, the superficial zones of the spinal trige-
the central bulbar afferents is cholinergic. minal nucleus and of the spinal dorsal horn,
Most of these fibres originate from neurons and the spinal substantia intermedia centra-
situated in the nucleus of the horizontal limb lis. Moreover, it has become clear that all
of the diagonal band, but such elements are of the "neuromediator-rich" centres are in-
also found in the substantia innominata. The terconnected by a number of "neuromedia-
nucleus of the horizontal limb of the diagonal tor-rich" fibre systems. These fibre systems,
band contains, in addition, a number of which are largely composed of thin, unmye-
GABA-ergic bUlbopetal neurons [1549]. linated, varicose axons, include the dorsal
longitudinal fascicle of Schlitz, the medial spectively. The posterior pituitary hormone
forebrain bundle, the diagonal band of Bro- vasopressin, which is involved in the mainte-
ca, the stria terminalis, the tubero-infundibu- nance of the body fluid balance, is produced
lar tract and the hypothalamohypophysial in the supraoptic and paraventricular hypo-
projection (cf. Nieuwenhuys [997], Fig. 87). thalamic nuclei. The medial preoptic area
2. Overall functional significance of the and the parvocellular part of the paraventri-
"neuromediator-rich" centres. Current neu- cular nucleus are involved in mediating thirst
rophysiological and neuroethological knowl- and drinking behaviour, whereas numerous
edge, part of which has been reviewed in the centres, including the amygdaloid complex,
present and previous chapters, enables us to the medial preoptic area, the dorsomedial
attach functional labels to practically all of and ventromedial hypothalamic nuclei, the
the limbic and other "neuromediator-rich" lateral hypothalamic area and the periaque-
centres. Many of these centres playa key role ductal grey, are known to playa role in the
in the regulation of visceral effector mecha- control of feeding.
nisms. Thus, the anterior, paraventricular Attack and defense-escape reactions, i.e.
and dorsomedial hypothalamic nuclei, the agonistic behaviours, involve highly charac-
medial preoptic area and the bed nucleus of teristic patterns of autonomic and skeletomo-
the stria terminalis have been implicated in tor responses (cf. Descending Reticular Sys-
cardiovascular control. Modification of re- tem, p. 279, [889]). Stimulation of a particu-
spiratory functions can be induced by stimu- larly large number of neuromediator-rich
lation of the anterior hypothalamic nucleus centres has been found to elicit these re-
and the bed nucleus of the stria terminalis. sponses. We confine ourselves here to men-
Finally the motility and secretory activity of tioning the hippocampus, the amygdaloid
the digestive thract is influenced by, among complex, the bed nucleus of the stria termina-
other structures, the periaqueductal grey, the lis, the lateral septal nucleus, several hypo-
ventromedial and dorsomedial hypothalamic thalamic centres, including the anterior and
nuclei and the lateral hypothalamic area. The ventromedial nuclei, and the periaqueductal
nucleus centralis amygdalae, the lateral para- grey.
brachial nucleus and the dorsal vagal nucleus Locomotion or forward progression is an
are involved in all three functions. essential component of various adaptive
The entire preoptico-hypothalamic con- behaviours, like fight and flight reactions and
tinuum is crucially involved in the mainte- food and water procurement. Several neu-
nance of homoiostasis, i.e. the regulation of romediator-rich centres, including the hippo-
the milieu interieur. This task is accomplished campus, the amygdaloid complex, the nucle-
by complex action patterns, encompassing us accumbens, and the medial and lateral
(a) visceromotor responses, (b) modulation preoptic areas, have been shown to be in-
of the release of anterior and posterior pitui- volved in the initiation and modulation of
tary hormones, and (c) feeding and drinking locomotor activity [1339].
behaviour. The most important centres in- The substantia gelatinosa and its rostral
volved in visceromotor responses have al- continuation in the spinal trigeminal nucleus
ready been mentioned. Several anterior pitui- are recipient centres of nociceptive stimuli,
tary hormones, including the growth hor- whereas the periaqueductal grey plays a key
mone and the adreno-corticotropic hormone, role in the modulation of nociception.
play an important role in metabolic regula- Several neuromediator-rich centres are in-
tion [1123]. The neurons producing the hypo- volved in reproductive behaviour. The cortico-
physiotropic peptides which regulate the se- medial nuclei of the amygdaloid complex, the
cretion of these hormones are mainly located bed nucleus of the stria terminalis and the
in the preoptic region and in the parvocellu- medial preoptic area playa role in the regula-
lar part of the paraventricular nucleus, re- tion of male sexual behaviour, whereas the
medial preoptic area, the ventromedial hypo- volved most directly in processes aimed at
thalamic nucleus and the periaqueductal grey the survival of the individual organism and
form nodal points in the circuitry underlying of the species, and which, hence, may be des-
feminine sexual behaviour. It is important to ignated as the core of the neuraxis [997].
note that (a) the release of gonadotropic hor- 3. One of us (Nieuwenhuys [997]) has ad-
mones from the pituitary gland is steered by duced evidence suggesting that paracrine or
the hypothalamus, and that (b) numerous non-synaptic neurotransmission may well
neurons in the central nervous system are tar- playa prominent role in the core of the neu-
gets of the gonadal steroid hormones which raxis. In one typical core region, viz. the par-
are produced under the control of the gona- ticularly neuropeptide-rich median eminence,
dotropic hormones. Stumpf and his asso- true synapses are entirely lacking [183, 184,
ciates [670, 1197, 1305, 1306, 1308, 1309, 1038].
1310, 1311] extensively studied the distribu- 4. Paracore zones. At the level of the brain
tion of neurons concentrating sex steroids in stem the core of the neuraxis has two ad-
the central nervous system of several mam- juncts, which may be designated as the medi-
malian groups. They found that (1) the male an paracore and the (bilateral) lateral para-
steroid hormone testosterone and the female core (Fig. 209). The median paracore is con-
steroid hormone estradiol are present in ap- stituted by the series of raphe nuclei, which
proximately the same brain regions and that extends throughout the brain stem (Figs. 155
(2) most of the testosterone and estradiol tar- and 183). In most places the raphe nuclei are
get neurons are situated close to the ventricu- directly adjacent to the core region, and in
lar surface, i.e. in a zone which they regarded some they even penetrate into it. Moreover,
as a phylogenetic ally ancient part of the neu- fibres of the core region project heavily to-
raxis. Stumpf [1306] held that this "periven- wards most raphe nuclei.
tricular brain" fulfils a neuroendocrine func- The lateral paracore consists of a series
tion throughout and suggested that a certain of grisea which in most, though not all, levels
proportion of the target cells of the sex hor- of the brain stem extend from the core region
mones are directly involved in the regulation ventrolaterally into the tegmentum. At the
of the production of hypothalamic regulating mesencephalic level this series includes the
hormones, whereas others may serve as continuum formed by the lateral part of the
threshold modulators on either the sensory tegmental grey, the substantia nigra and the
(estradioJ) or the motor side (testosterone). area tegmentalis ventralis. In the rhombence-
Remarkably, the set of estrogen- and andro- phalon it is the locus coeruleus or A6 group,
gen-hormone-concentrating neuronal groups the nucleus subcoeruleus (A6sc), the
which Stumpf included in his "periventricu- Kolliker-Fuse nucleus, the nucleus reticularis
lar brain" closely corresponds to the set of parvocellularis, the area reticularis superfi-
neuromediator-rich centres (cf. Nieuwenhuys cialis ventrolateralis and the cytoarchitecton-
[997], Fig. 88). ically ill-defined cell groups A1, A2, AS, A7,
The sense of smell and its central appara- C1, C2 and Ch5 which constitute this series.
tus, finally, plays an important role in both All of the rhombencephalic centres men-
feeding and reproductive behaviour. tioned form part of or are embedded in the
Taking the morphological and functional lateral reticular zone (Fig. 154).
data presented above together we venture the The median and lateral paracores have
conclusion that a set of neuromediator-rich several features in common:
centres and pathways, which corresponds a) As already mentioned, they are both
partly to the limbic system and which has directly continuous with the core region.
many features in common with the steroid- b) Both zones contain large numbers of
hormone-concentrating periventricular brain, monoaminergic elements. In the median para-
constitutes a functional complex which is in- core numerous serotoninergic elements are
c
A1-A7 Noradrenergic cell groups Ie Locus coeruleus (A6)
A8-A10 Dopaminergic cell groups leb Longitudinal catecholamine bundle
B1-B7 Serotoninergic cell groups lpc Lateral paracore
C1 , C2 Adrenergic cell groups mpc ~edian paracore
dacp Dorsal ascending catecholamine pathway pc [Paracrine ?] core
ddsp Dorsal descending serotonin pathway sca Subcoeruleus area (A6)
fld Fasciculus longitudinalis dorsalis of Schiitz vacp Ventral ascending catecholamine pathway
fr Formatio reticularis vdsp Ventral descending serotonin pathway
ip Interpeduncular nucleus
Fig. 209 A-C. Diagrammatic frontal sections through the human brain stem at mesencephalic (A),
rostral metencephalic (B) and myelencephalic levels (C) to show the positions of the [paracrine?]
core, the median paracore and the (bilateral) lateral paracores. In the [paracrine?] core the position
of the coarsest fibres in the fasciculus longitudinalis dorsalis is indicated; however, it is known that
this region contains large numbers of thin, unmyelinated fibres. Further explanation is given in the
text. From Nieuwenhuys [997]
found, but in the lateral paracore catechola- these elements might act as both neurons and
minergic neurons prevail. These include ad- endocrine-neural transducer cells.
renergic elements, which are found exclusive- g) Both paracores exchange numerous ax-
ly in the caudal part of the zone (cell groups ons with the core region. To give a few exam-
C1 and C2; Figs. 154, 159, 189), noradrener- pies: Numerous telencephalic and dience-
gic neurons, which occur throughout the phalic core regions project via the medial
length of the rhombencephalon (cell groups forebrain bundle to the paracore zones in the
A1-A7; Figs. 154, 159, 188, 189) and dopa- brain stem (Figs. 197B, 198B, 201) and many
minergic neurons, concentrated in the ventral of these connections are reciprocated by as-
part of the mesencephalic portion of the lat- cending projections (Figs. 159 and 198A).
eral paracore. The periaqueductal grey sends numerous
c) Both paracores lie clearly beyond the fibres to several raphe nuclei (Fig. 183B) and
trajectories of the large, myelinated ascend- the area reticularis superficialis ventrolatera-
ing and descending pathways. lis is reciprocally connected with the parabra-
d) Both paracores contain assemblies of chial nuclei, as well as with the nucleus of
thin, longitudinally oriented fibres. Ascend- the solitary tract (Figs. 157, 159, 186).
ing and descending serotoninergic axons Summarizing, it may be stated that the
(Figs. 155 and 183) contribute substantially neuromediator-rich, and in particular the
to the median paracore fibre assembly, neuropeptide-rich, core of the neuraxis is
whereas the large longitudinal catecholamine closely linked to two paracore zones, which
bundle (Figs. 159, 188, 189) occupies a cen- contain numerous monoaminergic cells and
tral position in the lateral paracore. More- fibres, and that there is evidence suggesting
over, the median and lateral paracore fibre that in the core as well as in the paracore
assemblies are both fostered by numerous zones large numbers of neurons are involved
fibres of telencephalic and diencephalic ori- in endocrine-neural transduction and/or neu-
gin which, after having followed the medial roendocrine regulation [997]. It should be
forebrain bundle through the lateral hypo- noted that the core-paracore concept just epi-
thalamus, enter the brain stem. tomized shows many points of resemblance
e) Both paracores contain sets of cells giv- to the "allocortex-brainstem-core (A-B-C)
ing rise to networks of fibres that pervade circuitry concept of endocrine-autonomic in-
virtually all grisea of the neuraxis, i.e. the tegration and regulation", enunciated by
serotoninergic cells in the median paracore Stumpf and lennes [1307].
and the catecholaminergic cells in the lateral Having introduced the terms" core of the
paracore. neuraxis", "median paracore" and "lateral
f) The perikarya and dendrites of the paracore" it now remains to present a brief
monoaminergic neurons contained within the overview of the structural and functional or-
median and lateral paracores frequently con- ganization of these entities. For practical rea-
tact blood vessels and tanycyte-like glia cells, sons the core of the neuraxis will be subdi-
i.e. elements with cell bodies forming part vided into a rostral, telencephalic part and
of the ependymal ventricular lining and pro- a caudal, mainly subtelencephalic part.
vided with one or a few long, peripherally 5. The caudal part of the core of the neur-
extending processes. Felten and colleagues axis includes the preoptic region, which is
[183, 273, 341, 342, 344], who first described of telencephalic origin, the entire hypothala-
these relationships, considered it likely that mus, the periaqueductal grey, the pontine
the direct neuron-vascular contacts and the central grey, the parabrachial nuclei, the dor-
tanycyte shafts represent communication sal vagal complex, which encompasses the
channels along which substances carried by nucleus solitarius and the dorsal motor vagal
the blood and cerebrospinal fluid influence nucleus, and the spinal substantia intermedia
monoaminergic neurons. They suggested that centralis (Fig. 210). The preoptic region and
the hypothalamus constitute together one and the latter projects via the fasciculus lon-
structural and functional complex, which gitudinalis dorsalis to the preoptic region and
may be designated as the preopticohypotha- to some hypothalamic centres. These direct
lamic continuum (PRC). The medial fore- projections from the nucleus solitarius are ac-
brain bundle may be considered as the princi- companied by several parallel pathways that
pal conduction channel of this continuum, are probably also involved in the transmis-
whereas the fasciculus longitudinalis dorsalis sion of interoceptive information to the
of Schutz interconnects the various subdien- PRe. These pathways, which also follow the
cephalic centres of the core and also contains fasciculus longitudinalis dorsalis, are synapti-
fibres which link these centres with the hy- cally interrupted in the parabrachial nuclei
pothalamus. or in the periaqueductal grey. Interoceptive
The PRC is active in the following func- information attaining the central nervous
tional and behavioural domains: (1) body system via the glossopharyngeal and vagal
fluid homeostasis, thirst and drinking behav- nerves may also by relayed to the area reticu-
iour, (2) metabolic regulatory functions and laris superficialis ventrolateralis, a highly im-
feeding behaviour, (3) thermoregulation, (4) portant lateral paracore component, before
neural regulation of the cardiovascular sys- it is conveyed to the PRe.
tem, (5) modification of respiratory func- Exteroceptive information may reach the
tions, (6) agonistic behaviour, (7) the genera- PRC via different routes. Direct visual input
tion and regulation of circadian rhythms, and attains the hypothalamus via the retino-
(8) reproductive and maternal behaviour. As hypothalamic tract, which terminates in the
has been discussed in previous sections of the suprachiasmatic nucleus. This centre, which
present chapter each of these functions can projects to various intra- and extrahypotha-
be related to one or more PRC loci or lamic core centres, may be designated as an
centres. The functional state of these centres endogenous circadian pacemaker. It influ-
is influenced by humoral information and by ences a number of circadian rhythms, includ-
neural messages from the internal milieu as ing patterns of general activity, foraging
well as from the external environment. behaviour and hormonal secretions. The ol-
As regards humoral information, the sodi- factory system has a relatively direct access
um concentration of the blood, as well as to the PRC via the cortical and medial amyg-
the concentration of glucose, free fatty acids daloid nuclei. The latter are directly innervat-
and other nutrients are monitored by special- ed by the olfactory bulb and project via the
ized hypothalamic and extrahypothalamic stria terminalis to several preoptic and anteri-
(e.g. nucleus solitarius) neurons. Moreover, or hypothalamic nuclei (Fig. 211). Righer
numerous hormones, like angiotensin II and order visual, auditory, tactile and gustatory
the gonadal and adrenal steroids, act directly information attains the PRC via long poly-
on endocrine - neural transducer cells which synaptic conduction lines that involve the
are found in large numbers in the core of various "lemniscal" systems and their corti-
the neuraxis. The circumventricular organs, cal projections, the primary sensory cortices,
i.e. loci which typically lack a blood-brain the various modality specific association cor-
barrier, represent trigger zones for humoral tices, and finally the amygdaloid complex
factors. These small organs, which constitute and its major efferent systems. Multimodal
important nodal points in the circuitry of the sensory information is conceivably conveyed
core region, may also be influenced by cere- to the PRC from the mesencephalic reticular
brospinal fluid-borne substances. formation (Fig. 156) and, in a highly pro-
Cardiovascular, gastrointestinal and respi- cessed form, from the hippocampus via its
ratory information is conveyed via the glos- various subcortical efferents (Fig. 205).
sopharyngeal and vagal nerves to the caudal In response to the afferent stimuli just dis-
interoceptive part of the nucleus solitarius cussed the PRC exerts modulatory and regu-
1 Organum subfornicale
2 Organum vasculosum laminae terminalis
3 Fibrae retinohypotbalamicae
4 Eminentia mediana
5 Hypophysis, lobus anterior
6 Hypophysis, lobus posterior
7 Preoptico-hypotbalamic continuum
10 Fibres ascending from tbe mesencephalic reticular formation
14 Nucleus dorsalis nervi vagi} d 1 1 I
IS Nucleus solitarius orsa vaga comp ex
16 Area postrema
17 Substantia intermedia centra lis
Fig. 210. The caudal part of the core of the neuraxis

1 Fornix
2 Stria terminalis
4 Nuclei septi
5 Nuclei gyri diagonalis
6 Preoptico-bypothalamic continuum
9 Nuclei cortical is + medialis amygdalae
10 Nuclei basaUs + lateraUs amygdalae
12 Formatio bippocampi
13 Areae 28, 35, 36
14 Unimodal cortical association areas
16 Nucleus rapbes dorsalis
18 Locus coeruleus
21 Nucleus raphes pallidus
Fig. 211. The rostral part of the core of the neuraxis

