Table Of ContentAdvances in Anatomy
Embryology and Cell Biology
Vol. 107
Editors
F. Beck, Leicester W. Hild, Galveston
W. Kriz, Heidelberg R. Ortmann, K6ln
J.E. Pauly, Little Rock T.H. Schiebler, Wiirzburg
H.J. ten Donkelaar G.C. Bangma
H.A. Barbas-Henry
R de Boer-van Huizen J.G. Wolters
The Brain Stem in a Lizard,
Varanus exanthematicus
With 69 Figures
Springer-Verlag
Berlin Heidelberg New York
London Paris Tokyo
Dr. Hendrik Jan ten Donkelaar
Roelie de Boer-van Huizen
Dr. Jan G. Wolters
Department of Anatomy and Embryology
Faculty of Medicine, University of Nijmegen
P.O.Box 9101, NL-6500 HB Nijmegen
Dr. Gesineke C. Bangma
Hubrecht Laboratory
Netherlands Institute for Developmental Biology
NL-Utrecht
Dr. Heleen A. Barbas-Henry
Department of Anatomy, Vrije Universiteit
NL-Amsterdam
ISBN -13: 978-3-540-t 7948-t e-ISBN-13: 978-3-642-72763-4
DOl: 10.1007/978-3-642-72763-4
Library of Congress Cataloging-in-Publication Data
The Brain stem in a lizard, Varanus exanthematicus.
(Ad~ances in anatomy, embryology, and cell biology; vol. 107) Bibliography: p.
Includes index. 1. Varanus exanthematicus - Anatomy. 2. Brain stem -
Anatomy. 3. Reptiles-Anatomy. I. Donkelaar, H.J. ten (Hendrik Jan), 1946-.
II. Series: Advances in anatomy embryology, and cell biology; v. 103.
QL80l.E67 vol. 107 574.4 s 87-16347 [QL666.L29] [597.95]
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© Springer-Verlag Berlin Heidelberg 1987
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Contents
1 Introduction 1
2 Materials and Techniques 3
2.1 Cytoarchi tectonic Analysis 3
2.2 Immunohistochemical Procedures 4
2.3 Tract-Tracing Studies 4
2.3.1 Anterograde Degeneration Studies 4
2.3.2 Horseradish Peroxidase Retrograde and
Anterograde Transport Studies 5
2.3.3 Retrograde Fluorescent Tracer Studies 5
2.3.4 Anterograde [3 H] Leucine Tracing Studies 6
3 Gross Anatomy 7
3.1 The Brain Stem 7
3.2 The Cranial Nerves 8
4 Spinal Projections to the Brain Stem 19
4.1 The Dorsal Column Nucleus and its Afferent and
Efferent Connections 19
4.2 Fiber Systems Ascending in the Lateral Funiculus 21
4.2.1 The Spinal Lemniscus . . . . . 21
4.2.1.1 Spinorhombencephalic Projections 21
4.2.1.2 Spinomesencephalic Projections 23
4.2.1.3 Spinothalamic Projections 24
4.2.2 Spinocerebellar Projections . . 26
5 Organization and Connections of the Sensory
Trigeminal Nuclei 28
5.1 Cytoarchitecture 28
5.2 Primary Afferents 29
5.3 Central Trigeminal Projections 32
6 The Solitary Tract and Related Nuclei 36
6.1 Cytoarchitecture: Viscerosensory Nuclei 36
6.2 Primary Afferents . . . . . . . . 37
6.3 Central Visceral and Taste Pathways . . 41
V
7 Cranial Nerve Motor Nuclei 43
7.1 Somatomotor Nuclei . 43
7.2 Branchiomotor Nuclei 44
7.3 Visceromotor Nuclei . 45
8 The Vestibular Nuclear Complex and Related
Structures 47
8.1 Cytoarchitecture 47
8.2 Primary Afferents 48
8.3 Central Vestibular Projections 49
8.3.1 The Fasciculus Longitudinalis Medialis 49
8.3.2 Vestibulospinal Pathways 54
9 Acoustic Projections 56
9.1 Primary Acoustic Projections and Nuclei 56
9.2 The Superior Olive 57
9.3 The Torus Semicircularis . . 59
10 Visual Input to the Brain Stem 61
10.1 The Tectum Mesencephali 61
10.1.1 Tectal Lamination 61
10.1.2 Tectal Afferents . . 63
10.1.2.1 Visual Input 63
10.1.2.2 Nonvisual Afferents 65
10.1.3 Tectal Efferents . . 68
10.1.3.1 Ascending Tectal Efferents 68
10.1.3.2 Descending Tectal Efferents 69
10.2 The Nucleus of the Basal Optic Root 74
10.3 The Isthmic Nuclei ....... . 74
11 Forebrain Projections to the Brain Stem 76
11.1 Notes on the Organization of the Reptilian
Forebrain .............. 76
11.2 Basal Ganglia Projections to the Brain Stem 77
11.2.1 The Substantia Nigra and Ventral Tegmental Area 82
11.3 Diencephalic Projections to the Brain Stem. 87
11.3.1 Thalamic Projections to the Brain Stem 87
11.3.2 Hypothalamic Projections to the Brain Stem 87
11.4 The Habenulointerpeduncular Tract 88
11.4.1 The Nucleus Interpeduncularis 88
12 Cerebellar Connections . . . . 89
12.1 Notes on the Cerebellar Cortex 89
12.2 The Cerebellar Nuclei . . . . 90
12.3 Afferent Connections of the Cerebellum 91
12.3.1 Spinal Afferents . . . . . . . . . . . 91
VI
12.3.2 Rhombencephalic Afferents . . . . 91
12.3.2.1 Vestibular Afferents . . . . . . . 92
12.3.2.2 The Nucleus Prepositus Hypoglossi 92
12.3.2.3 The Inferior Olive . . . . . . . 92
12.3.2.4 Other Rhombencephalic Afferents 94
12.3.3 Mesencephalic Afferents . . . . 95
12.3.4 Prosencephalic Afferents . . . . 95
12.4 Efferent Connections of the Cerebellum 96
12.4.1 Corticonuclear Projections . . . . 96
12.4.2 Projections of the Cerebellar Nuclei 99
12.4.3 The Red Nucleus 103
13 The Reticular Formation 106
13.1 Cytoarchi tecture 106
13.2 Monoaminergic and Peptidergic Components 109
13.2.1 Distribution of Catecholamine Neurons in the
Brain Stem .............. . 109
13.2.2 Distribution of Serotonin-Containing Neurons in
the Brain Stem . . . . 111
13.2.3 Peptidergic Components . . . . . . . . . . . 113
13.3 Ascending Projections ........... 117
13.3.1 Ascending Projections of the Brain Stem Reticular
Formation . . . . . . . . . . . . . 117
13.3.2 Ascending Monoaminergic Systems in the
Reptilian Reticular Formation 121
13.4 Descending Projections . . . . . . . . 123
13.4.1 Reticulospinal Projections ..... . 123
13.4.2 Descending Monoaminergic Projections to the
Spinal Cord ..... 129
13.4.2.1 Raphespinal Projections 129
14 Concluding Remarks . . 133
14.1 Ascending Projections 133
14.2 Descending Projections to the Brain Stem 134
14.3 Descending Projections from the Brain Stem 137
14.4 Cerebellar Connections 140
14.4.1 Cerebellar Afferents . 140
14.4.2 Cerebellar Efferents 141
14.5 Research Perspectives 144
Acknowledgments 145
References . 146
Subject Index 166
VII
Abbreviations Used in the Figures *
Acc nucleus accumbens
ADVR anterior dorsal ventricular ridge
cl caudolateral (somatosensory) area
m medial (acoustic) area
rl rostrolateral (visual) area
Alh area lateralis hypothalami
Am amygdala
Amb nucleus ambiguus
AmbX vagal part of the nucleus ambiguus
bc brachium conjunctivum
Bor nucleus of the basal optic root
C caudal
ca commissura anterior
cb cerebellum
cc corpus callosum
cereb cerebellum
Cerl nucleus cerebellaris lateralis
Cerm nucleus cerebellaris medialis
cho chiasma opticum
Cm corpus mamillare
Coa nucleus cochlearis angularis
Codm nucleus cochlearis dorsalis magnocellularis
Colm nucleus cochlearis laminaris
cosp corticospinal tract
cp commissura posterior
cpa commissura pallii anterior
CPC caudate-putamen complex
Cs colliculus superior
Cun nucleus cuneatus
Cx cortex telencephali
CXe cingulate cortex
CXr frontal cortex
CXsm somatomotor cortex
D nucleus dentatus
dh dorsal horn
DI nucleus dorsolateralis thalami
* Abbreviations with roman numerals are grouped separately at the end
of the entries for the respective letters or at the very end in the case of
roman numerals not preceded by a letter.