latory influences on (a) its own humoral and It is noteworthy that all of these cortico-
neural inputs, (b) the endocrine system, (c) amygdaloid projections are reciprocated by
the visceromotor apparatus, and (d) the ske- amygdaloid efferents. The bed nucleus of the
letomotor apparatus. All of these activities stria terminalis and the septal complex are
are mostly integrated into complex action reciprocally connected with the amygdala,
patterns. and the hippocampus, respectively.
As regards the control of humoral input, The subcortical telencephalic nuclei under
it has been found that all of the circumventri- discussion do not exert their influences on
cular organs receive strong projections from the lower parts of the brain only via the PHC.
other core centres. It is interesting to note It has been demonstrated that fibres emanat-
in this context that the area postrema receives ing from the bed nucleus of the stria termina-
a direct projection from the dorsal part of lis and from the amygdala project directly
the hypothalamus [556]. Descending fibres, to a large number of core and paracore
along which the PHC presumably controls centres in the brain stem, among which the
its own neural input, include hypothalamic periaqueductal grey, several raphe nuclei, the
projections to the nucleus of the solitary tract parabrachial nuclei, the locus coeruleus and
and to the parabrachial area and the peri- the dorsal vagal complex (Figs. 198 Band
aqueductal grey. Hypothalamic fibres also 201). Conversely, ascending projections aris-
descend to the marginal zone of the spinal ing from the nucleus solitarius, the area reti-
dorsal horn, where they may influence the cularis superficialis ventrolateralis, the para-
transmission of nociceptive impulses, and to brachial region, the locus coeruleus and the
the substantia intermedia centralis, another periaqueductal grey bypass the PHC to ter-
spinal core region. Hypothalamic fibres de- minate in the amygdala and in the bed nucle-
scending to the dorsal motor vagal nucleus, us of the stria terminalis (Figs. 198 A, 199),
as well as to other preganglionic cell groups and most of these brain stem centres also
of both the sympathetic and parasympathetic project to the septal region.
divisions of the autonomic system, mediate The allocortical hippocampal formation is
influences on heart rate, blood pressure, and reciprocally connected with the neocortex on
gastrointestinal motility and secretion. How- the one hand, and with the amygdala, the
ever, as will be discussed below, the PHC bed nucleus of the stria terminalis and the
exerts its influences on the skeletomotor sys- septal complex, i.e. the three subcortical te-
tems via the median and lateral paracores. lencephalic core centres, on the other hand
6. The rostral part of the core of the neu- (Figs. 203, 204, 206). Moreover, the hippo-
raxis is constituted by the septal region, the campus projects massively to the corpus ma-
amygdala and the bed nucleus of the stria millare, but its direct output to what is called
terminalis, subcortical telencephalic centres here the PHC is confined to a projection to
which together form a more or less continu- the ventromedial hypothalamic nucleus and
ous chain around the most rostral part of it is also noteworthy that the hippocampus
the PHC, and by the allocortical hippocam- does not entertain any direct fibre relations
pal formation (Fig. 211). with more caudal parts of the brain. With
The most salient common feature of the regard to neocortical afferents, it may be stat-
subcortical centres mentioned is that all of ed that the same unimodal association areas
them entertain strong and bidirectional con- that project to the amygdala are major
nections with the adjacent PHC. The amyg- sources of input to the hippocampal forma-
dala receives both direct and indirect inputs tion. However, whereas the various sensory
from the olfactory bulb and, as already men- modalities remain separated at the level of
tioned, higher order visual, acoustic, tactile the amygdala, the efferents from the unimo-
and gustatory information from the pertinent dal association areas directed towards the
modality-specific cortical association areas. hippocampus converge upon the entorhinal
and perirhinal cortices and these polymodal the habenulointerpeduncular tract. Within
sensory areas project in their turn to the hip- this system two separate subsystems, viz. the
pocampus (Fig. 211). Because of this particu- medial and lateral habenular paths, can be
lar position in relation to the flow of sensory distinguished. The medial habenular path,
information, Swanson [1326] considers the which is synaptically interrupted in the later-
hippocampus as a supramodal association al septal, medial habenular and interpedun-
area. cular nuclei, conveys information of hippo-
It is worth noting that the insular cortex, campal antecedency. The lateral habenular
which encompasses a viscerosensory area, path, which originates mainly from the later-
projects to the amygdala as well as to the al preopticohypothalamic area, is synapti-
hippocampus. Hence, it may be concluded cally interrupted in the lateral habenular
that highly processed information concerning nucleus, but bypasses the interpeduncular
both the external and internal environment nucleus. The major efferent pathway from
attains the hippocampus and the amygdala, the dorsal raphe and central superior nuclei
and is ultimately funnelled, partly via the sep- ascends to the diencephalon and joins the
tal complex and the bed nucleus of the stria medial forebrain bundle in the lateral hy-
terminalis, to the PHC, and that conversely pothalamus. Fibres leaving this main ascend-
information concerning the functional state ing pathway are distributed to a large
of the various PHC mechanisms is trans- number of diencephalic and telencephalic
ferred to the several cortical association areas structures, with a distinct quantitative prefer-
via sequential return pathways. The func- ence for core regions.
tions of this highly complex bidirectional cir- The rostral raphe nuclei, and the seroton-
cuitry have appeared difficult to clarify and inergic cell groups B6-B8 embedded therein,
can certainly not be epitomized in a few have been implicated in a variety of func-
words. However, clinical and experimental tions, such as the regulation of sleep, ther-
evidence suggest that the hippocampus and moregulation, neuroendocrine modulations,
the amygdala play an important role in mem- aggression and "enabling the organism to ar-
ory function. Past experience is of great im- range or tolerate delay before acting" [1274].
portance for an optimal transfer of many spe- We confine ourselves to stating that the as-
cific motivational states or drives into goal- cending raphe projections, particularly dur-
oriented adaptive behavioural patterns. ing their course through the lateral hypotha-
Moreover, newly arriving sensory stimuli lamus, are in an optimal position to influence
may be affective (e.g. fearful) in light of past many different behavioural mechanisms.
experience and lead to feelings which find The nucleus raphes magnus receives affer-
their expression in emotional behaviour as ents from the medial part of the hypothala-
such, or in emotional components of behav- mus, the periaqueductal grey and the nucleus
ioural patterns like defence or flight. These raphes dorsalis, and projects primarily to the
few examples may suffice to give some idea caudal part of the spinal trigeminal nucleus
how the higher core centres influence the exe- and to the superficial zone of the spinal dor-
cution of action patterns. sal horn. This descending projection plays an
7. The basic circuitry of the median para- important role in the regulation of the trans-
core, which encompasses the raphe nuclei and mission of nociceptive stimuli and may also
their afferent and efferent connections, is control the input of other sensory stimuli.
summarized in Fig. 212. All of the raphe nu- The nucleus raphes pallidus also receives af-
clei receive their principal input from one or ferents from the medial hypothalamic zone
more core centres. The dorsal raphe and cen- and projects chiefly to the anterior horn of
tral superior nuclei receive massive afferents the spinal cord. The substantial serotoniner-
via the dorsal diencephalic conduction sys- gic component of the latter projection is
tem, which includes the stria medullaris and known to facilitate all types of somatomotor
---------
1 Fornix
2 Stria medullaris
3 Diencephalic core tructure
4 Nuclei septi
5 Nuclei gyri diagonali
6 Preoptico-hypothalamic continuum
8 Subiculum
9 ucleus habenulae latera lis
10 ucleus habenulae medialis
12 ucleus raphes dorsalis
13 ucleus interpeduncularis
14 ucleus centralis superior
15 ucleus raphes magnus
16 Nucleus raphe pallidus
17 Nucleus raphes obscurus
18 Nucleu spinaLis nervi trigemini
+ substantia gelatinosa spinalis
Fig_ 212_ The median paracore

1 Fasciculus telencephaIicus medialis

2 Lateral paracore pathway
3 Mesencephalic" attack" sites
4 Mesencephalic locomotor area
5 Area subcoerulea (A6 sc)
6 Locus coeruleus
7 M-region of Holstege et al. [537]
8 L-region of Holstege et aI . [537]
9 Pontine" attack" si te
10 Nucleus of Kiill iker- Fuse
11 Pontine swallowing centre
12 Area reticularis superlicialis ventrolateralis
13 Dorsomedial swallowing area
14 Ventrolateral swallowing arca
15 Dorsal respiratory group
16 Ventral respiratory group
At , A2 etc. Noradrenergic cell groups
Cl , C2 Adrenergic cell groups
Fig. 213. The lateral paracore

neurons [35]. Thus, mass excitation of the periaqueductal grey projects to the lateral
bulbo-spinal projections from the nucleus part of the reticular formation and the para-
raphes magnus and the nucleus raphes palli- brachial nuclei distribute numerous fibres to
dus may be expected to simultaneously sup- the area reticularis superficialis ventrolatera-
press sensory input and enhance motor lis. Given the facts that (1) all core centres
responsiveness. contain several different populations of pep-
In summary, it seems likely that the medial tidergic neuronal elements, and that (2) a
zone of the hypothalamus, via descending considerable number of these centres project
pathways to the caudal parts of the median to the lateral paracore zone, it is not
paracore, is able to exert general, level-setting surprising that within the central longitudinal
influence on sensory and motor parts of the pathway of this zone several different sets
spinal cord [534]. of peptidergic fibres are present. Axons, con-
8. The lateral paracore consists of a series taining vasopressin, oxytocin, somatostatin,
of tegmental grisea, arranged along and corticotropin, corticotropin releasing factor
around a large, thin-fibred pathway. This and neurotensin have all been observed with-
pathway extends throughout the brain stem. in the trajectory of this pathway [997].
Rostrally it is directly continuous with the Physiological studies have shown that the
medial forebrain bundle and caudally it ex- lateral paracore contains numerous centres
tends into the lateral funiculus of the spinal which can be clearly related to specific physi-
cord (Fig. 213). This lateral paracore pathological or behavioural mechanisms. All of
way is composed of (a) ascending and de- these centres project, either directly or via
scending fibres originating from the various intercalated elements, to visceromotor or ske-
lateral paracore grisea, (b) fibres which de- letomotor neuron pools. These functionally
scend from telencephalic and diencephalic defined centres receive their principal affer-
core regions, and (c) fibres interconnecting ents from one or more core districts and
the brain stem segment of the core with the these, mainly descending, connections are of-
lateral paracore. Prominent among the fibres ten reciprocated by ascending fibres. Core
contributed by the lateral paracore are nor- and lateral paracore constitute one single
adrenergic and adrenergic axons and axonal continuum of which the complex medial fore-
ramifications, the parent cell bodies of which brain bundle and its substantial caudolateral
are confined to the rhombencephalon. To- extension into the brain stem constitute the
gether these fibres constitute a subset of the principal communication channel. This con-
lateral paracore pathway, which is known as tinuum harbours a large number of discrete
the longitudinal catecholamine bundle (Figs. functional circuits within which the lateral
159, 188, 189). The fibres descending from paracore centres referred to above form es-
higher levels of the brain originate mainly sential links. Some of these circuits may now
from the hypothalamus, particularly the lat- be briefly discussed.
eral hypothalamic area ([534,553,557, 1412]; The cardiovascular regulating circuit en-
Fig. 197B), and from the bed nucleus of the compasses afferents which enter the central
stria terminalis ([542, 1412]; Fig. 198B), and nervous system with the glossopharyngeal
the nucleus centralis amygdalae ([550, 1103, and vagal nerves and terminate in the nucleus
1412]; Fig. 201). The interrelationships be- solitarius. Efferents from the latter project
tween the core and the lateral paracore zones massively to a cardiovascular neuronal group
within the brain stem are manifold. To give within the area reticularis superficialis ven-
a few examples: The locus coeruleus and sev- trolateralis (ARSVL), which in turn projects
eral other noradrenergic cell groups project to the nucleus intermediolateralis in the thor-
to the periaqueductal grey, the parabrachial acic spinal cord. This pathway represents the
nuclei and the dorsal vagal complex morphological substrate of baroreceptive
(Fig. 158, 159, 189) whereas conversely, the and chemoreceptive reflexes. A comparable
circuit subserves the regulation of respira- in the rostrocaudal arrangement of the perti-
tion, but the efferent limb of this reflex arc nent centres along the rostral core - lateral
is formed by ARSVL fibres descending to paracore axis.
the nucleus phrenicus at the C4-C6 level and As has already been mentioned, locomo-
the neurons innervating the intercostal mus- tion is an essential component of the procure-
cles in the thoracic cord. Three additional ment phase of various adaptive behaviours.
rhombencephalic lateral paracore centres The functionally defined mesencephalic loco-
form part of the respiratory regulatory net- motor area, which roughly corresponds to
work, viz. dorsal and ventral respiratory the nucleus tegmentalis pedunculopontinus,
groups and the Kolliker-Fuse nucleus. The as well as to the cholinergic Ch5 group
dorsal and ventral respiratory groups are lo- (Figs. 100 and 154), is a prominent compo-
cated ventrolateral to the nucleus solitarius nent of the lateral paracore. The activity of
and close to the nucleus ambiguus, respec- this area is strongly influenced by a number
tively [1175]. Both of these centres project of core regions. More specifically, the nucleus
directly to respiratory motoneurons [1030]. accumbens, which receives its principal affer-
The pontine Kolliker-Fuse nucleus, which is ents from the amygdala and the hippocam-
considered to represent the pneumotaxic pus, projects directly and via the substantia
centre, projects strongly to the lower medulla innominata to the mesencephalic locomotor
oblongata (Fig. 186), but hardly to respirato- regions. Several other centres, including the
ry spinal motoneurons [1030]. Physiological medial and lateral preoptic region, the zona
experiments indicate that the rhombence- incerta and the area tegmentalis ventralis,
phalic cardiovascular and respiratory centres also form part of the circuitry converging
are under the control of centres forming part upon the mesencephalic locomotor area
of the rostral core, such as the nucleus centra- [1338, 1339]. It may be added that the nucleus
lis amygdalae and the bed nucleus of the stria accumbens receives, in addition to its strong
terminalis. Both of these centres are known afference from the hippocampus and the
to project via the medial forebrain bundle amygdala, also a substantial input from the
and the lateral paracore pathway to the nu- area tegmentalis ventralis, a largely dopamin-
cleus solitarius (Figs. 198B, 201). ergic centre, which forms part of the mesen-
Two pontine lateral paracore centres, the cephalic lateral paracore. Activation of the
M and L regions of Holstege and collabora- projection from the area tegmentalis ventralis
tors [537], are involved in the reguiation of to the nucleus accumbens enhances locomo-
micturition. As mentioned in a previous tor activity [1090].
chapter, the M region projects to the para- Swallowing, i.e. the final act of the execu-
sympathetic preganglionic neurons in the tion or consummatory phase of foraging
sacral cord that innervate the detrusor mus- behaviour, is centrally programmed by three
cle, whereas the L region sends its afferents lateral paracore centres, viz. the dorsomedial,
to the, also sacral, somatomotor nucleus of ventrolateral and pontine swallowing areas.
Onuf. The latter supplies, among other mus- The dorsomedial area is situated in the imme-
cles, the urethral sphincter. The M region, diate vicinity of the nucleus solitarius, where-
which roughly corresponds to the mictura- as the ventrolateral area is embedded in the
tion reflex centre of Barrington, is known to reticular formation surrounding the nucleus
receive numerous afferents from the anterior ambiguus [29, 617, 914]. The pontine swal-
hypothalamic area [534]. lowing area is also located in the reticular
Turning now to some behavioural mecha- formation, and lies just dorsal to the superior
nisms it should be stated first of all that the olivary complex [536]. The efferents of these
initiation, procurement and consummatory areas pass to the motor trigeminal, hypoglos-
phases through which motivated behavioural sal and ambiguus nuclei. The specific projec-
patterns in general pass is roughly reflected tions impinging upon the three swallowing
areas are unknown. However, it may be ex- elicited by electrical stimulation of the lateral
pected that the various core centres involved hypothalamus and of some loci situated in
in eating (e.g. the amygdaloid complex, the the mesencephalic and pontine lateral para-
medial preoptic area, the dorsomedial and core [1234, 1260]. Experimental neuroana-
ventromedial hypothalamic nuclei and the tomical studies have revealed that all of these
lateral hypothalamic area) and drinking (e.g. "attack sites" are interconnected and that
the subfornical organ, the medial preoptic the pontine locus projects to the vicinity of
area, the zona incerta and the lateral hypo- the principal sensory and motor trigeminal
thalamic area) form part of a network that, nuclei [235, 236]. Moreover, it has been
via the main core - lateral paracore channel, shown that the mesencephalic as well as the
ultimately converges upon these swallowing pontine attack sites receive afferents from nu-
areas. Other neuron pools governing the var- merous core regions, including the amygdala,
ious stereotyped ingestive performances pre- the bed nucleus of the stria terminalis, several
ceding swallowing are most probably also in- hypothalamic regions and the periaqueductal
tegrated in this network. grey [236, 1219].
Agonistic behaviour, as for instance de- The foregoing overview renders it plausi-
fense reactions, involves skeletomotor as well ble that within the core - lateral paracore
as visceromotor responses. The morphologi- continuum a number of function-specific sys-
cal substrate of this behaviour includes the tems are present. Although long through-
amygdala and a specific pathway, which conducting projections are not lacking within
passes from that centre to the ventromedial this continuum, most of the functional sys-
hypothalamic nucleus and thence, via the an- tems singled out are characteristically com-
terior hypothalamic area, to the periaqueduc- posed of series of consecutive neuronal links.
tal grey [365, 366]. The premamillary nuclei, Transmission through such neuronal chains
which are likewise intercalated between the depends in great part on how each station
amygdala and the periaqueductal grey, have along the way is modulated by the influences
also been implicated in agonistic behaviour converging upon them, and it stands to rea-
[825]. The caudal part of the periaqueductal son that the influence of systems having ac-
grey has been demonstrated to contain an cess to all of the sequential stations will be
area from which an integrated skeletomotor considerable. Another point that should be
and visceromotor defense reaction can be eli- mentioned at this juncture is that most
cited. The autonomic component of this be- behaviours can only be safely or optimally
havioural pattern has been designated as the executed within the setting of general states
visceral alerting response (cf. Descending Re- such as relaxation or stress, and that in some
ticular System, p. 279). Physiological evi- situations actions of vital priority have to be
dence strongly suggests that the ARSVL, i.e. continued, despite the arrival of distracting
the most caudal part of the lateral paracore, sensory cues, whereas in other situations sur-
receives a direct input from hypothalamic vival requires the prompt cessation of a given
and mesencephalic " defence areas", and behaviour, e.g. in light of threatening stimuli.
plays an essential role in the effectuation of It is in the functional domain just outlined
this complex response, which includes respi- that the noradrenergic and adrenergic neuro-
ratory activation, an increase in blood pres- nal popUlations, which are embedded in the
sure and a redistribution of cardiac output lateral paracore, fulfil a regulatory role.
in favour of skeletal muscles. The locus coeruleus, which comprises the
With regard to the neural organization of major aggregation of noradrenergic neurons
the skeletomotor components of agonistic in the central nervous system, projects to
behaviour, it may be noted that attack behav- widely varied target areas ranging from the
iour without signs of autonomic arousal - cerebral cortex to the spinal grey matter.
hence designated as "quiet attack" - can be Such an ubiquitous distribution of efferents
suggests a very global function. It has been fibres ascending from these medullary
suggested that this locus represents a central centres, as for instance the projection from
analogue of the peripheral sympathetic sys- Ai to the parvocellular part of the paraven-
tem, with the brain as its target organ [26]. tricular hypothalamic nucleus, sub serve spe-
Several physiological studies lend support to cific functions, but other may exert more gen-
this hypothesis. Thus, it has been shown that eralized influences. The same holds true for
in unrestrained behaving cats the activity of the large noradrenergic A5 group. Morpho-
locus coeruleus neurons generally parallels logical and physiological evidence has been
that of the peripheral sympathetic tone presented indicating that this centre projects
[1115], and it has also been demonstrated directly to the thoracic nucleus intermedio-
that these neurons are strongly activitated by lateralis and subserves a vasomotor function
stimuli considered to be significant challenges [806, 808]. However, it has been recently es-
to the integrity of the organism. These find- tablished that the A5 group entertains wide-
ings suggest an important general role of the spread and mostly reciprocal connections
locus coeruleus in the initiation and shaping with virtually all brain regions known to be
of adaptive responses of the organism to involved in cardiovascular control [199]. On
physiological and environmental challenges the basis of this finding it has been suggested
[952, 953, 954, 1110, 1111]. As Aston-Jones that the A5 group" may effect integrated car-
[63] has pointed out, this may mean that an diovascular response patterns by determining
individual under influence of increased locus the global orientation of the cardiovascular
coeruleus activity interrupts automatic, inter- regulatory system", and by doing so would
nally driven or vegetative behaviours (such exert a function analogous to that of the lo-
as sleep or ingestive behaviour), in order to cus coeruleus in the domain of sensory-motor
initiate an adequate response to imperative integration [199]. However this may be, there
(e.g. threatening) external events, and that, can be no doubt that, by way of the large
conversely, internal inhibitory mechanisms longitudinal catecholamine bundle, the nor-
may lead to a decrease of the activity of the adrenergic and adrenergic cell groups play
locus coeruleus, and thus to a favouring of an important role in the gating of the infor-
autonomic preprogrammed behaviour by mation flow through the various functional
allowing external stimuli to be relatively systems contained within the core - lateral
ignored. paracore continuum.
A most remarkable recent finding con-
cerning the connections of the locus coeru-
leus is that, contrary to what has been re-
ported in previous studies (cf Ascending Re-
ticular System, p.216), this centre receives
a substantial input from only two sources,
viz. the nucleus prepositus hypoglossi and the
nucleus paragigantocellularis lateralis ([64];
Fig. 154 B). This finding calls for a detailed
study of the afferents and of the integrative
capacities of these two medullary nuclei.
The noradrenergic A1 and A2 groups and
the adrenergic C1 and C2 groups, which all
form part of the lateral paracore, project to
several basal telencephalic, diencephalic, me-
sencephalic and rhombencephalic core
centres (Fig.-1-59). There is evidence that
some components of the large assembly of
Long Association and Commissural Connections
(Figs. 214-217)
General Features temporal gyrus and the temporal operculum.