IX
Dm nucleus dorsomedialis thalami
dp dorsal pallidum
dso decussatio supraoptica
Enp nucleus entopeduncularis posterior
Ep nucleus entopeduncularis
EW Edinger-Westphal nucleus
FB Fast Blue
fd funiculus dorsalis
Fl nucleus funiculi lateralis
fl funiculus lateralis
flm fasciculus longitudinalis medialis
fpd fasciculus predorsalis
fr fasciculus retroflexus
FRmes mesencephalic part of the reticular formation
fu fasciculus uncinatus
Fun nucleus funiculi dorsalis
fv funiculus ventralis
Gc griseum centrale
gl granular layer of the cerebellum
Glv nucleus geniculatus lateralis pars ventralis
GP globus pallid us
Gp nucleus geniculatus pretectalis
Gr nucleus gracilis
Hb habenula
hip hippocampus
hyp hypothalamus
Ico nucleus intercollicularis
Iflm nucleus interstitialis of the flm
Ila intra laminar nuclei
Ip nucleus interpeduncularis
Ipd nucleus interpeduncularis pars dorsalis
Ipv nucleus interpeduncularis pars ventralis
Ism nucleus isthmi pars magnocellularis
Isp nucleus isthmi pars parvocellularis
lZ intermediate zone
L lateral
Lc locus coeruleus
Le nucleus lentiformis thalami, pars extensa
Ifb lateral forebrain bundle
Li nucleus linearis intermedius
Ll nucleus lemnisci lateralis
11 lemniscus lateralis
Lm nucleus lentiformis mesencephali
1m lemniscus medialis
Lp nucleus lentiformis thalami, pars plicata
Lr nucleus linearis rostralis
Is lemniscus spinalis
M medial
Md nucleus mediodorsalis
x
Med nucleus medialis
mfb medial forebrain bundle
ml molecular layer of the cerebellum
Ncp nucleus of the commissura posterior
Ndb nucleus of the diagonal band
Nflm nucleus of the flm
Nlot nucleus of the lateral olfactory tract
NY Nuclear Yellow
nl nervus olfactorius
nIl nervus opticus
nlIl nervus oculomotorius
nlV nervus trochlearis
nV nervus trigeminus
nVI nervus abducens
nVII nervus facialis
nVIII nervus vestibulocochlearis (octavus)
nVIIla anterior root of the nervus vestibulocochlearis
nVIIlp posterior root of the nervus vestibulocochlearis
nIX nervus glossopharyngeus
nX nervus vagus
nXII nervus hypoglossus
oc olivocerebellar fibers
Oli oliva inferior
Ols oliva superior
Pal pallidum
Pb parab rachial region
Pd nucleus posterodorsalis
Ph nucleus periventricularis hypothalami
Phg nucleus prepositus hypoglossi
PI Purkyne cell layer
Prmc nucleus profundus mesencephali, pars caudalis
Prmr nucleus profundus mesencephali, pars rostralis
Pt nucleus pretectalis
Pv nucleus paraventricularis
Pvo paraventricular organ
R rostral
Rai nucleus raphes inferior
Rai nucleus raphes inferior, magnocellular part
m
Raip nucleus raphes inferior, "pallidal" part
Ram nucleus raphes magnus
Rap nucleus raphes pallidus
Ras nucleus raphes superior
Rasl nucleus raphes superior, pars lateralis
Rasm nucleus raphes superior, pars medialis
rasp raphespinal projections
Rd nucleus raphes dorsalis
resp reticulospinal projections
Rf reticular formation
Ri nucleus reticularis inferior
XI
Ril nucleus reticularis inferior, pars lateralis
Ris nucleus reticularis isthmi
Riv nucleus reticularis inferior, pars ventralis
Rm nucleus reticularis medius
rm ramus muscularis n. trigemini
Rml nucleus reticularis medius, pars lateralis
Ro nucleus raphes obscurus
Rot nucleus rotundus
Rpa nucleus raphes paUidus
Rpo nucleus raphes pontis
Rs nucleus reticularis superior
Rsl nucleus reticularis superior, pars lateralis
Rub nucleus ruber
rusp tractus rubrospinalis
rVme ramus mesencephalicus nervi trigemini
sac stratum album centrale
Sc subcoeruleus area
sept septum
sgc stratum griseum centrale
sgfs stratum griseum et fibrosum superficiale
sgp stratum griseum periventriculare
Si nucleus salivatorius inferior
Sn substantia nigra
Sne substantia nigra, pars compacta
Sn substantia nigra, pars reticulata
r
So subcommissural organ
so stratum opticum
Sol nucleus tractus solitarii
spcd tractus spinocerebellaris dorsalis
spcv tractus spinocerebellaris ventralis
Ss nucleus salivatorius superior
Sth nucleus subthalamicus
Str striatum
tb tractus tectobulbaris
tbd tractus tectobulbaris dorsalis (neuron)
tbv tractus tectobulbaris ventralis
thd dorsal thalamus
tm tectum mesencephali
to tractus opticus
tol tractus olfactorius lateralis
Torc torus semicircularis, nucleus centralis
TorI torus semicircularis, nucleus laminaris
Tpp nucleus tegmenti pedunculopontinus
tr opt tractus opticus
tsol tractus solitarius
tth tractus tectothalamicus
Tub olf tuberculum olfactorium
tV ds tractus descendens nervi trigemini
Va nucleus ventralis anterior
XII