These short connections will be reviewed ac-
cording to their modality.
Association and commissural fibres, which 2. Modality-specific areas of parasensory
interconnect different cortical areas, make up association cortex are connected with multi-
most of the white matter of the cerebral modal sensory areas located at their borders.
hemisphere. Commissural fibres cross in the These areas constitute a belt which extends
corpus callosum, the caudal part of the ante- from the junction of the occipital and parietal
rior commissure and the hippocampal com- lobes, through the caudal part of the superior
missures. Association systems can be rather temporal gyrus, into the superior temporal
arbitrarily subdivided into short and long as- sulcus. Long association systems connect the
sociation fibres. Short association fibres may modality-specific parasensory assocIatIOn
remain within the grey matter of the cortex cortex and the multimodal areas in the occi-
or pass through the superficial white matter pital, temporal and parietal lobes with the
between neighbouring cortical areas as U- premo tor and prefrontal cortex of the frontal
fibres. Long association systems are located lobe. Short association fibres interconnect
in deeper parts of the white matter, lateral the prefrontal cortex, the premotor area and
and medial to the corona radiata and the in- the motor cortex. Short association fibres
ternal capsule. In the human brain, long asso- which interconnect the motor cortex and the
ciation systems are mainly known from gross primary somatosensory cortex are relatively
dissection [290, 821], as illustrated in scarce.
Fig. 215. Not much is known about their po- 3. Connections from the parasensory and
larity, origin or termination. The following multimodal association cortices and the pre-
description is mainly based on the experimen- frontal cortex to limbic structures pass
tal literature, which gives a fairly complete through the cingulate gyrus, the hippocampal
account of these connections in primates. gyrus and the insula. The pathways from the
Some general principles apply to the organi- paralimbic association cortex of the cingulate
zation of the corticocortical connections: gyrus and the caudal parahippocampal gyrus
to the hippocampal formation pass through
1. Cascades of short association fibres in- the perirhinal and entorhinal cortices, which
terconnect the primary, modality-specific ar- cover the rostromedial portion of the para-
eas of the cortex, which receive their sensory hippocampal gyrus. The connections of the
input from the thalamus, with the modality- insula are targeted on olfactory limbic areas.
specific para sensory association areas. The 4. Most association connections are recip-
somatosensory association cortex is found in rocal.
the parietal lobe, the visual association areas 5. Connections from the primary sensory
in the occipital lobe and the temporal lobe areas to their neighbouring association areas
below the superior temporal sulcus, and the usually originate from the supragranular
auditory association areas are in the superior layers and terminate in and around layer IV.
2 Forceps minor
3 Genu corporis callosi
5 Rostrum corporis callosi
6 Commissura anterior,
crus anterius
7 Commissura anterior,
crus posterius
11 Tapetum
12 Splenium corporis callosi
13 Radiatio corporis callosi
14 Forceps major
Fig. 214. Commissural connections of the telencephalon as seen from the basal side of the brain
(1 / 1 x)
1 Fasciculus occipitofrontalis superior

2 Site of corona radiata
3 Fasciculus longitudinalis superior
4 Fasciculus longitudinalis superior, brachium posterius
5 Fasciculus longitudinalis superior, brachium anterius
6 Outline of insula
7 Fasciculus occipitofrontalis inferior
8 Fasciculus longitudinalis inferior
9 Site of commissura anterior
10 Fasciculus uncinatus
Fig. 215. Long association bundles of the right cerebral hemisphere in a lateral view (1/1 x). Part
of the superior longitudinal fascicle has been removed to show the superior occipitofrontal fascicle
B Broca's speech area

MT Middle temporal visual association area
W Wernicke's speech area
Fig. 216. Long association connections of the cerebral cortex of the left hemisphere. The multimodal
association cortex in the superior temporal sulcus and the parietooccipital sulcus is stippled
17
p.h. Parahippocampal gyrus

pl.t. Planum temporale
W Wernicke' peech area B
Fig. 217 A, B. Short association connections of the cerebral cortex. A lateral view. The frontal and
part of the parietal operculum has been removed; the temporal operculum is retracted . B Medial
view. The primary and secondary sensory areas (Sl and S2), the primary and secondary motor areas
(Ml and M2), the primary visual and auditory cortices, the paralimbic association cortex and the
insula, and the limbic cortex are indicated with different shadings. The multimodal cortex in the
superior temporal sulcus and the parietooccipital sulcus is stippled
This type of connection is therefore consid- strips, which are separated from each other
ered to transfer information in a forward di- by strips which lack these particular connec-
rection. The reverse, (feed-back) connections tions. This pattern of alternating positive and
originate in the infragranular layers and ter- negative strips has a periodicity of 200-
minate in layer I. The laminar analysis of 1000 !.lm [415, 589, 620, 632]. In the somato-
association connections may therefore reveal sensory and the auditory cortices, the strips
the direction of information transfer. This extend perpendicular to the borders of the
analysis renders it possible to arrange associ- somatotopic or the isofrequency maps, but
ation cortices in logical, hierarchical se- a periodicity in the origin and termination
quences. These sequences have been worked of corticocortical connections has also been
out in some detail for the visual [1136, 1374, observed for visual association areas and for
1398, 1400, 1418] and auditory [375, 1048] the multimodal, frontal and paralimbic asso-
para sensory association areas. For other ciation cortices. Cells of origin and their ho-
parts of the cortex, the data are still far from motopic terminations are located in the same
complete. strips. It is not certain whether the ipsilateral
6. Commissural connections are homotop- association and commissural pathways are
ic or heterotopic. Homotopic connections in- complementary with respect to the strips [32,
terconnect the same cortical areas in both 181,417,628,632,637,1224].
hemispheres. Strictly speaking, homotopic 8. Association and commissural connec-
connections should interconnect bilateral tions are involved in many higher functions
representations derived from the same parts of the nervous system. Damage to the prima-
of the body in the two hemispheres, such as ry sensory or to the parasensory association
the midline of the body or the vertical meridi- areas may result in perceptual deficits. In
an of the visual field. However, the term "ho- contrast, the interruption of commissural
motopic" is generally used for the com- connections or the association connections
missural connections of symmetrical cortical between the unimodal areas and multimodal
areas, which may represent different, but or paralimbic association areas may lead to
symmetrical parts of the body or of the visual disconnection syndromes. These syndromes
or auditory periphery. Commissural connec- have been extensively studied in experimental
tions between cortical areas that represent animals and in humans [899, 1278].
distal parts of the body or the periphery of
the visual or auditory fields are fewer than
those between areas that represent proximal
The Somatosensory Cortex
and axial parts. The density of the com-
missural connections increases from the pri-
mary sensory and motor areas to the more The first somatosensory area (S1) is subdi-
distal association areas [277]. vided into the narrow cytoarchitectonic areas
Heterotopic commissural connections 3B, 1 and 2 [177], which extend parallel to
connect a cortical area with other areas in the central sulcus. Area 3A is usually in-
the contralateral hemisphere. The pattern of cluded in the motor cortex. Areas 3B and
heterotopic connections of a particular corti- 1 contain separate and systematic representa-
cal area often mimics its association connections of the body surface; areas 2 contains
tions with other areas in the ipsilateral hemi- additional representations of predominantly
sphere. It is sometimes possible to distinguish non-cutaneous receptors [651, 652, 1095,
between forward and backward types of 1315]. The lower limb is represented in the
heteropic connections on the basis of their dorsal part, the face in the ventral part, and
laminar origin and termination [277]. the trunk and upper limb in the intermediate
7. Association and commissural connec- part of S1. The second somatosensory area
tions often originate from and terminate in (S2) is located in the parietal operCUlum.
Short, caudally-directed fibres connect of the truncus of the corpus callosum, fibres
area 3B with areas 1 and 2, and area 1 with from Sl are located dorsal to those from S2
area 2 [634, 637, 638, 639, 640, 735, 1040, [1044]. Homotopic connections of the hand
1095, 1451]. Connections to the parasensory and foot representations in area 3B are few;
association cortex in -the caudal part of the there are more for representation of the mid-
superior parietal lobule (area 5) arise from line, trunk, arm and face [277, 632, 686,
the dorsal (hindlimb) part of S1. Those from 1095]. The density of the commissural con-
the more ventral (forelimb, face) parts of Sl nections increases and the sparing of the
terminate in the caudal part of the inferior hand and foot areas decreases in the more
parietal lobule. Brodmann's [177] cytoarchi- caudal somatosensory areas 1 and 2 and in
tectonic subdivision of the posterior parietal the superior and inferior parietal lobules.
cortex into a small dorsally and rostrally lo- Heterotopic commissural connections termi-
cated area 5, a large area 7 occupying most nate in areas which also receive ipsilateral
of the superior parietal lobule, and areas 40 association fibres from the same region. This
and 39 in the inferior parietal lobule remains is also true for the commissural connections
difficult to reconcile with the subdivisions of of the superior and inferior parietal lobules.
the posterior parietal cortex in the monkey These fibres are located in the presplenial
[583, 899]. Cascades of short fibres intercon- part of the corpus callosum and terminate
nect rostral parts of the superior parietal lob- heterotopically in S2, the cingulate gyrus and
ule with the medial parietal cortex of the pre- the superior temporal gyrus and sulcus.
cuneus (area PGM; [1042,1043,1048]). Ros- There are no heterotopic connections be-
tral and caudal (area PG) parts of the inferior tween the parietal cortex and the frontal lobe,
parietal lobule are connected in a similar which receives a prominent ipsilateral projec-
fashion. Connections from the medial parie- tion from the partietal cortex [203, 277, 489,
tal and caudal inferior parietal cortex con- 1043, 1231].
verge with visual and auditory association
fibres in the multimodal sensory cortex locat-
ed at the junction of the parietal and occipital
The Visual Cortex
lobes, in the intraparietal sulcus and the an-
gular gyrus; [1048]).
Rostrally directed pathways interconnect The primary visual area (striate area or V1,
the caudal parietal and S2 cortex with Sl and area 17; [177]) is located in the walls of the
with the primary motor (Ml), premotor, and calcarine sulcus on the medial side of the oc-
supplementary motor (M2) areas of the fron- cipitallobe. It contains a complete represen-
tal lobe [231, 410, 583, 634, 735, 945, 1047, tation of the contralateral visual hemifield,
1303]. Long association fibres from the cau- with the vertical zero meridian located at the
dal parietal cortex, which contribute to the outside border with the visual association
superior longitudinal fascicle, bypass Sl, Ml cortex. It is surrounded by the circumstriate
and the premo tor area and are distributed visual association cortex of area 18 (second-
to the frontal eye field (area 8) and the pre- ary visual area V2) and area 19 (part of it
frontal cortex. The caudal parietal cortex, in- corresponds to the tertiary visual area V3).
cluding the multimodal cortex at the junction Both V2 and V3 contain alternating represen-
of the parietal and occipital lobes, also gives tations of the contralateral hemifield, with
rise to connections with paralimbic associa- the vertical zero meridian located at the inner
tion areas in the cingulate gyrus (area 23), border of V2 with V1 and at the outside
the insula and the parahippocampal gyrus border of V3 and the horizontal meridian at
(area 35; [32, 76, 306, 900, 911, 1224, 1230]). the common border of V2 and V3 [1421].
Commissural connections of the primary Visual association cortex extends beyond the
somatosensory area occupy the middle part circumstriate belt to the temporal lobe, below
the superior temporal sulcus, where it in- Commissural connections of Vl are lim-
cludes areas 20 and 21 [177]. A visual associa- ited to the border region of areas 17 and 18,
tion area is also found in the caudal part along the representation of the vertical zero
of the superior temporal sulcus (the middle meridian. Commissural connections are more
temporal visual area: MT; [1344, 1400, numerous for areas 18 and 19, the inferotem-
1419]). Dorsally, the visual association cortex poral cortex and the multimodal belt. The
extends into the caudal parietal lobule and distribution of the commissural connections
the walls of the intraparietal sulcus. over these areas is uneven; they lie concen-
When the flow of information in the visual trated along the borders but display a patchy
cortex is deduced from the laminar origin distribution within some areas such as 18
and termination of its short corticocortical (V2) and the middle temporal area [276, 277,
connections, two prominent streams can be 680, 994, 1136, 1374, 1421]. Commissural
distinguished [1136, 1374, 1398, 1401, 1418, fibres from areas 18 and 19 occupy the splen-
1421]. One pathway, directed towards the in- ium. Fibres from the inferotemporal cortex
ferior temporal cortex, is mainly concerned cross rostral to the splenium and in the ante-
with the analysis of form and colour and is rior commissure [1044].
crucial for recognition of objects. The other
pathway includes the middle temporal area
and is directed towards the caudal parietal
The Auditory Cortex
lobe. It is involved in visuomotor perfor-
mance, spatial recognition and the analysis
of visual motion. The auditory cortex occupies the superior
Short association fibres interconnect area temporal gyrus and the temporal operculum.
17 (Vl) with area 18 (V2) and the middle The latter consists of Heschl's transverse gy-
temporal area [19, 1344, 1400, 1419, 1420, rus and the temporal plane, which extends,
1493]. A sequence of short connections leads caudal to the transverse gyrus and lateral to
from V2 into V3 (area 19) and the successive- the insula, to the parietal operculum. The
ly more rostral, infratemporal areas 20 and koniocortex of the primary auditory area
21 [1048, 1136, 1137]. V2 and V3, the middle (area 41 and 42; [177]) occupies the trans-
temporal visual area and the inferotemporal verse gyrus. Short association fibres connect
visual cortex project to the adjacent multimo- it with a belt of auditory association areas
dal cortex, which extends from the parieto- surrounding it on all sides [1041, 1229]. Most
occipital junction and the intraparietal of these short connections are directed cau-
sulcus, through the angular gyrus and dally into the temporal plane up to the parie-
Wernicke's area in the caudal superior tem- tal operculum and laterally into the superior
poral gyrus, into the superior temporal sulcus. temporal gyrus (area 22). The next auditory
Long association fibres, which are prob- association area occupies the most caudal
ably located within the posterior branch of part of area 22 (area Tpt of Galaburda and
the superior longitudinal fascicle and the in- Sanides [376] or TAl of Von Economo and
ferior occipitofrontal fascicle, interconnect Koskinas [1455]) in the superior temporal
areas 18 and 19 and the middle temporal area gyrus and in the temporal plane. It corre-
with the frontal eye field (area 8). The cortex sponds to most ifnot all of Wernicke's speech
of the inferotemporal visual areas 20 and 21 area [770]. The auditory association areas
is connected with prefrontal and orbitofron- surrounding the primary auditory cortex and
tal areas that are still more rostral [1136, caudal area 22 are connected, through a se-
1374, 1398]. In addition the visual association ries of short steps, with more rostral parts
cortex is connected with the parahippocam- of the superior temporal gyrus and the cortex
pal area [1048, 1151]. of the temporal pole.
The auditory association cortex in the su- dependent upon the corpus callosum, be-
perior temporal gyrus projects to the multi- cause the auditory pathways which connect
modal cortex in the superior temporal sulcus, the cochlear nuclei with the thalamus are
which also receives visual information from both crossed and uncrossed.
the inferotemporal region and somatosen-
sory projections from the parietal lobe. Part
of this multimodal belt should be included
The Frontal Lobe
in Wernicke's area, which receives both audi-
tory and visual association fibres.
Long association fibres connect Wer- The frontal lobe can be subdivided into: the
nicke's area, through the anterior limb of the primary motor cortex (M1), which corre-
superior longitudinal fascicle (arcuate fasci- sponds to area 4 and is located in the rostral
cle), with the premotor region of the frontal wall of the central sulcus and the caudal part
lobe, including Broca's speech area (area 44) of the precentral gyrus; the premotor area,
in the inferior frontal gyrus and area 8 [231, which includes area 6, area 8 (the frontal eye
375, 640, 770, 1048]. The fibres of the anteri- field) and Broca's speech area (area 44) in
or limb are located beneath the supramargin- the opercular part of the inferior frontal gy-
al gyrus. More rostral parts of the superior rus (for other definitions of the premotor
temporal gyrus are connected with succes- area see [945]); and the prefrontal cortex,
sively more rostral parts of the prefrontal which includes the orbitofrontal cortex. The
cortex. The temporal pole is connected with supplementary motor area (M2; [1065]) be-
the mediofrontal and orbitofrontal cortex longs to area 6 and is located on the medial
through the uncinate fascicle. The auditory aspect of the superior frontal gyrus, in front
association areas, especially the more rostral of the paracentral lobule.
ones, also establish connections with the Short association fibres successively link
paralimbic association cortex of the cingulate the orbitofrontal cortex, the prefrontal areas
and parahippocampal gyri. 9, 10 and 46, the premo tor area 8 and area
A strong asymmetry has been described 6. Dorsal and ventral streams, passing
for the human auditory cortex. The asym- through the dorsal and ventral parts of areas
metry mainly concerns the size of the tempo- 46 and 8 and directed towards the dorsal and
ral plane, which is usually larger in the left, ventral parts of area 6, have been distin-
dominant hemisphere [377, 396]. Wernicke's guished in the monkey [78, 607, 608, 945,
area (area Tpt [376]) displays the most asym- 1048]. Connections with the motor cortex
metry, but other auditory areas are also in- arise from area 6, including the supplementa-
volved [770]. ry motor cortex, but not from area 8 or more
Commissural connections of the auditory rostral parts of the frontal lobe [79, 410, 735,
cortex are predominantly homotopic. They 736, 750, 775, 1040, 1047, 1303].
are located in the caudal part of the truncus Ventral area 6 is topically and reciprocally
of the corpus callosum, together with the connected with the face and hand representa-
commissural fibres of the parietal lobe [1044]. tions in the primary motor cortex, although
In the cat they interconnect columns of bila- the premotor-motor pathway to the mouth
terally excited neurons, which extend perpen- representation in ventral area 4 is not recip-
dicular to the borders of the isofrequency rocal and contains an extra link in rostral
maps in the auditory cortex [589, 590]. Inter- area 6 [945]. Premotor connections to the
mediate columns, which contain contralater- foot representation in the dorsal motor cor-
ally excited neurons, do not receive callosal tex originate from the caudal part of dorsal
afferents. The bilateral representation of the area 6 [79, 410]. The supplementary motor
auditory field in the auditory cortex is not area (M2) is connected with the entire prima-
ry motor cortex and with the dorsal and ven- the multimodal sulcal cortex project more
tral parts of area 6 [647, 735]. The short con- rostrally into the prefrontal cortex through
nections of Broca's speech area (area 44) are the inferior occipitofrontal bundle. The cor-
unknown. tex of the temporal pole is connected with
The primary motor cortex is thus topically the orbitofrontal cortex through the uncinate
connected with premo tor area 6 and somato- fascicle. Convergence of (parietal) somato-
sensory area 2. In addition Ml is linked with sensory and (temporal) auditory projections
the supplementary motor and sensory cor- occurs in the dorsal part of area 8 and area
tices. Area 3A is located in the base of the 46; somatosensory and visual projections
central sulcus. Its cytoarchitecture is similar converge more ventrally, in areas 46 and 45.
to area 4, but it receives information from Somatosensory projections from the caudal
muscle receptors through the thalamus. Its parietal cortex join auditory and visual pro-
short association connections are the same jections from the temporal lobe in the more
as those of the primary motor cortex [634, rostral parts of the prefrontal cortex.
637,638,639, 1095]. The prefrontal cortex maintains strong
Long association fibres interconnect the connections with limbic and paralimbic asso-
premotor and the prefrontal cortices with the ciation areas. It projects through the superior
postcentral association cortex [231, 410, 415, occipitofrontal bundle, which is located ven-
607, 736, 1047, 1048, 1079]; see also previous tral to the radiation of the corpus callosum,
paragraphs). The somatosensory association and the cingulum, which is located dorsal to
cortex of the superior parietal lobule and the it, to the cingulate gyrus, the retrosplenial
rostral part of the inferior parietal lobule are cortex, the parahippocampal area and the
connected with dorsal and ventral parts of presubiculum [414, 417, 418, 607,1047]. Con-
area 6 and with the supplementary motor nections with the cingulate gyrus and also
cortex. The caudal parietal somatosensory the retrosplenial cortex have been described
association cortex and the multimodal (so- for the premotor area (area 8 and ventral
matosensory and visual) cortex, located at area 6) and for ventral area 6 connections
the parieto-occipital junction and in the an- with the insula have been reported [607, 945,
gular gyrus (area 39), project more rostrally 1047].
into the frontal lobe to areas 8 and 46, but The commissural fibres between the fron-
not to area 6 or the supplementary motor tal lobes occupy the rostrum and genu of the
cortex. corpus callosum. Fibres of the premo tor area
Similar patterns of connections are found are located next to those from M2 in the ros-
between caudal and rostral parts of the occi- tral part of the truncus. Fibres from the pre-
pitotemporal association cortex and the cau- central region occupy the middle part of the
dal (premotor) and rostral (prefrontal) cor- truncus of the corpus callosum [1044]. Ho-
tex. The auditory association cortex in caudal motopic connections interconnect all parts of
area 22 projects to the premotor area. This the frontal lobe, except for the hand and foot
connection, through the anterior limb of the areas of the motor cortex, which give rise
superior longitudinal fascicle, probably in- to relatively few commissural fibres [277, 318,
cludes the pathway between the speech areas 1047]. Heterotopic projections from the mo-
of Wernicke in the superior temporal gyrus tor cortex connect it with the supplementary
and Broca in the frontal operculum [770]. motor cortex and area 6. The premo tor and
Caudal visual areas (18 and 19 and the mid- supplementary motor areas are connected
dle temporal visual area) are reciprocally with each other, with the parietal association
connected with the frontal eye field (area 8). cortex, the cingulate gyrus and the cortex of
Rostral parts of the auditory association cor- the superior temporal sulcus [647, 735]. Het-
tex in area 22, the visual fields below the su- erotopic connections of the prefrontal cortex
perior temporal sulcus (areas 20 and 21) and link it with other parts of the prefrontal cor-
tex, the visual and auditory association cor- poral pole, the inferotemporal cortex and the
tex (areas 21 and 22) in the temporal lobe insula [1048, 1228, 1426]. Moreover, prefron-
and the cingulate gyrus [608, 1224]. tal and premotor areas are connected with
the parahippocampal area through the ros-
tral part of the cingulate gyrus (areas 32 and
24; [177]) and the cingulum. The caudal part
The Paralimbic Areas of the cingulate gyrus (area 23) serves as the
relay for similar, indirect connections to the
parahippocampal area from the para sensory
The neocortex has access to limbic structures and multimodal association areas (visual
through the paralimbic association cortex of area 19, the caudal parietal somatosensory
the cingulate gyrus, the parahippocampal association cortex and the auditory and visu-
area and the insula. The caudal part of the al fields in the temporal lobe, including the
cingulate gyrus (area 23) and the rostrome- multimodal cortex in the superior temporal
dial, perirhinal (area 35; [177]) and entorhi- sulcus; [74, 1046, 1048, 1450]). Connections
nal (area 28) parahippocampal cortices main- from the insula are directed towards the
tain direct connections with the hippocampal olfactory limbic areas [903].
formation. Connections from the neocortex Commissural connections of the cingulate
to the hippocampus travel through the super- gyrus and the insula are distributed widely
ficial layers of the perirhinal and entorhinal over the truncus of the corpus callosum
cortices; the reciprocal connections from the [1044]. Heterotopic projections connect the
hippocampus and the subiculum to the neo- cingulate gyrus with areas 17 and 18 [1450].
cortex synapse in their deep layers [1521]. Fibres from the parahippocampal area cross
The parahippocampal gyrus, including the in the splenium, the anterior commissure
perirhinal area 35, receives direct connections and, from the entorhinal and perirhinal cor-
from the prefrontal cortex and the parasen- tices, in the hippocampal commissures [645,
sory association areas, but also from the tem- 646, 1044].
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Subject Index
In this index the first number (in Sans Serifs) refers to the figure; the second number corresponds to
the number under which the pertinent structure can be found in the figure. Numbers following the abbrevia-
tion "p." refer to text-pages.
accessory optic system 145; p. 184 anteromedian medullary vein, see: vena spinalis
accomodation reflex p. 185 anterior
adenohypophysis, see: hypophysis, lobus anterior apertura lateralis ventriculi quarti, syn,' foramen of
adhesio interthalamica 13.10 Luschka 15.41
adrenergic cell groups p. 205, 290 apertura mediana ventriculi quarti, syn,' foramen of
- ascending projections 159; p. 220 Magendie 12.29; 15.45
A-fibre(s) of dorsal root 123.4; 129.A; p. 150 aqueductus cerebri 3.4; 12.22; 21.12; 74.24
agonistic behaviour p. 293, 349, 362 arachnoidea (mater) 50 (legend); 52.5+12;
ala cisternae ambientis, syn,' wing of the ambient 53.7; 58.18+38
cistern; retropulvinar cistern 55.g archipallium p. 334
ala lobuli centralis 18.16; 75.15 arcuate fascicle, see: fasciculus longitudinalis
ala major ossis sphenoidalis 41.19 superior
ala minor ossis sphenoidalis 38.9 area
allocortex p. 334 - lateralis hypothalami 91.6; 93.9; 192.3;
alveus hippocampi 94.33 197.3; 1988.10; p.302, 319
Alzheimer's diseae p. 276 - ovalis, see: nucleus ovalis
amiculum (olivae) 107.25; 163.16 - postrema p. 164, 310
amygdala, see: corpus amygdaloideum - pretectalis, syn,' pre tectum 86.30; 97.3;
amygdala, afferents p. 324 146.10; 147.9; 149.4; p. 189
amygdala, efferents p.328 - reticularis parvocellularis p. 202
analgesia p. 160 - reticularis parvocellularis, connections p.211,
anastomoses 284
- cerebrocerebral, see: anastomoses, encephalo- - reticularis superficialis ventrolateralis p. 202,
encephalic 212
- cerebromeningeal 30.1 - reticularis superficialis ventrolateralis, connec-
- encephaloencephalic (anastomoses of brain and tions p. 211, 212, 286
of brain stem arteries) 31.5+28; 36; 48.6 - striata, syn,' primary visual cortex (area 17)
- extracranio-orbital 30.9 + 22 + 26 72.15; 75.10; 144.21; 145.17; 147.13;
- extracraniomeningeal 30.38 + 40 p. 181, 244, 371
- meningeo-orbital and orbitomeningeal 30.14+ - subcallosa 11.7; 89.12; 193.10; 203.5
19; 43.10 - subcoerulea, see: nucleus subcoeruleus
angulus oculi lateralis, see: ektokanthion - tegmentalis, syn,' tegmentum 80.34
ansa anastomotica 56.16 - tegmentalis dorsalis p. 323
ansa lenticularis 92.36; 171.13; 173.14; p.261 - tegmentalis H, syn,' field H of Forel 93.27
ansa peduncularis 91.32; 169.15; 191.16 - tegmentalis pontis lateralis p. 286
anterior, central and posterior insular veins, see: - tegmentalis ventralis (Tsai) 97.16; 192.5;
venae insulares 198A.7; p.302, 320
anterior cerebellar vein, see: vena pontis lateralis - vestibularis 15.13
anterior cerebral veins, see: venae cerebri anter- area 4, see: motor cortex, primary
iores area 6, see: premotor area
anterior extravertebral veins, see: plexus venosi area 8, see: frontal eye field
vertebrales externi area 17, see: area striata
anterior interposed nucleus, see: nucleus emboli- area 18 + area 19, see: visual cortex, secondary
formis area 20 5.20; 217.20
anterior, peripeduncular (paired) part of the ambi- area 21 5.21; 217.21
ent cistern, see: cisterna cruralis area 44, see: Broca's speech area
anterolateral bundle of Gowers, see: fasciculus arteria
anterolateralis - Adamkiewiczi, see: arteria radicularis magna
anterolateral system p. 151 - angularis 30.27
420 Subject Index
arteria arteria
- basilaris 38.17; 40.34; 43.37 - cerebri pars rami thalamici, syn: rami diencepha-
- callosomarginalis, syn: arteria cingulomarginalis; lici (posteriores + superiores) 31.24 + 27;
ramus cingularis 31.11; 39.3 35.33 + 34; 40.21
- - ramus frontalis anteromedialis, syn: arteria - - - ramus choroideus posterior lateralis, syn:
frontalis interna anterior 31.10; 32.4 rami choroidei posteriores laterales; arteria
- - ramus frontalis intermediomedialis, syn: arter- choroidea posterior lateralis 35.30; 40.20;
ia frontalis interna intermedia 31.9; 32.3 51.18
- - ramus frontalis posteromedialis, syn: arteria - - pars precommunicalis 31.20; 35.24
frontalis interna posterior 31.8; 32.2 - - - rami thalamici, syn: rami diencephalici
- candelabra, see: arteria prefrontalis (inferiores) 35.32
- carotis communis 30.49; 38.43; 39.43 - - - ramus choroideus posterior medialis, syn:
- carotis externa 30.47; 38.28; 39.41 rami choroidei posteriores mediales; arteria
- carotis interna 30.48; 41.28 choroidea posterior medialis 35.27; 40.18;
- - pars cavernosa 38.40; 39.21 51.17
- - pars cerebralis, syn: pars subarachnoidealis; - - pars terminalis (corticalis), see: arteria occipi-
pars supracavernosa 37.16; 39.20; 43.6 talis lateralis/medialis
- - pars cervicalis 38.42; 39.38 - cervicalis ascendens 56.8
- - pars petrosa, syn: pars canalis carotici - cervicalis profunda 56.9
38.41; 39.27 - choroidea anterior 35.28; 37.25; 40.14;
- centralis, see: arteria sulci centralis 51.16
- centralis longa, syn: arteria recurrens; arteria - - rami capsulae internae 40.17
Heubneri 37.9 - - rami choroidei (ventriculi lateralis) 35.29
- cerebelli inferior anterior 39.30; 40.38; 43.40 - choroidea posterior lateralis, see: arteria cerebri
- cerebelli inferior posterior 38.21; 40.45; posterior, pars postcommunicalis, ramus choroi-
46.35 deus posterior lateralis
- - rami medullares (mediales, laterales et poster- - choroidea posterior medialis, see: arteria cerebri
iores), syn: rami ad medullam oblongatam; posterior, pars precommunicalis, ramus choroi-
(laterales:) rami retroolivares 49.4+ 5+ 10 deus posterior medialis
- - ramus choroideus ventriculi quarti 46.33; - cingulomarginalis, see: arteria callosomarginalis
51.27 - cingulothalamica 31.26; 40.24
- - ramus lateralis (rami laterales), syn: arteria - - ramus corporis callosi dorsalis 31.28; 40.26
hemispherii cerebelli inferior lateralis 34.28; - - ramus thalamicus (superior) 31.27; 40.25
38.20; 40.46 - collicularis, see: arteria quadrigemina
- - ramus medialis, syn: rami mediales; arteria - communicans anterior 31.15; 38.37; 43.4
hemispherii cerebelli inferior medialis 34.29; - communicans posterior 31.19; 35.21; 40.12;
38.19; 46.28 43.13
- - ramus tonsillae cerebelli 46.32 - - ramus hypothalamicus, syn: arteria tuberotha-
- cerebelli superior 38.16; 39.17 lamica; arteria premammilaris 35.22; 40.13
- - ramus lateralis 38.14; 40.30 - - ramus thalamicus 35.23; 40.15
- - ramus medialis 38.7; 40.29 - dorsalis nasi 30.9
- - ramus mesencephalicus, syn: ramus laminae - ethmoidalis anterior 30.14; 43.32
tecti; ramus quadrigeminus; ramus colliculi in- - ethmoidalis posterior 30.16; 43.32
ferioris 40.32 - facialis, syn: arteria maxillaris externa 30.28
- cerebri anterior 37.11; 39.7; 43.5 - frontalis ascendens 33.10
- - pars ascendens 31 .14 - frontalis interna, see: arteria callosomarginalis
- - pars postcommunicalis 31.14; 38.36 (rami)
- - pars precommunicalis 35.20; 38.38 - frontalis medialis, see: arteria supratrochlearis
- cerebri media 43.11 - frontobasalis lateralis, syn: ramus orbitofrontalis
- - pars insularis 30.34; 35.13; 38.35 lateralis 33.9; 35.12
- - pars sphenoidalis 35.17; 38.39; 40.9 - frontobasalis medialis, syn: ramus orbitofrontalis
- - pars terminalis 38.34 medialis (arteria orbitalis + arteria frontopolaris)
- - rami striati 64.33 31.16; 35.11
- - truncus anterior, syn: frontal trunk 33.10; - gyri angularis, syn: artere parieto-oocipitale
39.8 33.22
- - truncus intermedius, syn: parietal trunk - hemispherii cere belli inferior lateralis, see: arteria
33.13; 39.10 cerebelli inferior posterior, ramus lateralis (rami
- - truncus posterior, syn: temporal trunk laterales)
33.14; 39.11 - hemispherii cere belli inferior medialis, see: arter-
- cerebri posterior 38.15; 39.18 ia cerebelli inferior posterior, ramus medialis
- - pars postcommunicalis 31.23; 35.26 - Heubneri, see: arteria centralis longa
- - - rami pedunculares 47.20 - hypophysialis inferior 37.13; 192.24+25
Subject Index 421
arteria arteria
- hypophysialis superior 37.12; 192.16 - recurrens, see: arteria centralis longa
- infraorbitalis 30.24 - sphenopalatina 30.23
- intercostalis suprema 56.12 - spinalis anterior 43.42; 56.2; 58.29
- intercosticalis posterior 56.1jXI; 58.6 - spinalis posterior (dextra et sinistra) 56.3;
- - ramus spinalis 58.31 58.13
- labyrinthi, syn: ramus meatus acustici interni - splenialis, see: arteria pericallosa, ramus
40.40 posterior
- lacrimalis 30.12; 43.33 - subcostalis 56.17
- laminae tecti, see: arteria quadrigemina - sulci centralis, syn: arteria centralis; Rolandic
- lumbalis 56.18 artery 33.2
- magna spinalis, see: arteria radicularis magna - sulci postcentralis, syn: arteria postcentralis;
- maxillaris, syn: arteria maxillaris interna arteria parietalis (anterior) 33.20
30.29; 38.23 + 26; 39.39 - sulci precentralis, syn: arteria precentralis;
- maxillaris externa, see: arteria facialis prerolandic artery 33.5
- maxillaris interna, see: arteria maxillaris - supratrochlearis, syn: arteria frontalis medialis
- mediana corporis callosi 31.13; 46.47 30.8
- meningea anterior 30.11; 43.1 - temporalis anterior 33.11; 35.3
- meningea media 30.30; 38.24 - temporalis intermedia 33.15; 35.4
- - ramus anastomoticus 30.19; 43.10 - temporalis posterior 33.16; 35.5
- - ramus frontalis 30.20; 38.11; 43.15 - temporalis superficialis 30.45
- - ramus parietalis 30.33; 38.12; 43.17 - temporo-occipitalis 33.18
- meningea posterior 30.42; 43.28 - temporopolaris 33.12; 34.21
- occipitalis 30.39 - tuberothalamica, see: arteria communicans
- occipitalis lateralis, syn: arteria cerebri posterior, posterior, ramus hypothalamicus
pars terminalis (corticalis) 31.35; 35.8; 38.8 - vermis inferior 46.27
- - rami temporales anteriores 31.37; 35.2 - vermis superior, syn: ramus vermis superior
- - rami temporales intermedii mediales 31.34; 40.28
35.6 - vertebralis 30.50
- - rami temporales posteriores 31.33; 35.9 - - pars atlantis, syn: pars atlantica 38.27;
- occipitalis medialis, syn: arteria cerebri posterior, 39.37
pars terminalis (corticalis) 31.25; 35.7; 38.6; - - pars intracranialis 38.22; 39.36; 56.1
40.23 - - - rami medullares (anterolaterales + antero-
- - ramus calcarinus 31.32 mediales) 49.11 + 14
- - ramus parietalis 31.29 - - - ramus meningeus posterior 43.24
- - ramus parieto-occipitalis 31.31; 38.2 - - pars transversaria, syn: pars cervicalis 38.30;
- operculofrontalis, see: arteria prefrontalis 39.42; 56.5
- ophthalmica 30.17; 41.15; 43.35 - - - rami spinales, syn: rami radiculares 56.4
- paracentralis 31.6; 38.31 arteriae
- parietalis (anterior), see: arteria sulci - centrales anterolaterales, syn: arteriae thalamo-
postcentralis striatae anterolaterales; rami diencephalici
- parietalis interna, see: arteria precunealis lateroinferiores
- parietalis posterior 33.21 - - rami laterales 35.15; 37.7; 40.7
- parieto-occipitalis, see: arteria gyri angularis - - rami mediales 35.16; 37.8; 40.8
- pericallosa 31.12; 39.2 - centrales anteromediales, syn: arteriae thalamo-
- pericallosa, ramus posterior, syn: ramus corporis striatae anteromediales; rami diencephalici
callosi posterior; arteria splenialis 31.5 anteroinferiores 35.18
- pharyngea ascendens 30.46 - centrales posteromediales, syn: rami diencephalici
- postcentralis, see: arteria sulci postcentralis posteroinferiores; rami interpedunculares profun-
- precentralis, see: arteria sulci precentralis di 35.25; 40.16; 46.42; 47.11
- precunealis, syn: rami precuneales; arteria - insulares 7
parietalis interna (inferior + superior) 31.4 - mesencephalicae, see: arteria quadrigemina
- prefrontalis, syn: arteria operculofrontalis; - pontis, syn: rami ad pontem
arteria candelabra 33.8 - - rami laterales, syn: superolateral + inferolateral
- premamillaris, see: arteria communicans pontine arteries 40.36; 43.39; 48.9+10
posterior, ramus hypothalamicus - - rami mediales 40.35; 48.13
- quadrigemina, syn: arteria laminae tecti; arteria - thalamostriatae anterolaterales, see: arteriae cen-
collicularis 47.1 + 19 trales anterolaterales
- radicularis anterior (arteriae vertebralis) 56.7 - thalamostriatae anteromediales, see: arteriae cen-
- radicularis magna, syn: arteria magna spinalis; trales anteromediales
arteria Adamkiewiczi 56.15; 58.28 ascending reticular pathways p. 206
- radicularis posterior (arteriae vertebralis) 56.6 ascending reticular systems p. 197
422 Subject Index
ascending sacral vein, see: ramus dorsalis venae capsula interna 28; 81.23; 172.4
sacralis lateralis - crus anterius 28.1; 85.5; 90.20; 169.11 + 12
"aspecific" thalamocortical projection p. 210, - crus posterius 28.10; 85.9; 92.28; 169.6/10;
238 173.5
association connections p. 365 - genu 85.8
association fibres 215; 216; 217; p.365 - pars retrolentiformis 28.9; 86.11 ; 95.27;
atrial part of the vena terminalis (inferior), see: 169.17; 172.11
vena atrii lateralis - pars sublentiformis 28.8; 67.31; 94.28;
auditory cortex p. 372 169.16; 173.17
auditory projection, ascending p. 177 caput costae: facies articularis capitis costae 58.2
auditory projection, descending p. 178 caput nuclei caudati 28.4; 29.3; 63.16; 80.29
auditory reflex pathways p. 179 cardiorespiratory function p. 218
auditory system p. 172 carotissiphon, see: siphon arteriae carotidis
internae
Babinski's sign p. 258 cauda equina 19
bandeletta diagonalis (Broca) 89.25; 90.30; cauda nuclei caudati 29.9; 68.33; 85.30; 95.3
196.8 cavitas epiduralis (spinal epidural space) 58.21 +
basal amygdaloid nucleus, see: corpus amygdaloi- 40; 60.1
deum, nucleus basalis cavitas subarachnoidealis 50.3; 53.8; 58.18; 60.2
basal ganglia 27; 28; p. 269 cavum septi pellucidi 91.1
basal olfactory area p.344 cavum trigeminale (Meckeli) 41.25; 43.36
basal pontine nuclei, see: nuclei pontis cell group Y p. 233
basis cranii 43 cell group Z p.154
bed nucleus of the stria terminalis, see: nucleus cellulae marginales (cornus posterioris) 117.1;
interstitialis striae terminalis 121.21; p. 150
bicommissural line (Talairach) 39.BC cellulae motoriae cornus anterioris 174.25
bordercell 123.10; 161.24 cellulae motoriae laterales (comus anterioris)
brachial vein, see: vena pedunculi cerebellaris 117.6; 121.27; 124.13
superioris cellulae motoriae mediales (cornus anterioris)
brachium colliculi inferioris 15.6; 99.15; 117.7; 121.29; 124.14
100.13; 141A.4; 142.3 central grey matter, see: substantia grisea centralis
brachium colliculi superioris 15.4; 86.13; central limbic continuum p. 296, 302, 309
97.19; 144.13; 145.13 centre median (Luys), see: nucleus centromedianus
brachium conjunctivum, see: pedunculus cerebel- centroposterior orbitofrontal cortex p. 345
laris superior centrum ciliospinale 146.15
brachium pontis, see: pedunculus cerebellaris cerebellar ataxia p. 236
medius cerebellar nuclei, see: nuclei centrales cere belli
Broca, diagonal band of -, see: Bandeletta cerebellar serotoninergic pathway p. 224
diagonalis cerebellopontine angle cistern, see: cisterna ponto-
Broca's speech area p. 373 cerebellaris
Brodmann, cytoarchitectonic fields of - 5 cerebellovestibular fibres 166; p.232
bulbospinal serotoninergic projections 183; cerebellum 2.2; 16; 18; 160-166; p.221
p.281 - corticonuclear projection p. 228
bulbus oculi 144.1 - function and malfunction p. 236
bulbus olfactorius 9.8; 196.1 207.9; p.344 - zonal organization p. 228
bulbus venae jugularis inferior 57.10 cerebrum, syn: telencephalon 1.1 ; 2.1; 4
bulbus venae jugularis superior 41.29 C-fibre(s) of dorsal root 123.5; 129.C
bundle of Rasmussen, see: fasciculus olivocochlearis chiasma opticum 9.2; 13.18; 64.36; 144.3;
bundle of Vicq d'Azyr, see: tractus mamillo- 145.7
thalamicus cholinergic cell groups 154C; 180; p.205, 273
cholinergic fibers 180; p. 205, 273
calvaria 30.2; 38.1; 53.2+3+4 cingulum 92.22; 191.1; p.335
canales semicirculares 140.19+20+21 circuit of Papez 206; p.303, 341
canalis caroticus 38.41; 39.27; 51.32 circulus arteriosus cerebri (Willisii) 43.4 + 5 + 11 +
canalis centralis 3.6; 12.30; 21.16; 121.10 13+14
canalis hypoglossalis 43.41 circumventricular organs p. 309
canalis infraorbitalis 30.25 cistern of the Gasserian ganglion, see: cisterna
canalis opticus 30.21 trigemini
canthus externus, see: ektokanthion cistern of the roof of the third ventricle, see:
canthus-meatus line 39.CM cisterna fissurae transversae
capsula externa 67.22; 92.29 cistern of the Sylvian fissure, see: cisterna fossae
capsula extrema 67.20; 92.30 lateralis cerebri
Subject Index 423
cistern of the vein of Galen, see: cisterna venae claustrum 67.21; 85.19; 91.4
cerebri magnae climbing fibres p. 222, 229
cisterna coeruleospinal system p. 256
- ambiens = cisterna cruralis + cisterna laminae collateral circulation, arterial 30
tecti 54.1; 55.f + g + p collateral circulation, venous 42
- cerebelli superior 52.27; 54.k colliculo-cochleonuclear projection p. 179
- cerebellomedullaris, syn: cisterna magna 32.27; colliculus facialis 15.12
34.17; 52.36 54.q colliculus inferior 14.5; 15.7; 141A.5
- chiasmatis 34.4; 52.18; 54.g - nucleus centralis 101.2; 142.11
- corporis callosi, see: cisterna pericallosa - zona lateralis 101.3; 142.9
- cruralis, syn: anterior, peripeduncular (paired) colliculus superior 14.3; 15.5; 145.21; 149.8;
part of the ambient cistern 55.f p. 194
- fissurae transversae, syn: cisterna veli interpositi; - stratum album profundum 98.19
cistern of the roof of the third ventricle 52.24; - stratum griseum medium 98.3
54.i; 55.q - stratum griseum profundum 98.4
- fossae lateralis cerebri, syn: cisterna fossae Syl- - stratum griseum superficiale 98.2
vii; cistern of the Sylvian fissure 32.20; 54.e; - stratum lemnisci 98.18
55.r - stratum opticum 98.17
- fossae Sylvii, see: cisterna fossae lateralis cerebri - stratum zonale 98.1 + 16
- intercruralis, see: cisterna interpeduncularis columna Clarki (Stilling-Clarke), see: nucleus
- interpeduncularis, syn: cisterna intercruralis thoracicus
34.8; 52.19; 54.h; 55.d columna fornicis 22.7; 79.29; 90.26; 92.35;
- laminae tecti, syn: cisterna quadrigemina; poste- 195.6
rior, unpaired part of the ambient cistern; name commissura
also used including the cistern of the vein of - alba (medullae spinalis) 116.14; 120.14;
Galen (cisterna venae cerebri magnae) 52.25; 121.12; p.151
55.p - anterior 13.11; 27.10; 86.3; 90.27; 92.38
- laminae terminalis 34.1; 52.11; 54.d; 55.b - - crus anterius 214.6
- magna, see: cisterna cerebellomedullaris - - crus posterius 214.7
- meatus acustici (interna) 34.13; 55.i - cerebelli 103.16
- medullae oblongatae et spinalis, see: cisterna - colliculi inferioris 141A.6; 142.10
medullaris - colliculi superioris 98.20
- medullaris, syn: cisterna medullae oblongatae et - fornicis 22.10; 194.3; 195.20
spinalis 32.25; 34.16; 52.22; 54.r - grisea anterior 120.7; 121.11
- pericallosa, syn: cisterna corporis callosi 52.9; - grisea posterior 120.6; 121.9
54.c - habenularum 13.5; 194.10; 196.22
- pericallosa posterior, see: cisterna retrosplenialis - posterior 13.8; 68.29; 97.21; 146.11; 147.10
- pontis (pars medialis et pars lateralis) 32.22; - supramamillaris 97.36
34.11; 52.20; 54.m commissural connections p.370
- pontocerebellaris, syn: cerebellopontine angle commissural fibres 214
cistern; "Kleinhirnbruckenwinkelzisterne" commissure of the nuclei of the lateral lemniscus,
32.24; 34.15; 54.p; 55.j see: decussatio lemniscorum lateralium
- quadrigemina, see: cisterna laminae tecti confluens sinuum 42.20; 43.31; 52.29
- retrosplenialis, syn: cisterna pericallosa posteri- conus medullaris 20
or = posterior part of the pericallosal cistern core of the neuraxis p. 350
55.n - caudal part p. 352
- suprasellaris, see: cisternae suprasellares - rostral part p. 356
- trigemini, syn: cistern of the Gasserian ganglion cornu ammonis 23.12+22; 74.31; 93.16;
34.12; 54.n; 55.h 195.12+ 14+ 15; 203.10; p.334
- valleculae cerebri (" root" of the cistern of the cornu anterius, syn: ventral horn 121.8
Sylvian fissure) 34.3; 55.c cornu posterius, syn: dorsal horn 121.5; p. 150,
- veli interpositi, see: cisterna fissurae transversae 160
- venae cerebri magnae, syn: cistern of the vein of corona radiata 66.20; 82.23; 172.2; 173.1
Galen; may be considered to constitute one corpus
cistern together with the quadrigeminal cistern - amygdaloideum, syn: amygdala 74.32; p. 323
52.26; 54.j; 55.0 - - afferents p.324
cisternae subarachnoideales 32; 34; 52; 54; - - efferents p. 328
55 - - functions p. 329
cisternae suprasellares = cisterna chiasmatis + cister- - - nucleus anterior, syn: nucleus anterior amygda-
na interpeduncularis + cisterna laminae termina- lae 90.12; 193.22; 196.13
lis 34.1+4+8; 52.11+18+19; 54.d+g+h - - nucleus basalis, syn: nucleus basalis amygda-
Clarke's thoracic nucleus, see: nucleus thoracicus lae 91.15; 196.17; 199.17; 201.18
424 Subject Index
corpus costa 58.1

- amygdaloideum nucleus basalis accessorius, syn: cranial nerve nuclei p. 144
nucleus basalis accessorius amygdalae 91.14; crista galli 52.13
93.14; 196.17 crus cerebri, see: pedunculus cerebri
- - nucleus centralis, syn: nucleus centralis amyg- crus fornicis 22.12; 69.21; 95.25; 195.19
dalae 92.15; 196.15; 199.16; 201.16 culmen 13.28; 16.1; 75.12; 162
- - nucleus corticalis, syn: nucleus corticalis amyg- cuneus 11.27; 75.3
dalae 91.13; 193.21; 196.12; 199.15;
201.17 declive 13.30; 16.2; 162
- - nucleus lateralis, syn: nucleus lateralis amygda- decussatio
lae 91.16; 196.14; 199.17; 201.18 - fasciculorum uncinatorum cerebelli 103.17
- - nucleus medialis, syn: nucleus medialis amyg- - lemniscorum lateralium, syn: Probst's com-
dalae 91.12; 196.16; 199.15; 201.17 missure 142.14
- callosum 12; 23.3; 214.3+5+9+12 - nervorum trochlearium 102.12
- cerebelli 18(8); 162 - pedunculorum cerebellarium superiorum
- ciliare 147.2 79.32; 95.35; 100.24; 160.3; 163.6
- fornicis 22.9; 67.14; 91.20; 195.16 - pyramidum 17.37; 109.16; 170.20; 172.18;
- geniculatum laterale 14.4; 15.33; 68.32; p.247
145.16; 167.16; p.241 - tegmentalis dorsalis (Meynert) 99.24
- - laminae magnocellulares 97.6 - tegmentalis ventralis (Forel) 99.27
- - laminae parvocellulares 97.7 - tractuum trigeminothalamicorum ventralium
- geniculatum mediale 15.32; 68.31; 141A.3; 103.31
167.17; p.154, 241 deep middle cerebral vein, see: vena cerebri media
- - pars dorsalis 97.4; 142.4 profunda
- - pars medialis 142.6 Deiters' nucleus, see: nucleus vestibularis lateralis
- - pars ventralis 97.5; 142.5 descending reticular systems p. 279
- mamillare 9.6; 13.15; 192.11; 194.20; Deutsche Horizontale, see: horizontal line or plane
205.13; p.303 of Frankfurt (Reid)
- - nucleus lateralis 97.17; 156.14 diaphragma sellae 52.16 + 17
- - nucleus medialis 97.18 diencephalon 2.3
- medullare cerebelli 71.24; 82.17 digitationes hippocampi 23.13
- nuclei caudati 28.5; 29.1; 65.17; 84.18; diploe 53.3
92.2 direct fastigiobulbar tract, see: tractus fastigiobul-
- parabigeminum 101.8 baris
- pineale 13.6; 15.3; 85.34 dopaminergic cell groups (and pathways) p.260
- pontobulbare 105.15; 106.11 dorsal ascending serotoninergic pathway p. 207
- restiforme, see: pedunculus cerebellaris inferior dorsal column nuclei p. 154, 160
- striatum 26.1 + 2 dorsal cuneocerebellar tracts, see: tractus cuneo-
- subthalamicum (Luys), see: nucleus subthalami- cerebellaris
cus dorsal diencephalic conduction system p. 208
- trapezoideum 104.28+32; 105.25; 141A.11; dorsal horn, see: cornu posterius
142.20 - laminar arrangement 122
- vertebrae 58.49 dorsal noradrenergic bundle 158; p.213, 290
cortex - caudal limb 188 p. 290
- cere belli p. 222, 225 dorsal pathway of Shute and Lewis p. 205
- cerebri, columnar organization (positive and dorsal periventricular pathway 158; p.213,
negative strips) p. 370 218, 290
- - cortical layers I-VI p.238 dorsal raphespinal projection p.281
- entorhinalis 93.21; 204.14; 206.11; p.335 dorsal root fibres, syn: fibrae radicis dorsalis nervi
- frontalis medialis 205.7 spinalis p. 150
- orbitofrontalis 190; 202.13 a, 14; 207.1 + 2 dorsal tegmental region, see: area tegmentalis
- periamygdalaris 201.19 dorsalis
- perirhinalis p. 375 dorsal thalamus p. 237
- prepiriformis 89.15; 193.24; 207.16; p.344 dorsum sellae 43.8
- retroplenialis 205.2 dura mater 50 (legend)
"cortex visceralis" 187.1 dura mater encephali 52.6; 53.5
cortical veins, see: venae cerebri superficiales dura mater spinalis 58.20 + 39; 60.4
corticocochlear projection p. 178
corticonuclear projection p. 228 ektokanthion, syn: canthus externus; angulus oculi
corticopontine connections p. 258 lateralis 39.24
corticostriate projection 176; p. 260 eminentia medialis 15.10
corticothalamic connections p. 237 encephalon 2.1/7
Subject Index 425
endosteum 53.16 fibrae

epicritic conduction pathway p. 154, 155 - amygdalofugales 91.33
epicritic systems p. 149 - amygdalofugales ventrales 193.12; 196.11;
epidural veins, see: plexus venosi vertebrales interni p. 324
epithalamus p. 296 - arcuatae externae 106.27; 108.30; 160.15
epithelium olfactorium 207.11 - arcuatae internae 108.24; 130.3; 131.5;
extrageniculate visual pathway 145 p. 154
extraocular muscles 140.1/6 - bulbospinales 186.11
"extrapyramidal" motor system p. 258 - caudatopallidales 90.21
eye movements 140; p. 169, 185 - cerebellovestibulares 104.19
- corticonucleares, see: tractus corticonuclearis
falx cerebelli 43.29; 52.32 - corticotegmentales 101.26; 111.20; 169.10;
falx cerebri 44.11 ; 52.7; 53.21 184.2
fascia dentata 94.20; 203.fd; p.334 - nervi facialis 105.22
fasciculi proprii 121.26 - olivocerebellares, see also tractus olivocerebellaris
fasciculus 106.21 ; 107.23; p. 222
- anterolateralis, syn: anterolateral bundle of - pallidotegmentales, see: tractus pallidoreticularis
Gowers 99.14; 106.22; 117.15; 121.31; - pontocerebellares, syn: fibrae transversae pontis
129.13; p.151 101.28; p. 225
- cuneatus 15.21; 109.10; 121.17; 130.6; - pontospinales 187.13
131.9; p.154 - radicis dorsalis nervi spinalis, see: dorsal root
- dorsolateralis (Lis sauer) 116.9; 121.18; 123.2 fibres
- gracilis 15.23; 109.9; 121.16; 130.7; - reticulospinales 188.16
131.8; p.154 - strionigrales 97.30; 171.4; 173.4
- lenticularis 91.25; 93.28; 171.8; 173.10; - transversae pontis, see: fibrae pontocerebellares
p.261 field H of Forel, see: area tegmentalis H
- longitudinalis dorsalis (Schiitz) 98.21 136.6; fila olfactoria, see: nervus olfactorius
191.9; 197.27; p.308 fila radicularia 19 ; 20
- longitudinalis inferior 215.8 filamentum terminale, syn: filum terminale 19;
- longitudinalis medialis 100.20; 106.20; 20; 50.17
117.17; 121.32; 138.5 filum terminale, see: filamentum terminale
- longitudinalis superior, syn: arcuate fascicle fimbria hippocampi 22.14; 23.11; 69.24;
84.4; 215.3; p.372 94.32; 195.17
- - brachium anterius 215.5 fissura
- - brachium posterius 215.4 - ansoparamediana 16.14; 18.26
- mamillaris princeps 66.34; 93.30; 97.37; - choroidea 193.16
194.19 - horizontalis 13.33; 16.12; 18.24
- mamillotegmentalis, see: tractus mamillotegmen- - intrabiventeris 18.30
talis - longitudinalis cerebri 6.1; 67.12
- mamillothalamicus; bundle of Vicq d'Azyr, see: - mediana anterior 17.41; 20; 121.14
tractus mamillothalamicus - orbitalis superior 30.18
- medialis telencephali, see: fasciculus telencephali- - posterolateralis 13.39; 18.34; 162
cus medialis - prebiventeris 18.28
- occipitofrontalis inferior 91.29; 215.7 - precentralis 13.25
- occipitofrontalis superior 84.2; 91.19; 215.1; - preculminata 13.27; 18.13
p. 374 - prepyramidalis 13.35
- olivocochlearis (Rasmussen) 143.9 - prima 13.29; 16.8; 18.18; 162
- ovalis 103.20; 105.21 - secunda 13.27; 18.31
- retroflexus (Meynert), see: tractus habenulointer- - superior posterior 13.31; 16.10; 18.20
peduncularis flocculus 18.23; 77.14; 166.25; p. 232
- tegmentalis dorsolateralis, see: tractus trigemino- folium vermis 13.32; 16.3; 162
thalamicus dorsalis (Wallenberg) foramen
- tegmentalis ventralis 132.17 - costotransversarium 58.10
- telencephalicus medialis, syn: medial forebrain - ethmoidale anterius 30.13
bundle (M.F.B.) 191.13; 197; 207.8 - ethmoidale posterius 30.15
- thalamicus 93.26; 171.3; 173.8 - infraorbitale 30.26; 42.36
- uncinatus cerebelli 112.17; 138.4; 165.10; - interventriculare (Monroi) 3.3; 12.6; 21.4;
166.22; p.232 65.21
- - ramus ascendens 165.6; 166.11 - intervertebrale 58.24
- uncinatus (cerebri) 89.24; 215.10; p.374 - jugulare 30.44; 51.33
fasciola cinerea 23.30; 69.9; 193.15 - magnum 51.34; 54.20
fastigium 13.42; 18.3 - mastoideum 30.41
426 Subject Index
foramen granulationes arachnoideales (Pacchioni) 50.2;

- of Luschka, see: apertura lateralis ventriculi 52.2; 53.17
quarti granule cells 174.20; p.222
- of Magendie, see: apertura mediana ventriculi griseum centrale mesencephali, syn: substantia
quarti grisea centralis mesencephali; periaqueductal grey
- ovale 41.26 matter 98.5; 100.2; p.302, 321
- parietale 30.38 griseum centrale metencephali, syn: substantia
- spinosum 30.31 grisea centralis metencephali 103.6
foramina sacralia dorsalia 59.28 gyri
foramina sacralia pelvina 59.27 - Andreae Retzii 23.21
forceps major 214.14 - breves insulae 8.9; 9.17; 65.2; 86.20
forceps minor 214.2 - frontales 85.17
Forel's field H, see: area tegmentalis H - occipitales 7.11; 10.11 ; 75.2; 85.37
formatio - orbitales 10.2; 62.8; 82.36
- hippocampi, see: hippocampus - temporales transversi (Heschl), see: gyrus
- reticularis p. 197 temporalis transversus
- - lateralis 103.11; 105.8; 153; 185.6; gyrus
p.202 - ambiens 10.18; 11.14; 65.13; 74.17
- - medialis 129.7; 153; 184; p.202 - angularis 7.8
- - mesencephali 156.17; 184.7 - cinguli 11.3; 23.14; 66.5; 84.13; 180.3;
- - pontis paramedianus 148A.ll; 149.11; p. 334
p.258 - dentatus 23.24; 69.10; 74.10; 193.17;
fornix 12.3; 23.4; 191.2; 205.3; p.335 195.18
fornix precommissuralis 194.14; 195.4 - diagonalis 9.15; 87.21; 193.11
fossa interpeduncularis 13.16; 17.30; 67.35 - fasciolaris 11.29; 23.6+ 19; 69.8; 193.14;
foveola granularis (Pacchioni) 53.14 195.22
frontal eye field p. 373 - frontalis inferior 7.23-25; 62.6
fronto-orbital veins, see: venae gyrorum - - pars opercularis 7.23
orbitalium - - pars orbitalis 7.25
funiculus - - pars triangularis 7.24
- anterior 17.40; 20; 121.13 - frontalis medius 6.4; 7.21; 62.4
- anterolateralis 121. 7 - frontalis superior 6.2; 7.20; 11.1; 62.2
- lateralis 15.19; 17.38; 20; 121.6+7 - intralimbicus 11.10; 23.28; 74.12
- posterior 19; 121.3; 123.1; p.154 - longus insulae 8.7; 9.18; 65.4; 86.22
- posterolateralis 121.6 - occipitotemporalis lateralis 10.9; 11.35;
70.7; 75.7
- occipitotemporalis medialis 10.25; 11.33;
Ganglion cervicale superius 146.13 70.9; 75.6
ganglion - parahippocampalis 10.20; 11.16; 67.7;
- ciliare 146.6; 147.6 74.11; 190; p.334
- habenulae, see: nuclei habenulae - paraterminalis 11.5
- semilunare Oasseri, see: ganglion trigeminale - postcentralis 6.11; 7.2; 66.9; 84.19
- spinale 19 - precentralis 6.8; 7.17; 66.3; 84.16
- trigeminale, syn: ganglion semilunare Gasseri - rectus 10.12; 11.9; 62.14; 205.9
132.3; 133.10 - semilunaris 11.13; 65.14; 81.34; 91.11
general sensory systems p. 149 - supramarginalis 7.9
general somatic afferent system p. 144, 149 - temporalis inferior 7.39; 10.8; 65.9; 75.8
general somatic efferent system p. 144; see also - temporalis medius 7.35; 65.7; 74.6
motor systems p. 247 - temporalis superior 7.33; 65.5
general visceral afferent system p. 144, 164 - temporalis transversus (Heschl) 68.8; 141.2;
general visceral efferent system p. 144 142.2
geniculocalcarine tract, see: radiatio optica - uncinatus 11.12; 23.31; 66.11; 74.15
genu corporis callosi 12.4; 85.2; 214.3
genu nervi facialis 104.22; 126.13
glabella 39.25 habenula 13.4
glabella-inion line 39.01 hemiballism p. 264
globus pallidus, syn: pallidum dorsale 28.6; hemispherium cerebelli 7.16; 9.25; 16;
64.28; 81.27 163.12; 164.8; p.221
- pars lateralis 27.8; 65.26; 86.23; 91.7; - pars intermedia 16; 18; 162; 164.7; p.229
176.6 - pars lateralis 164.8
- pars medialis 27.9; 65.28; 86.24; 91.8; Heschl's transverse gyrus, see: gyrus temporalis
176.7 transversus
Subject Index 427
hippocampus, syn.' formatio hippocampi 23; lamina

68.35; 190; 204.10+11; p.334 - interna (of calvaria), syn.' inner table 53.4
- afferents p. 336 - medullaris externa (thalami) 92.25; 94.23;
- efferents p. 340 p.237
- intrinsic connections p. 335 - medullaris interna (thalami) 25.8; 80.27;
- precommissuralis 23.8 92.26; 94.22; p.237
- retrocomrnissuralis 23.22-24 - medullaris lateralis (nuclei lentiformis) 65.25;
- supracommissuralis 23.16-18 86.7; 90.23; 92.31
homoiostasis p. 293, 349 - medullaris medialis (nuclei lentiformis) 65.27;
horizontal line or plane of Frankfurt (Reid), syn.' 86.8; 90.24; 92.32
Deutsche Horizontale 39.FH - quadrigemina, syn.' tectum mesencephali 12.21
Huntington's disease p. 268 - terminalis 12.8; 87.1
hypophysis 192 laminae albae cerebelli 71.25; 82.16
- lobus anterior, syn.' lobus anterior hypophyseos; lateral epidural vein 59.4
adenohypophysis 192.18 + 19 + 20 lateral medullary vein, see: vena spinalis lateralis
- - pars distalis 192.19 lateral motor column p. 253
- - pars infundibularis 192.18 lateral motor system p. 253
- - pars intermedia 192.20 lateral paracore p. 350, 360
- lobus posterior, syn.' lobus posterior hypophy- lateral pontine vein, see: vena pontis lateralis
seos; neurohypophysis 192.21 lateral sinus, see: sinus transversus
hypothalamohypophyseal pathways p. 308 lateroposterior orbitofrontal cortex p. 345
hypothalamus 12.9; 65.34; 87.22; 192; p.296 lemniscal systems p. 149
lemniscus lateralis 101.18; 103.28; 141A.7;
incisura preoccipitalis 7.15; 10.10 142.15
incisura tentorii 41.12; 44.27 lemniscus medialis 95.37; 99.22; 106.25;
incisura unci 11.15; 93.18 130.2; 131.4; p.154
indusium griseum 23.17; 66.7; 92.1; 193.2; lemniscus trigeminalis 132.14; 133.4; p. 160
195.3 lens crystallina 147.1
inferior petrosal vein (Dandy; inconstant), see: leptomeninges = arachnoidea + pia mater 50
vena petrosa inferior (legend)
inferior striate veins, see: venae thalamostriatae ligamentum arcuatum mediale 59.6
inferiores ligamentum denticulatum 58.23
infrapedicular vein, see: vena intervertebralis ligamentum longitudinale posterius 58.41; 59.17
superior ligth reflex (pathway) p. 185
infundibulum 9.4; 13.20; 64.37; 192.17 limbic lobe p. 334
inion (protuberantia occipitalis externa) 39.45; limbic midbrain area p. 302
52.31 limbic system p. 293, 348
inner table, see: lamina interna - midbrain circuit p. 302
insula (Reili) 8; 89.9; p. 241 limbus Giacomini 11.11; 23.29; 74.13; 195.13
intermediate grey matter, see: substantia interme- limen insulae 8.12; 9.16
dia line of Gennari p. 181
intermediate raphespinal projection p. 281 lingula cerebelli 13.24; 15.35; 18.17; 162.Li
interneurons in the thalamic nuclei p. 239 liquor cerebrospinalis 50 (legend)
intumescentia cervicalis 20 lobulus
intumescentia lumbosacralis, syn.' intumescentia - biventer 18.29; 162
lumbalis 20 - centralis 18.15; 75.13; 162
iris 146.2 - gracilis 16.15; 18.27; 162
isocortex, see: neocortex - paracentralis 6.10 11.20
isthmus gyri cinguli 11.31; 70.11 - parietalis inferior 6.15; 7.6
- parietalis superior 6.14; 7.4
juxtallocortex p. 334 - quadrangularis 16.7; 18.14
- semilunaris inferior 16.13; 18.25; 162
"Kleinhirnbriickenwinkelzisterne", see: cisterna - semilunaris superior 16.11; 18.21; 162
pontocerebellaris - simplex 16.9; 18.19; 162
Kolliker-Fuse nucleus 154.nKF; p.203, 212 lobus
- connections 157.16; p.211 - anterior cerebelli 16; 18 162
- anterior hypophyseos, see: hypophysis, lobus an-
lacunaelaterales 44.5; 52.1; 53.15 terior
lamina - flocculonodularis 18; 161.12; 166.24+ 25;
- affixa 15.26; 17.2; 22.1; 84.8; p.237 p.232
- cribrosa 52.14 - frontalis 4.9
- externa (of calvaria), syn.' outer table 53.2 - limbicus 4.11; p.334
428 Subject Index
lobus monoaminergic cell groups

- occipitalis 4.2 - B1 p.197
- parietalis 4.3 - B3 1548; 155.48; p.197
- posterior cerebelli 16; 18; 162 - B5 154C; 155.54; p.202
- posterior hypophyseos, see: hypophysis, lobus - B6 155.44; p.202
posterior - B7 155.40; p.202
- temporalis 4.4 - B8 154C; 155.44; p. 202
locomotion p. 349, 361 - Cl 154A; 159.25; 189.3; p.205
locus coeruleus 79.9; 102.3; 158.37; 188.1; - C2 154A; 159.25; 189.4; p.205
p.204 mossy fibres p. 222, 224
- afferents 159; p.216 motor cortex 170.3; p.247, 374
locus coeruleus complex 158.36+37; p.213 - primary, syn: Mi-region p.373
- ascending efferents 158; p. 213 - supplementary, syn: M2-region 217.M2;
- descending efferents 158; 188; p.256, 287, p.373
290 motor systems (medial, lateral, "third") 175;
long corticofugal pathways p. 247 p. 247, 253, 256
L-region of Holstege et al. 187.8; p. 287, 290 M-region of Holstege et al. 187.7; p.287, 290
multimodal association cortex p. 365, 372
macula lutea 145.5 myelencephalon, see: medulla oblongata
magnocellular basomedial telencephalic complex
180; 181; p. 273 neocortex, syn: isocortex p. 334
marginal cells, see: cellulae marginales nervi ciliares breves 146.4; 147.4
margo liber alae minoris ossis sphenoidalis 38.9 nervus
margo liber falcis cerebri 52.8 - abducens (n. VI) 14.30; 17.15; 104.24
margo superior partis petrosae ossis temporalis - accessorius (n. XI) 14.34; 17.21
38.13; 39.29 - - radices craniales 14.37
meatus acusticus externus, see: porion - - radices spinales 14.38
meatus acusticus internus 43.20 - cochlearis, syn: nervus octavus (n. VIII), pars
medial epidural vein 59.3 cochlearis 141A.14; 142.21
medial forebrain bundle (M.F.B.), see: fasciculus - facialis (n. VII) 14.27; 17.16; 104.26;
telencephalicus medialis 105.28; p.164
medial lemniscus system p. 154 - glossopharyngeus (n. IX) 14.31; 17.19;
medial motor column p. 253 106.23; p.164
medial motor system p. 253 - hypoglossus (n. XII) 14.35; 17.22; 108.28
medial nucleus of the diagonal band of Broca, see: - intermedius (n. VII) 14.28; 17.17
nucleus gyri diagonalis, pars dorsalis - mandibularis (n. V3) 132.V3
medial temporal cortical veins, see: venae tempor- - maxillaris (n. V2) 132.V2
ales mediales - octavus, pars cochlearis, see: nervus cochlearis
median paracore p. 350, 357 - - pars vestibularis, see: nervus vestibularis
medulla oblongata, syn: myelencephalon 2.7; - - see: nervus vestibulocochlearis
12.15; 19; 20 - oculomotorius (n. III) 12.13; 17.11; 99.26;
medulla spinalis, syn: spinal cord 1.4; 2.8; 19; 146.8; 147.7
20 - olfactorius (n. I), syn: fila olfactoria 207.10
- pars cervicalis 20 - ophthalmicus (n. Vl) 132.Vl
- pars lumbalis 20 - opticus (n. II) 12.11; 14.22; 144.2;
- pars thoracica 20 145.6
- substantia grisea 121.5+8+28; 122 - spinalis 19
mesencephalic locomotor area 213.4; p. 267, 361 - - radix dorsalis 14.18; 15.47; 19
mesencephalon 2.4 - - radix ventralis 14.39; 17.23
mesocortex p. 334, 335 - trigeminus (n. V) 14.24+25; p.155
metathalamus p. 237 - - portio major, see: nervus trigeminus, radix
metencephalon 2.5 + 6 sensoria
molecular layer of cerebellar cortex p. 222 - - portio minor, see: nervus trigeminus, radix
monoaminergic cell groups motoria
- Al 154A; 159.27; p.204 - - radix motoria, syn: nervus trigeminus, portio
- A2 154A; 159.28; p.204 minor 14.24; 103.26; 112.14+25; 132.4;
- A5 1548; 189.2; p. 204 133.8+9
- A6 154C; p.204 - - radix sensoria, syn: nervus trigeminus, portio
- A7 154C; 189.1; p.204 major 14.25; 17.14; 103.27; 132.5;
- A8 p.260 133.11; p. 155
- A9 p.260 - trochlearis (n. IV) 14.7; 15.34; 102.11
- Al0 p.260, 320 - vagus (n. X) 14.33; 17.20; 107.20; p.164
Subject Index 429
nervous nucleus
- vestibularis, syn: nervus octavus (n. VIII), pars - basalis amygdalae, see: corpus amygdaloideum,
vestibularis 137.8; p.165, 224 nucleus basalis
- - fibrae afferentes 138.13 - caudatus 26.1 ; 28; 29; 176.2
- - fibrae efferentes 138.14 - centralis amygdalae, see: corpus amygdaloideum,
- - ramus descendens 106.17 nucleus centralis
- vestibulocochlearis, syn: nervus octavus (n. VIII) - centralis superior (mesencephali, Bechterew)
14.29; 17.18; 105.27 102.8; 153.2; 194.25
neurohypophysis, see: hypophysis, lobus posterior - centromedianus, syn: centre median (Luys)
nigroreticular projection p. 267 25.10; 80.30; 94.9; 156.7; 167.11
nigrostriatal dopaminergic projection p. 264 - cochlearis dorsalis 106.1; 141A.16; 142.23
nigrotectal projection p. 267 - cochlearis ventralis 106.10; 141A.15; 142.22
nigrothalamic projection p. 265 - cornucommissuralis 119.6
nociceptive units p. 151 - corporis trapezoidei, syn: nucleus corporis trape-
nodulus 13.40; 18.33; 76.8; 162; 166.24; zoidei dorsalis, nucleus corporis trapezoidei ven-
p.232 tralis 104.16; 141A.12; 142.19
non-specific thalamic nuclei p. 238 - corticalis amygdalae, see: corpus amygdaloi-
noradrenergic cell groups (and pathways) p. 204, deum, nucleus corticalis
290 - cuneatus lateralis 107.2; 108.2; 161.15;
- ascending efferents p. 217 p.154, 225
nuclei - cuneatus medialis 108.3; 130.4; 131.7
- arcuati 106.14; 108.16 - cuneiformis 99.5; 101.7; 153.6
- centrales cerebelli (nucleus dentatus, nucleus - Darkschewitschi 97.11
emboliformis, nucleus fastigii and nucleus - dentatus 71.23; 105.1; 165.12; 166.19;
globosus) 161 9; p.222 170.17; p. 222
- habenulae, syn: ganglion habenulae 24.14; - dorsalis nervi vagi, syn: nucleus parasympathicus
68.28; 85.27; 95.8; 196.23 nervi vagi 107.5; 126.18
- intralaminares (thalami) 93.5; 128.1; 129.3; - dorsomedialis (hypothalami) 192.8; 197.12;
156.4; 165.2+3+4; 176.4; p.233, 238, 246 p. 316
- laterales thalami 24.4+7; p.237, 244 - emboliformis 71.20; 76.18; 103.2; 163.10;
- lemnisci lateralis 141.9 p.222
- mediani thalami 93.6 - fastigii 71.21; 103.4; 164.11; 165.11;
- periolivares 142.16 166.21; p.222
- pontis 102.10; 103.15; 161.6; 174.14; - funiculi anterioris, syn: nucleus reticularis para-
p.225 medianus 107.10; 108.13; p.225
- posteriores thalami p.237, 241 - funiculi lateralis, syn: nucleus reticularis latera-
- preoptici, see: nucleus preopticus lis 107.13; 108.12; p.225
- pulvinares 24.15; 25.15; 85.28; 97.1 - gigantocellularis 104.12; 153.10; 156.28;
- raphes 153; p. 197, 206, 207, 279 184.12
- salivatorii, syn: nucleus salivatorius 126.16 - globosus 71.22; 76.17; 103.3; 163.11;
- septi 89.6; 197.4+7; 198A.l0+11; 201.11 p.222
- tuberales 92.12 - gracilis 108.1; 130.5; 131.6
- ventrales thalami 25.2+5+12+13; p.237, - gyri diagonalis 89.10; 90.7; 198.14+16;
239 203.8
- vestibulares 148.15 - - pars dorsalis 198.14; p.310
nucleo-olivary fibres p. 232 - - pars ventralis 198.16, p. 310
nucleus - habenulae lateralis 155.11; 197.6; p.312
- accessorius nervi oculomotorii (Edinger-West- - habenulae medialis 155.12; 197.41; p. 312
phal), syn: nucleus parasympathicus nervi oculo- - infundibularis 92.13; 192.15; 197.17;
motorii 95.13; 98.11; 126.9; 146.9; 147.8 p.302, 318
- accessorius olivae superioris, see: oliva superior, - intercalatus 107.6
nucleus medialis - intercollicularis, syn: zona intercollicularis
- accumbens 27.12; 87.19; 89.11; 205.10 100.1
- ambiguus 107.12; 126.17 - intermediolateralis 118.4; 125.3
- anterior amygdalae, see: corpus amygdaloideum, - interpeduncularis 100.10; 101.12; 194.23;
nucleus anterior 198.49
- anterior hypothalami 91.7; 184.4; 192.7; - interpositus (nucleus emboliformis plus nucleus
197.14; 198.19; p. 313 globosus) 164.12; 166.20; p.222
- anterior thalami 24.10; 92.3; 167.4; 194.5; - interstitialis (Cajal) 95.12; 98.10; 137.1;
195.8; p.238, 240 138.1; 149.7
- basalis accessorius amygdalae, see: corpus amyg- - interstitialis rostralis fasciculi longitudinalis me-
daloideum, nucleus basalis accessorius dialis 149.6; 150.2; p.258
430 Subject Index
nucleus nucleus
- interstitialis striae terminalis, syn: bed nucleus of - paratenialis (thalami) p. 238
the stria terminalis 91.10; 198.4; 199.2; - paraventricularis (hypothalami) 91.5; 192.2;
201.3; p. 332 197.8; 198.18; p.314
- lateralis amygdalae, see: corpus amygdaloideum, - peripeduncularis 97.9; p.296, 313
nucleus lateralis - pontis centralis caudalis, see: nucleus reticularis
- lateralis dorsalis 24.12; 25.6; 94.2 pontis caudalis
- lateralis olivae superioris, see: oliva superior, - pontis centralis oralis, see: nucleus reticularis
nucleus lateralis pontis oralis
- lateralis posterior 24.13; 25.14; 94.3; 95.6; - posterior hypothalami 92.9; 93.8; 192.4;
167.10; p.244 198.22; p.318
- lemnisci lateralis dorsalis 102.4; 142.12 - posterior thalami p. 238
- lemnisci lateralis ventralis 103.12; 142.13 - pregeniculatus 94.16
- lentiformis 26.2 - premamillaris dorsalis p. 318
- limitans (thalami) p. 238 - premamillaris ventralis p. 318
- medialis amygdalae, see: corpus amygdaloideum, - preopticus (nuclei preoptici) 184.3; 192.6+
nucleus medialis 12; p.296
- medialis olivae superioris, see: oliva superior, - preopticus lateralis 90.5; 192.12; p.312
nucleus medialis - preopticus medialis 90.6; 192.6; 197.15;
- medialis thalami 24.11; 94.4; 167.5; 194.7; p.311
201.2; p.237, 240 - prepositus hypoglossi 105.6; 106.4; 148.16;
- - pars magnocellularis 207.5 149.15
- medialis ventralis, see: nucleus ventralis postero- - pretectalis principalis, see: regio pretectalis
medialis, pars parvocellularis - proprius 117.3; 121.23; 129.12; p.150
- medullae oblongatae centralis 107.9; 109.7; - radicis spinalis nervi accessorii 126.20
153.14; 156.29; 184.13 - raphes dorsalis 153.1 ; p. 279
- mesencephalicus nervi trigemini 99.3; 126.2; - raphes magnus 104.13; 106.9; 153.4;
133.6; p. 160 p.279
- motorius nervi trigemini 103.9; 126.12; - raphes obscurus 107.11; 153.5; p.279
133.12; 185.7 - raphes pallidus 183.10; p.279
- nervi abducentis 104.8; 126.14; 148.12; - raphes pontis 103.13; 153.3
149.13 - reticularis gigantocellularis, see: nucleus giganto-
- nervi facialis 104.11; 105.10; 126.15; cellularis
185.8 - reticularis lateralis, see: nucleus funiculi lateralis
- nervi hypoglossi 107.7; 126.19; 185.9 - reticularis paramedianus, see: nucleus funiculi
- nervi oculomotorii 98.12; 137.2; 148.7; anterioris
149.9 - reticularis parvocellularis p. 203
- nervi trochlearis 100.6; 126.11; 137.3; - reticularis pontis caudalis, syn: nucleus pontis
148.9; 149.10 centralis caudalis 103.10; 153.9; 156.23;
- of Onuf p. 290 184.11
-olfactoriusanterior 89.14; 205.11; 207.13; - reticularis pontis oralis, syn: nucleus pontis cen-
p. 344 tralis oralis 102.7; 153.7; 156.20; 184.8
- olivaris accessorius dorsalis 79.17; 107.14; - reticularis tegmenti pontis (Bechterew), syn: nu-
163.17; 164.17 cleus papillioformis 102.9; 103.14; 153.8;
- olivaris accessorius medialis 79.18; 107.15; p.225
163.18; 164.18 - reticularis thalami 67.18; 93.4; 171.9;
- olivaris inferior, syn: principal olive 79.16; 173.6; p. 237, 239
107.16; 163.15; 164.16 - retroambiguus 109.6
- ovalis, syn: area ovalis 105.7; 135.6; 136.8 - reuniens (thalami) p.238
- papillioformis, see: nucleus reticularis tegmenti - ruber 67.28; 160.2; 164.1; 165.5; 194.21
pontis (Bechterew) - - pars magnocellularis 99.10; 166.9; 174.10
- parabrachialis lateralis 102.5 ; 153.12; - - pars parvocellularis 97.12; 166.8; 174.9;
199.24; p.211 p.232
- parabrachialis medialis 102.6; 136.7; 153.13; - salivatorius (inferior, superior), see: nuclei sali-
p.164, 211 vatorii
- parafascicularis 25.11; 94.10; 167.12 - sensorius principalis nervi trigemini 103.8;
- paralemniscalis 99.4; 100.3 132.7; 133.13; p.155
- parasympathicus nervi oculomotorii, see: nucleus - septi lateralis 197.4; 198.10; 205.6; p.310
accessorius nervi oculomotorii (Edinger-West- - septi medialis 197.7; 198.11; p.310
phal) - solitarius 107.4; 135.8; p.164
- parasympathicus nervi vagi, see: nucleus dorsalis - - pars cardiorespiratoria 136.11
nervi vagi - - pars gustatoria 136.9
Subject Index 431
nucleus oliva (inferior), see also nucleus olivaris inferior

- spinalis nervi trigemini 132.8; p.155 14.36; 17.35 p. 154
- - pars caudalis 108.8-10; 133.17; p.155 oliva superior (complex) 142.16/19; p.l72
- - pars interpolaris 107.8; 133.16 - nucleus lateralis, syn: nucleus lateralis olivae
- - pars oralis 105.9; 133.15 superioris 104.14; 141.10; 142.18
- subcoeruleus, syn: area subcoerulea 158.37; - nucleus medialis, syn: nucleus medialis olivae
188.1; p. 204 superioris, nucleus accessorius olivae superioris
- subcuneiformis 153.6 104.15; 141.11; 142.17
- subthalamicus, syn: corpus subthalamicum olivocerebellar fibres 160.14; 163.14; 164.19;
(Luys) 80.35; 93.10; 173.11; 174.8; 176.8 p.229
- suprachiasmaticus 192.14; 197.60; p.316 olivocochlear bundle, see: fasciculus olivocochlearis
- suprageniculatus (thalami) p. 238 (Rasmussen)
- supraopticus 90.9; 91.8; 192.13; 197.61; operculum frontale 4.7; 8.10
p.314 operculum frontoparietale 4.5; 8.4
- supraspinalis 109.8 operculum temporale 4.6; 8.11
- tegmentalis dorsalis (Gudden) 135.3; 136.5; orbitale 39.26
194.24; 197.23; p.302 organum subfornicale 197.58; p.309
- tegmentalis pedunculopontinus, pars compacta organum vasculosum of the lamina terminalis
101.9; 153.11 197.59; p.309
- - pars dissipata 100.8 os petrosum, see: pars petrosa ossis temporalis
- tegmentalis ventralis, see: area tegmentalis ven- outer table, see: lamina externa
tralis (Tsai)
- terminalis (dorsalis, lateralis, medialis) 145.14+ pachymeninx = dura mater 50 (legend)
12+11 pain conduction, see: protopathic systems
- thoracicus, syn: columna Clarki (Stilling- pallidohabenular fibres 179.11-6; p.272
Clarke) 118.5; 123.9; 161.23; p.224 pallidothalamic projection p. 261
- ventralis anterior 24.5; 92.5; 167.6; 177.4; pallidum dorsale, see: globus pallidus 182A
p.240 pallidum ventrale = part of substantia innominata
- ventralis lateralis 24.9; 93.3; 166.6; 167.7; 1828
176.5; p.233, 240 paralimbic association cortex p. 375
- ventralis posterior (nucleus ventralis postero- parallel fibres p. 222
inferior, nucleus ventralis posterolateralis and parasensory association cortex p. 365
nucleus ventralis posteromedialis) 167.8; parasubiculum p. 335
p.154, 239 parietal cortex p. 371
- ventralis posteroinferior 138.19 Parkinson's disease p. 264
- ventralis posterolateralis 24.6; 25.12; 94.8; pars atlantica, see: arteria vertebralis, pars atlantis
128.2; 129.4; 131.3; p.151 pars canalis carotici, see: arteria carotis interna,
- ventralis posteromedialis 25.13; 94.11 ; pars petrosa
132.1+13; 133.3; 135.1; p.160 pars cervicalis, see: arteria vertebralis, pars trans-
- - pars parvocellularis, syn: nucleus medialis ven- versaria
tralis 94.12; 136.3; 167.9; p.164, 238 pars intermedia cerebelli, see: hemispherium cere-
- ventromedialis (hypothalami) 92.10; 192.9; belli, pars intermedia
197.13; 198.21; p.316 pars petrosa ossis temporalis, syn: os petrosum;
- vestibularis inferior 105.5; 106.2; 137.7; petrous bone 38.13; 39.29
138.12 pars subarachnoidealis, see: arteria carotis interna,
- vestibularis lateralis (Deiters) 104.6; 105.3; pars cerebralis
137.9; 138.10; p.228 pars supracavernosa, see: arteria carotis interna,
- vestibularis medialis 104.7; 106.3; 137.6; pars cerebralis
138.11; p. 228 pediculus arcus vertebrae 58.3; 59.8 + 26
- vestibularis superior 103.5; 104.5; 137.5; pedunculopontino-reticular projection p. 267
138.9 pedunculus
- cerebellaris inferior, syn: corpus restiforme
Obere Horizontale, see: upper horizontal line or 18.5; 80.14; 106.18; 160.9; 163.13; p.222
plane (Kronlein) - cerebellaris medius, syn: brachium pontis 18.6;
obex 14.15; 15.46 81.16; 103.33; 160.10; 161.7; p.222
occipital eye field, see: visual cortex - cerebellaris superior, syn: brachium conjuncti-
ocular dominance strips p. 244 vum 18.2; 79.10; 98.31; 164.9; 165.7
oculomotor pathways 148/152 - - ramus descendens 102.21; 161.3; 164.4;
olfactory projections, primary -, secondary -, terti- 165.8
ary - 207; p. 345 - cerebri, syn: crus cerebri 14.21; 67.33;
olfactory system p. 344 74.29; 94.30; 172.13-15; p.247
olfactory vein, see: vena gyri olfactorii - corporis mamillaris 99.28; 191.14; p.308
432 Subject Index
pedunculus posterior, unpaired part of the ambient cistern,

- flocculi 18.22; 105.18 see: cisterna laminae tecti
- nuclei lentiformis 29.12 posteromedian medullary vein, see: vena spinalis
- thalami anterior 80.28; 86.4; 90.22; posterior
169.12; p.237 precuneus 11.23
- thalami inferior 169.14; 194.12; p.237 premotor area (areae 6, 8 + 44) p. 373
- thalami posterior 169.8; p.237 premotor cortex 170.4+5; p.247, 373
- thalami superior 169.7; p.237 prerolandic artery, see: arteria sulci precentralis
periaqueductal grey matter, see: griseum centrale presubiculum p. 335
mesencephali pretectum, see: area pretectalis
periolivary cochleonuclear projection p. 178 primary visual cortex, see: area striata
periosteum 53.1; 58.22 Probst's commissure, see: decussatio lemniscorum
pes hippocampi 66.42 lateralium
petrous bone, see: pars petrosa ossis temporalis processus reticularis 117.4; 121.24
pia mater 50 (legend); 52.10; 53.9; 58.17 + 37 processus transversus 58.4
planum polare 141.19 propriobulbar projections p.218
planum temporale 68.8; 141.1 prosencephalon 2.1-3
plexus protopathic pathway p. 151, 155
- basilaris 41.32; 43.37; 52.21 protopathic systems, syn: pain conduction system
- choroideus ventriculi lateralis 46.6; 51.4; p.149, 155, 164
65.19; 67.32; 68.19; 84.9 protuberantia occipitalis externa 52.31
- choroideus ventriculi quarti 14.12; 15.42; protuberantia occipitalis interna 52.30
18.11; 51.26 pulvinar thalami 14.2; 15.31; 144.12; 145.22;
- choroideus ventriculi tertii 13.9; 46.10; 51.6; 167.13+14+15; p.238, 244
68.22 pupilla 146.1
- pterygoideus, syn: plexus venosus pterygoideus Purkinje-cells 174.19; p.222
42.32 putamen 29.4; 65.24; 86.18; 91.6; 176.3
- venosi vertebrales externi, syn: anterior + posteri- pyramidal tract syndrome p. 258
or extravertebral veins 58.48; 59.16 pyramis (medullae oblongatae) 9.21; 17.36;
- venosi vertebrales interni, syn: anterior + posteri- 172.17
or epidural veins 42.47; 58.15+46 pyramis vermis 13.36; 16.5; 162.Py
- venosus foraminis ovalis 41.26
- venosus pterygoideus, see: plexus pterygoideus
- venosus vertebralis internus anterior 41.52; quadrigeminal vein, see: venae mesencephalicae
58.46; 59.20
- venosus vertebralis internus posterior 58.15;
59.9 radiatio acustica 141A.2
plica petroclinoidea anterior 44.19 radiatio corporis callosi 63.11; 71.15; 214.13
plica petroclinoidea posterior 44.20 radiatio optica, syn: geniculocalcarine tract
polus frontalis 4.10 85.13; 95.32; 144.11; 145.23; 169.18;
polus insulae 8.13; 9.10; 82.37 p.181, 244
polus occipitalis 4.1 - genu occipitale 144.20
polus temporalis 4.8 - genu temporale 144.5
pons (Varoli) 2.5; 12.14; 14.26; 17.31; radices craniales nervi accessorii 14.37
75.19 radices spinales (or radix spinalis) nervi accessorii
pontes grisei caudatolenticulares 28.14; 29.8 14.38; 116.10
"pontine gaze centre" p. 188 radices ventrales (or radix ventralis) nervi spinalis
"pontine taste area" p.211 14.39; 17.23; 117.18; 121.15; 124.15
pontine venous plexus, see: venae pontis radix dorsalis nervi spinalis, see also: dorsal root
pontocerebellar fibres 161.7; 174.14-20; p.224 (fibres) 14.18; 15.47; 19; 117.8; 121.2
pontospinal tract, see: tractus reticulospinalis me- rami
dialis - ad medullam oblongatam, see: arteria cerebelli
porion, syn: meatus acusticus externus 39.31 inferior posterior, rami medullares
portio major nervi trigemini, see: nervus trigemin- - ad pontem, see: arteriae pontis
us, radix sensoria - choroidei posteriores laterales, see: arteria cere-
portio minor nervi trigemini, see: nervus trigemin- bri posterior, pars postcommunicalis, ramus
us, radix motoria choroideus posterior lateralis
posterior extravertebral veins, see: plexus venosi - choroidei posteriores mediales, see: arteria cere-
vertebrales externi bri posterior, pars precommunicalis, ramus chor-
posterior funiculus, somatotopically organised oideus posterior medialis
p.154 - diencephalici anteroinferiores, see: arteriae cen-
posterior interposed nucleus, see: nucleus globosus trales anteromediales
Subject Index 433
rami Rexed, subdivision of the spinal cord grey matter

- diencephalici (posteriores + superiores), see: ar- according to - 122.I/X
teria cerebri posterior, pars postcommunicalis, rhinencephalon p. 293
rami thalamici rhombencephalon 2.5/7
- diencephalici posteroinferiores, see: arteriae cen- rolandic artery, see: arteria sulci centralis
trales posteromediales rostral interstitial nucleus of the medial longitudi-
- interpedunculares profundi, see: arteriae cen- nal fascicle 139.5
trales posteromediales rostrum corporis callosi 12.5; 63.15; 214.5
- medullares, see: arteria cerebelli inferior posteri- rubro-bulbar and -spinal tract 161.4 ; 163.3;
or and: arteria vertebralis, pars intracranialis 174.21; 175.11
- precuneales, see: arteria precunealis
- radiculares, see: arteria vertebralis, rami spinales Schutz, fascicle of -, see: fasciculus longitudinalis
- striati arteriae cerebri mediae 64.33 dorsalis
ramus senile dementia of the Alzheimer's type p.276
- cingularis, see: arteria callosomarginalis septal region, see: septum and: nuclei septi
- colliculi inferioris, see: arteria cerebelli superior, septum medianum posterius 121.4
ramus mesencephalicus septum pellucidum 12.2; 63.13
- communicans (nervi spinalis) 146.14 septum precommissurale, see: septum verum
- corporis callosi posterior, see: arteria pericallosa septum verum, syn: septum precommissurale
posterior 193.9; 194.16; p.296, 310
- dorsalis venae sacralis lateralis, syn: ascending serotoninergic cell groups (and pathways) p.279
sacral vein 59.21 sexual behaviour p. 293, 349
- laminae tecti, see: arteria cerebelli superior, short association fibers p. 365
ramus mesencephalicus sinus
- mastoideus arteriae occipitalis 30.40 - cavernosus 41.22; 42.31
- meatus acustici interni, see: arteria labyrinthi - durae matris, syn: sinus venosi durales 42;
- orbitofrontalis lateralis, see: arteria frontobasalis 53.19
lateralis - intercavernosus anterior 52.16; 192.23
- orbitofrontalis medialis, see: arteria frontobasalis - intercavernosus posterior 43.9; 52.17; 192.22
medialis - marginalis 43.43; 52.33
- quadrigeminus, see: arteria cerebelli superior, - occipitalis 42.23; 43.30
ramus mesencephalicus - petrosus inferior 41.30; 42.28; 43.22
- sympathicus (plexus carotici) 146.7 - petrosus superior 41.34; 42.27; 44.25
- vermis superior, see: arteria vermis superior - rectus 42.18; 45.15; 52.28
raphe nuclei 153.1/5; p. 197 - sagittalis inferior 42.7; 44.9; 52.23
- ascending projections p. 206 -sagittalissuperior 42.2; 44.1; 52.3; 53.19
- afferents p. 207 - sigmoideus 41.49; 42.26; 43.25
- descending projections p. 279 - sphenoparietalis 41.13; 43.2; 44.16
raphespinal projection p. 279 - tentorii 41.45 + 48; 44.29; 45.38
raphespinal system p. 160, 256 - transversus, syn: lateral sinus 41.47; 42.22;
recessus 43.27
- infundibuli 13.19; 21.7 - venosi durales, see: sinus durae matris
- lateralis ventriculi quarti 14.11; 15.40; 18.9; siphon arteriae carotidis internae, syn: carotissi-
21.15 phon 38.40; 39.22
- opticus 13.17; 21.6; 64.35 siphon point 39.22
- pinealis 13.7; 21.10 somatosensory cortex p. 154, 155, 160, 240,
- suprapinealis 13.3; 21.9; 85.11 370
red nucleus, see: nucleus ruber - somatotopical localization p. 154, 155, 371
reference points, lines and planes 38; 39 somatosensory relay nuclei p. 160
relay nuclei p. 210, 238 somatotopic organization
regio preoptica p.296, 311 - of fasciculus anterolateralis p. 151
regio pretectalis 145.15 - of funiculus posterior p. 154
reticulothalamic fibres p. 210 - of somatosensory cortex p. 155, 371
reticulocerebellar fibres p. 225 - of cerebellar cortex p. 225
reticulospinal system p. 256 spatium subdurale 53.6; 58.19
retina 145.4 special sensory system p. 165
retinal ganglion cells p. 180 special somatic afferent system p. 144, 165
retinogeniculocortical projection 144; 145 special visceral afferent system p. 144, 164
retinohypothalamic projectioon p. 180 special visceral efferent system p. 144
retinopretectal projection 145 spinal cord, see: medulla spinalis
retino-topical localization p. 181 spinoreticular projection 128.5; 129.10; p.208
retropulvinar cistern, see: ala cisternae ambientis spinoreticular systems p. 151
434 Subject Index
spinovestibular fibres 138.17; p. 168 sulcus

splenium corporis callosi 12.17; 70.18; 214.12 - cinguli 11.2
stratum sagittale 144.18; 169.19 - - pars marginalis 11.21
stratum subependymale 64.21 - circularis insulae 8.5; 65.1
stria - collateralis 10.19; 11.34
- acustica dorsalis 141.14 - corporis callosi 11.4; 23.15; 66.6
- Gennari 72.14; 75.9 - fimbriodentatus 23.26
- longitudinalis lateralis 23.2 + 18; 63.10; - frontalis inferior 6.5; 7.22
89.18; 92.23; 195.2 - frontalis superior 6.3; 7.19
- longitudinalis medialis 23.1+16; 63.9; - hippocampi 23.25 + 30; 93.19
89.19; 92.24; 195.1 - hypothalamicus 13.13
- medullaris thalami 15.28; 92.27; 194.4; - intermedius posterior 15.22
p. 308 - intraparietalis 7.7; 8.13; 70.2
- olfactoria lateralis 9.14; 191.18; 196.4; - lateralis 7.32
207.15; p.344 - - ramus anterior 7.27
- olfactoria medialis 9.11; 196.3; p. 344 - - ramus ascendens 7.26
- terminalis 22.2; 93.23+34; 193.3; 196.21; - - ramus posterior 7.10
199.1; 201.1; p. 324 - - anterior 17.39
- - bed nucleus of the -, see: nucleus interstitialis - - posterior 14.17; 15.20
striae terminalis - limitans p. 144
- - postcommissuralis 194.15 - lunatus 7.12; 75.1
- - precommissuralis 194.14 - medianus (ventriculi quarti) 15.11
striae acusticae dorsales 106.15; 141A.13; - medianus posterior 15.24; 121.1
142.24 - occipitalis anterior 7.13
striae medullares ventriculi quarti 106.16 - occipitotemporalis 10.7; 68.14
striatal circuits 176; 177; 182; p.260 - olfactorius 10.13; 62.9
striatum, see also corpus striatum - paracentralis 80.1
- "limbic" p. 270 - parieto-occipitalis 6.16; 11.26; 72.5;
- "non-limbic" p.270 80.7
- dorsal 182A; p.269 - parolfactorius anterior 11.8; 87.14
- organization p. 267 - parolfactorius posterior 11.6
- ventral 182B; p.269 - postcentralis 6.12; 7.3
- - limbic connections p. 269 - precentralis 6.7; 7.18
striosomes p. 268 - rhinalis 10.17; 11.17; 64.16
subcoeruleo-spinal pathway 188.16; p.290 - subparietalis 11.24
subcortical veins, see: venae cerebri profundae - temporalis inferior 7.36; 10.5; 68.12
subfornical organ, see: organum subfornicale - temporalis superior 7.34; 68.10
subiculum 23.23; 93.20; 195.10; 203.sub; superior median collicular vein (Duvernoy), see:
205.16; p.334 venae mesencephalicae
substantia superior occipitofrontal bundle, see: fasciculus
- gelatinosa (Rolandi) 117.2; 121.22; p. 150 occipitofrontalis superior
- grisea centralis mesencephali, see: griseum cen- superior olivary complex, see: oliva superior
trale mesencephali (complex)
- grisea centralis metencephali, see: griseum cen- superior petrosal vein (Dandy), see: vena petrosa
trale metencephali superolateral + inferolateral pontine arteries, see:
- grisea intermedia, see: substantia intermedia arteriae pontis, rami laterales
- innorninata 64.32; 90.4; 91.9 supplementary motor area (M2) 217.M2; p.373
- intermedia, syn: substantia grisea intermedia suprapedicular vein, see: vena intervertebralis infe-
117.5; 121.28; 124.12; 174.24; p.150 rior
- nigra 67.34; 74.28; 173.12 Sylvian vein, see: vena cerebri media superficialis
- - pars compacta 98.14; 176.9
- - pars reticulata 98.15; 176.10 taenia choroidea 15.25; 17.1; 22.4
- perforata anterior 9.13; 64.18; p.344 taenia fimbriae 22.13
- perforata posterior 17.29 taenia fornicis 22.5
subthalamus, see: nucleus subthalamicus taenia thalami 11.30; 15.29; 17.3; 22.3
sulci occipitales 7.14 taenia ventriculi quarti 15.16
sulci orbitales 7.28; 10.1; 62.7 tapetum 214.11
sulcus taste p. 149, 164
- basilaris pontis 17.32 taste pathway p. 164
- calcarinus 11.28; 72.8; 75.4; 144.22; 145.24 tectal vein, see: venae mesencephalicae
- centralis 7.1; 8.9 tectospinal fibres 174.22; 178.12
- centralis insulae 8.8; 65.3 tectum mesencephali, see: lamina quadrigemina
Subject Index 435
tegmentum, see also: area tegmentalis tractus

- mesencephali 74.26; 79.8 - reticulospinalis 176.19; p. 256; see also fibrae
- mye1encephali 77.19 reticulospinales
- pontis 75.18 - - lateralis p. 282
- pontis laterale 187.6/12; p. 286 - - medialis p. 256, 282
- - connections 187; p.287 - rubrospinalis 163.3; 174.21; 185.5
tela choroidea ventriculi quarti 12.28; 15.43; - solitariospinalis 136.14
18.10; 77.12 - solitarius 107.18; 135.11
tela choroidea ventriculi tertii 12.19; 194.6 - spinalis nervi trigemini 132.9; 133.14
telencephalon, see: cerebrum - spinocerebellaris anterior 117.14; 121.25;
telencephalon impar 2.2 123.6; 160.8; 161.17; p.225
tentorium cerebelli 44.28; 52.28 - spinocerebellaris posterior 117.12; 121.19;
tertiary olfactory projections, see: olfactory projec- 123.3; 160.17; 161.18; p. 224
tions - spino-olivaris 163.19; 164.21
thalamic nuclei - spinoreticularis 123.7; 129.10; 156.18
- anterior (connections) p.240 - spinotectalis 152.24
- lateral (connections) p.241, 244 - spinothalamicus 97.22; 123.8; 128.5; 129.9;
- medial (connections) p.240 p.151
- posterior (connections) p. 241 - spinovestibularis 138.17
- ventral (connections) p. 239 - tectobulbaris 156.19
thalamocortical circuits p. 239 - - lateralis 152.14
thalamostriate fibres 176.1; 182.1; p.261 - - medialis 152.13
thalamus 12.18; 14.1; 17.5; 24; 167; p.237 - tectopontinus 101.19; p.228
tonsilla cerebelli 13.45; 18.32; 77.9; 162 - tectospinalis 99.18; 102.19; 156.19; 174.22;
tractus 178.12
- bulbospinalis 184.15; p.282 - tegmentalis centralis 79.13; 98.30; 102.17;
- corticobulbaris 175.4 163.5; 164.3; p.232
- corticospinalis, see: tractus pyramidalis - tegmentalis medialis 163.4; 164.5; p.229
- corticovestibularis 112.6 - temporopontinus 170.16; 172.13
- cuneocerebellaris 161.14; p.224 - trigeminothalamicus dorsalis (Wallenberg), syn:
- fastigiobulbaris 112.6; p. 232 fasciculus tegmentalis dorsolateralis 98.28;
- frontopontinus 97.35; 101.31; 170.6; 132.2; 133.5; 136.4
172.15; 174.12 - trigeminothalamicus lateralis 132.18
- habenulointerpeduncularis, syn: fasciculus retro- - vestibulomesencephalicus 103.21; 104.21 ;
flexus (Meynert) 95.28; 97.33; 191.12; 139.11
194.11; 198B.5; p.308 - vestibulospinalis 104.21; p.256
- interstitiospinalis 138.2 - - lateralis 137.10; 139.23; p.256
- mamillotegmentalis 79.27; 191.11 ; 194.22; - - medialis 139.22; p.256
p.308 - vestibulothalamicus 139.9
- mamillothalamicus, syn: fasciculus mamillothala- trigeminal system p. 155
micus; bundle of Vicq d'Azyr 93.29; 191.8; trigonum habenulae 15.30
194.8; 195.9 trigonum lemnisci 14.6
- mesencephalicus nervi trigemini 99.17; 126.1 ; trigonum nervi hypoglossi 15.14
133.7; p.160 trigonum nervi vagi 15.15
- occipitopontinus 170.10 trigonum olfactorium 9.12
- olfactorius 9.9; 62.15; 196.2; 208.7; p.344 truncus cerebri 1.3; 2.4/7
- olivocerebellaris, see also fibrae olivocerebe1- truncus corporis callosi 12.1; 63.12; 84.6;
lares 160.14; 163.14; 164.19 214.9
- opticus 9.3; 14.20; 144.4; 145.8 tuber cinereum 9.5
- - radix lateralis 144.7; 145.10 tuber vermis 13.34; 16.4; 162.Tu
- - radix medialis 144.8; 145.9 tuberculum anterius thalami 14.19; 17.4
- pallidoreticularis, syn: fibrae pallidotegmentales tuberculum nuclei cuneati 14.14; 15.17
97.29 tuberculum nuclei gracilis 14.16; 15.18
- parietopontinus 170.9 tuberculum olfactorium 196.6; 197.39; 207.13;
- parietotemporopontinus (tractus parieto-occipito- p. 344
temporopontinus) 97.31; 101.29; 174.11 tuberculum trigeminale 96A + B
- pontospinalis 184.14; p. 282
- pyramidalis, syn: tractus corticospinalis 97.34; V-fibres p. 365
108.31; 170.8; 172.14; 174.13; p. 247 uncus 11.10-12
- - anterior 117.16; 121.30; 172.20; 174.15 upper horizontal line or plane (Kronlein), syn:
- - lateralis 117.13; 121.20; 172.19; 174.23 Obere Horizontale 39.0H
- reticulocerebellaris 114.19 uvula vermis 13.38; 18.35; 77.10; 162.Uv
436 Subject Index
vallecula cerebelli 54.19 vena

veins of the intervertebral foramen, see: venae - pedunculi cerebellaris superioris, syn: brachial
. intervertebrales vein 41.38; 45.31
velum medullare inferius 13.41; 18.4 - petrosa, syn: superior petrosal vein (Dandy)
velum medullare superius 13.23; 15.9; 18.1 41.35; 45.29
vena - - syn: inferior petrosal vein (Dandy; incon-
- anastomotica inferior (Labbe) 42.16 stant) 41.50; 43.23
- anastomotica superior (Trolard) 42.6 - pontis lateralis, syn: lateral pontine vein .. anterior
- angularis 42.34 cerebellar vein 41.37; 45.32; 48.19
- atrii lateralis, syn: vena atrii ventriculi lateralis - pontis mediana anterior, see: vena pontomesen-
lateralis .. atrial part of the vena terminalis inferi- cephalica anterior, median branch
or 45.27 - pontomesencephalica anterior 41.10
- atrii medialis, syn: vena atrii ventriculi lateralis - - median branch, syn: vena pontis mediana ante-
medialis 45.12 rior 43.38; 47.30
- azygos 57.17; 58.50; 59.5 - - peripeduncular branch, syn: vena sulci ponto-
- basalis (Rosenthali) 41.5; 42.14; 44.21 mesencephalica 47.26; 48.21
- cerebri interna 42.10; 45.11; 46.11 - precen tralis cere belli 41.40; 46.17
- cerebri magna (Galeni) 41.7; 45.14; 46.14 - radicularis anterior 57.6; 58.36
- cerebri media profunda, syn: deep middle cere- - radicularis posterior 57.5; 58.30
bral vein 41.3; 42.12; 45.18 - recessus lateralis ventriculi quarti 45.33
- cerebri media superficialis, syn: Sylvian vein - retromandibularis 42.44
42.11; 44.18 - sacralis lateralis 59.22
- cervicalis profunda 42.46; 57.4 - septi pellucidi anterior 45.5; 46.3
- choroidea inferior 42.13; 45.24 - spinalis anterior, syn: anteromedian medullary
- choroidea superior 42.9; 45.8 vein 41.51; 57.1; 58.26
- communicans posterior 47.28 - spinalis lateralis, syn: lateral medullary vein
- corporis callosi dorsalis 44.10 41.51
- diploica frontalis 53.13 - spinalis posterior, syn: posteromedian medullary
- diploica temporalis anterior 43.34 vein 41.51; 57.2; 58.14
- emissaria condylaris, syn: vena emissaria condy- - subclavia 57.9
loidea 41.54; 42.25 - subcostalis 57.19
- emissaria mastoidea 41.53; 42.24; 43.26 - sulci pontomedullaris 41.31
- emissaria occipitalis 42.21 - sulci pontomesencephalica, see: vena pontome-
- emissaria parietalis 42.4 sencephalica anterior, peripeduncular branch
- facialis 42.39 - terminalis, see: vena thalamostriata superior
- faciei profunda 42.38 - - inferior (occasional inferior section of the vena
- gyri olfactorii, syn: olfactory vein 44.14 thalamostriata superior) 45.23
- hemiazygos 57.18; 58.34 - - superior (superior section of the vena thala-
- hemiazygos accessoria 57.16 mostriata superior) 45.26
- iliaca interna 59.25 - thalami superior 45.10
- infraorbitalis 42.37 - thalamostriata inferior, see: venae thalamostria-
- intercostalis posterior (venae intercostales poster- tae inferiores
iores) 57.1,11, etc; 58.5 - thalamostriata superior, syn: vena terminalis
- - ramus spinalis 57.7; 58.8+35 42.8; 45.7; 46.4; 65.18; 84.7
- - ramus dorsalis 58.33 - ventricularis inferior 45.22
- intercosticalis superior dextra 57.14 - vermis inferior 41.43; 45.37; 46.25
- intercosticalis superior sinistra 57.15 - vermis superior 41.41; 45.36; 46.18
- intercosticalis suprema 57.8 - vertebralis 57.3
- interpeduncularis (with inferior thalamic - basivertebrales 58.47; 59.19
branches) 41.4; 44.23; 47.29 venae
- intervertebralis inferior, syn: suprapedicular vein - cerebelli 41; 45 ; 46
58.45; 59.12 - cerebri anteriores 41.2; 44.13
- intervertebralis superior, syn: infrapedicular vein - cerebri inferiores 42.19; 44.26
58.43; 59.11 - cerebri profundae, syn: subcortical veins 41;
- jugularis externa 42.45 45
- jugularis intern a 42.43 - cerebri superficiales, syn: cortical veins 44
- lumbalis ascendens 57.20; 59.7 - cerebri superiores 42.3; 44.2 + 3 + 6 + 7; 53.20
- mesencephalica lateralis 41.9; 45.30; 47.25 - diploicae 53.13
- occipitalis 42.48 - emissariae 42.4 + 21 + 24 + 25
- ophthalmica inferior 41.18; 42.35 - frontales 44.3
- ophthalmica superior 41.16; 42.33; 43.3 - gyrorum orbitalium, syn: fronto-orbital veins
- palatina, syn: vena palatina externa 42.40 44.15
Subject Index 437
venae ventriculus cornu posterius 21.11; 28.15; 71.16

- hemispherii (cere belli) inferiores 41.46; 45.39; - pars centralis 21.1; 28.2; 67.13
46.26 ventriculus quartus 3.5; 21.14; 70.28; 77.11
- hemispherii (cerebelli) superiores 41.39; 45.40 ventriculus tertius 3.3; 15.2; 21.5; 66.30
- insulares, syn: anterior, central and posterior vermis cerebelli 12.26; 16; 18; 162; 164.6;
insular veins 41.1; 45.17 p.221, 228
- intervertebrales, syn: veins of the intervertebral vestibular system p. 165
foramen 58.43 + 45; 59.11 + 12 vestibulocerebellar fibres
- lumbales (I, II, etc) 57.21 - primary - 138.13-3; 161.10-12; p.165
- maxillares 42.41 - secondary - 139.16/21-13; 161.11-12;
- medullae oblongatae (lateralis, anterolateralis et p.224
mediana) 49.17+18+19 vestibulocerebellum p. 232
- meningeae 53.23 vestibulomesencephalic projection p. 169
- meningeae mediae 42.30; 43.16 vestibulo-ocular reflex pathways 140; p. 169,
- mesencephalicae, syn: quadrigeminal vein; tectal 170
vein; superior median collicular vein (Duvernoy) vestibulo-olivary fibers 139.16/19-21
41.8; 47.23 vestibulothalamic fibers 139.9
- nuclei caudati 45.6; 46.2 villi arachnoideales 53.22
- occipitales 44.7 visceral afferent fibre (of dorsal root) 125.1
- parietales 44.6 visceral afferent systems p. 164
- orbitae 42.33 + 35 + 37; 43.3 visceral efferent fibre (of ventral root) 125.5
- pedunculares (some authors include the vena in- visual area Vl, see: area striata
terpeduncularis) 47.27 visual area V2 and V3, see: visual cortex, second-
- pontis, syn: pontine venous plexus 41.33; 48.20 ary - and: tertiary
- - transversae (superior et inferior) 41.36; visual association cortex, see: visual cortex, sec-
48.17 ondary - and: tertiary
- prefrontales 44.2 visual cortex, primary-, see: area striata
- spinales 41.51 visual cortex, secondary-, syn: area 18+area 19
- temporales mediales, syn: medial temporal cortial 5; 145.18+19; p.371
veins 45.25 visual cortex, tertiary p. 371
- temporales superficiales 42.5 + 42 visual field 145.1/3
- thalamostriatae inferiores, syn: inferior striate visual pathway p. 179
veins 44.17 visual reflexes p. 185
- trunci encephalici 47; 48; 49 visual system p. 179
ventral amygdalofugal pathway p. 324
ventral ascending serotoninergic pathway p. 206 Wernicke's speech area 216.W; 217.W; p.372
ventral horn, see: cornu anterius wing of the ambient cistern, see: ala cisterna am-
ventral raphespinal projection 183.14; p.281 bientis
ventral thalamus p.237
ventriculus lateralis 3.1 zonaincerta 66.31; 86.25; 93.7; 156.8;
- cornu anterius 21.2; 28.3; 63.14; 85.4 173.9; p. 154, 296, 313
- cornu inferius 21.8; 28.17; 66.41; 74.18 zona intercollicularis, see: nucleus intercollicularis
R. Nieuwenhuys, University ofNijmegen
Chemoarchitecture
of the Brain
1985.85 figures. X, 246 pages. Soft cover.
ISBN 3-540-15349-7
This book provides a concise and yet comprehensive survey

of the present state of "chemical neuroanatomy". It:
- sketches the development of the main concepts concerning
chemical transmission in the central nervous system
- discusses and illustrates 28 neuronal populations, each
containing a different neuromediator
- examines the cell groups and fiber systems containing
acetylcholine, the various monoamines, the amino acid
neurotransmitters and 18 different neuropeptides
- explores the relations between "classical" neuroanatomy
and "chemical" neuroanatomy
- indicates how, by a synthesis of these two disciplines, new
functional entities in the brain can be delineated
The volume also covers the most recent literature in the field,
and will prove useful to all those working in the neurological
sciences, both fundamental and applied.
From the reviews:

Chemoarchitecture of the Brain is an attempt to bring
It•••
together the fruit of classical research with what has been

learned from a somewhat different vantage point. In this it is
largely successful. To date, the topic has been approached
with varying success in multi-author books and limited review
articles, but this volume is a critical review of the literature
which covers most of the eNS from a uniform perspective .,.
On the whole the book provides an excellent summary of
contemporary views in chemical neuroanatomy and of their
impact on what has often been a very conservative discipline.
It should be required reading for students of neuroanatomy,
Springer-Verlag while workers in other fields will find it a helpful introduction
Berlin Heidelberg New York to a most complex field of research."
London Paris Tokyo HongKong Nature, 15. May 1986
Current Topics in Neuroendocrinology
Editors: D. Ganten and D. Pfaff
Volume 9 Volume 8
D. Ganten, University of Heidelberg; D. Pfaff, Rocke- D.Ganten, University of Heidelberg; D.PfatT, Rocke-
feller University, New York (Eds.) feller University, New York; K. Fuxe, Stockholm
(Eds.)
Stimulus-Secretion Coupling
in Neuroendocrine Systems Neuroendocrinology of
With contributions by I. M. Cooke, H. Duve,
Mood
D.K.Hartlin~ G. I. Hatton, W.1. Malaisse, 1988. 80 figures. VI, 335 pages. Hard cover.
R W. Newcomb, L. 1. Parry, A. Poulain, F. Strum- ISBN 3-540-17892-9
wasser, E. L. Stuenkel, A. 1. S. Summerlee,
D. T. Theodosis, A. Thorpe Contents: Principles for the Honnone Regulation of
Wiring Transmission and Volume Transmission in
1988. 69 figures. Approx. 290 pages. Hard cover. the Central Nervous System. - Clinical Studies with
ISBN 3-540-19043-0 Corticotropin-Releasing Hormone: Implications for
Contents: Cellular Reorganization in Neuroendo- Hypothalamic-Pituitary-Adrenal Dysfunction in
crine Secretion. - Stimulus-Secretion Coupling in the Depression and Related Disorders. - Biological
Oxytocin System. - Coupling of Electrical Activity Rhythms and Mood Disorders. - Recurrent Affective
and Hormone Release in Mammalian Neurosecre- Disorders: Lessons from Limbic Kindling. - The
tory Neurons. - The Bag Cell Neuroendocrine Mechanisms of Action of Antipsychotics and Antide-
System of Aplysia. - Electrophysiological Character- pressant Drugs. - Catecholamines and Mood:
istic ofPeptidergic Nerve Terminals Correlated with Neuroendocrine Aspects. - Serotonin and Mood:
Secretion. - Changes in Information Content with Neuroendocrine Aspects. - Cholinergic Mechanisms
Physiological History in Peptidergic Secretory in Mood: Neuroendocrine Aspects. - The Psycho-
Systems. - Insect Neuropeptides. - Stimulus-Secre- biology ofNeurotensin. - Cholecystokinin and
tion Coupling in the Pancreatic B Cell. - Subject Mood. - Opioid Peptides and Mood: Neuroendo-
Index. crine Aspects. - The Neuroendocrinology of Ano-
rexia Nervosa. - Effects of Peripheral Thyroid
The role of electrical signalling in the control of Hormones on the Central Nervous System: Rele-
endocrine secretions by the brain has been clear for vance to Disorders of Mood.
many years. Recently, the influences of hormones on
synthetic events in neuroendocrine cells have raised This volume deals with the various neurotransmitters
new questions concerning the peptides released from (amino acids, biogenic arnines, peptides) and their
such neurons. role in the control of behaviour and mood, both from
This volume concentrates on the relation between the experimental and from the clinical and pharma-
these two fields and asks how electrical potentials cological point of view.
facilitate secretion of substances from nerve cells The book provides a detailed discussion of how
which control endocrine events. While stimulus- biological rhythms relate to changes in mood, and
secretion coupling has been studied extensively in how the hypothalamus and pituitary hormones can
other physiological contexts, this is the first treatment be related to mood changes. Basic hints for diagnosis
of the phenomenon in an exclusively neuroendocrine are also considered.
setting.
Springer-Verlag Berlin Heidelberg New York London Paris Tokyo HongKong

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R.

Third Revised Edition

With 217 Figures

Springer-Verlag Berlin Heidelberg GmbH

JAN VOOGD, M.D., Ph.D.

CHRISTIAAN VAN HUIJZEN, F.M.A.A.

1978 First English Edition

Acknowledgement is made to the artists: to Mr. T. VAN GERWEN, who

Summer 1988 R. NIEUWENHUYS

Summer 1981 R.NIEUWENRUYS

Part I Gross Anatomy

Part II Vessels and Meninges

Part III Brain Slices

Part IV Microscopical Sections

Part V Functional Systems

The Trigeminal System . . . . . . . . . . . . . . 155

Connections of the Anterior Nucleus and the Lateral Dorsal Nucleus

Olfactory and Limbic Systems . . . . . . . . . . . 293

The Circuit of Papez . . . . . . . . . . . . . . . 341

SUbject Index 419

Purpose and Plan between the basic neuroanatomy required by

Since the Latin terminology has the obvious

External and Medial Views

Fig. 1. The central nervous system in situ (1 /6 x)

1 Polus occipitalis 5 Operculum fronto-parietale 9 Lobus frontalis

I Fissura longitudinalis cerebri

Fig. 6. The brain seen from above (1/1 x )

Fig. 7. Lateral view of the brain (1 /1 x )

Fig. 8. Dissection of the right cerebral hemisphere to display the insula (1 /1 x)

8 Bulbus olfactorius 18 Gyrus longus insulae

Fig. 11. Medial aspect of the right cerebral hemisphere (1 /1 x)

1 Truncus corporis callosi

19 Thberculum anterius thalami

1 Ventriculus lateralis 25 Taenia choroidea

9 Velum medu.llare superius

1 Culmen 7 Lobulus quadrangularis 13 Lobulus semilunaris inferior

Fig. 16. Dorsal view of the cerebellum (6/5 x; diagram: 1/1 x)

1 Taenia choroidea 24 Ventriculus lateralis

1 Velum medullare superius 12 Culmen 24 Fissura horizontalis

hemis pher i um cere b elli

Fig. 18. Ventral view of the cerebellum (6/5 x; diagram : 1/1 x)

Fissura mediana anterior

1 Ventriculus lateralis, pars centralis 9 Recessus suprapinealis

1 Lamina affixa 9 Corpus fornicis

1 Stria longitudinalis medialis 14 Gyrus cinguli

11 Fimbria hippocampi 24 Gyrus dentatus retrocommJ urahs

9 Nucleus ventralis lateralis

I Nucleus anterior thalami

1 Corona radiata (some isolated fibres)

Fig. 27. The basal ganglia in medial view (6/5 x)

8 Capsula interna, pars sublentifonnis

1 Capsula interna, crus anterius

7 Ventriculus lateralis, pars centralis

Arteries of the Brain

Veins of the Brain

Vessels of the Brain Stem

Meninges, Cisterns and Liquor System

Vessels and Meninges of the Spinal Cord

1 Sulcus centralis 20 A. cerebri posterior, pars precommunicalis

17 Sulcus lemporaJis superior

1 Cis!. laminae terminalis

1 Nucleus caudatus 20 Nucleus dorsomedialis thalami

1 Calvaria (inner oulline)

36 A. vertebrali • pars intracranialis