Sensory Mechanisms of the Spinal Cord: Volume 1 Primary Afferent Neurons and the Spinal Dorsal Horn

603 Pages·2004·16.352 MB·English

by  William D. Willis Jr., Richard E. Coggeshall (auth.)

Sensory Mechanisms
of the Spinal Cord
Third Edition
Volume 1
Primary Afferent Neurons
and the Spinal Dorsal Horn
Sensory Mechanisms
of the Spinal Cord
Third Edition
Volume 1
Primary Afferent Neurons
and the Spinal Dorsal Horn
William D. Willis, Jr.
and
Richard E. Coggeshall
The University of Texas Medica/ Branch
Ca/veston, Texas
Springer-Science+Business Media, LLC
ISBN 978-1-4613-4894-8 ISBN 978-1-4615-0037-7 (eBook)
DOI 10.1007/978-1-4615-0037-7
©2004 Springer Science+Business Media New Yark
Originally published by Kluwer Academic/Plenum Publishers, New York in 2004
Softcover reprint of the hardcover 3rd edition 2004
\098765432
A C.I.P. record for this book is available from the Library of Congress
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No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any
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PREFACE
The third edition of this monograph continues to have the goal of providing an overview
of current thought about the spinal cord mechanisms that are responsible for sensory
processing. We hope that the book is of value to both basic and clinical neuroscientists.
Several changes have been made in the presentation, as well as additions because of
the research advances that have been made during the past decade. Chapters 3 and 4 in the
previous edition have been subdivided, and now the morphology of primary afferent neu
rons of the dorsal root ganglia is described in Chapter 3 and the chemical neuroanatomy
of these neurons in Chapter 4. The description of the dorsal hom in the previous Chapter 4
is now included in Chapter 5, and the chemical neuroanatomy of the dorsal hom in
Chapter 6. Furthermore, discussions of the descending control systems have now been
consolidated at the end of Chapter 12.
The authors would like to express their appreciation for the help provided by several
individuals. R.E.C. wishes to acknowledge the many things he learned about primary
afferent neurons from conversations with Dr S. N. Lawson. He also thanks Lyn Shilling
for her assistance with the typing. WDW thanks Dr Nada Lawand for her critical reading
of parts of the manuscript, Rosaline Leigh for help with the manuscript, and Griselda
Gonzales for preparing the illustrations.
William D. Willis and Richard E. Coggeshall
CONTENTS
Volume 1
Chapter 1: Introduction 1
Sensory Modalities and Channels 1
Specificity versus Pattern Theories of Sensation 2
Historical Perspective 2
Specificity Theory 2
Pattern Theory 4
Gate Theory of Pain 5
Supporting Evidence 7
Molecular Basis of Specificity of Primary Afferent Neurons 7
Sinclair's Challenge and Evidence for Specificity from
Microneurography 9
Sensory Role of Single Tactile Afferents in Human Subjects 9
Criticism of Microneurography Results 11
Rebuttal of this Criticism 12
Microneurography and Pain 14
Somatosensory System Dualities 14
Epicritic versus Protopathic Sensations 14
Lemniscal versus Non-Iemniscal Systems 15
Large- versus Small-Fiber Systems 15
Conclusions 16
Chapter 2: Sensory Receptors and Peripheral Nerves 19
Composition of Peripheral Nerves 19
Sensory Receptors 22
Cutaneous Receptors 31
Cutaneous Mechanoreceptors 32
Mechanoreceptors in the Glabrous Skin of the Primate Hand 32
Cutaneous Displacement and Velocity Detectors 34
SA I Receptors 34
SA II Receptors 37
Cutaneous Velocity Detectors 39
FA I receptors 39
Hair Follicle Receptors 41
Field Receptors 43
C Mechanoreceptors 44
Cutaneous Transient Detectors 44
vii
viii Contents
FA II Receptors 45
G Hair Follicle Receptors 46
1
Sinus Hairs 47
The Role of Growth Factors in the Regulation of Mechanoreceptors 47
SA I (Merkel Cell) Endings 47
FA I (Meissner Corpuscle) Receptors 48
FA II (Pacinian Corpuscle) Receptors 48
Hair Follicle Receptors 48
Transduction in Mechanoreceptors 48
Cutaneous Nociceptors 50
A3 Nociceptors 50
C Nociceptors 53
Transduction in Nociceptors 57
Sensitization of Nociceptors 58
Neurogenic Inflammation 63
Peptidergic and Non-Peptidergic Nociceptors 63
Nociceptors and Neurogenic Inflammation 63
Interaction of Growth Factors with Nociceptors 64
Inhibitory Interactions with Peripheral Cutaneous Nociceptor Endings 66
Cutaneous Chemoreceptors Possibly Responsible for Itch 67
Cutaneous Thermoreceptors 67
Cold Receptors 68
Warm Receptors 69
Transduction in Thermoreceptors 70
Muscle Receptors 70
Stretch Receptors 70
Muscle Spindles 71
Golgi Tendon Organs 72
Pressure-Pain Endings and Muscle Nociception 73
Group III Muscle Afferents 73
Group IV Muscle Afferents 74
Sensitization of High Threshold Muscle Afferents 74
Joint Receptors 75
Joint Mechanoreceptors 76
Slowly Adapting Joint Receptors 76
Rapidly Adapting Joint Receptors 77
Joint Nociceptors 77
Sensitization of Joint Nociceptors 77
Peptide Content of Fine Joint Afferents 78
Joint Inflammation 78
Role of Dorsal Root Reflexes in the Neurogenic
Inflammation of Arthritis 79
Visceral Receptors 80
Visceral Mechanoreceptors 81
Visceral N ociceptors 82
Peptides in Visceral Afferents 85
Growth Factors and Visceral Afferents 85
Contents ix
Sensitization of Visceral Afferents 85
Visceral Thermoreceptors 86
Conclusions 87
Chapter 3: Dorsal Root Ganglion Cells and their Processes 91
Dorsal Root Ganglion Cell Bodies 91
DRG Cell Classifications 92
Cytologic and Size Classifications 92
Myelination 93
Axon Conduction Velocity 93
Action Potentials and After Polarizations 94
Classifications by Chemical Neuroanatomical Criteria 94
Topographic Organization 95
Innervation of Ganglion Cell Bodies, Pericellular Baskets 96
Dorsal Root Ganglion Cell Processes 98
Dermatomes 98
Branching 98
Ventral Root Afferent Fibers 98
Spinal Cord 99
Blind Fibers 99
Looping Fibers 99
Pial Fibers 100
Function 100
Segregation of Dorsal Root Fibers 100
Conclusions 101
Chapter 4: Chemical Anatomy of Dorsal Root Ganglion Cells 103
Amines 103
Catecholamines (CAs) 103
Histamine 103
Serotonin 103
Amino Acids 104
Glutamate 104
Aspartate 104
GABA 104
Glycine 105
Channels 105
Calcium Channels 105
Sodium Channels 105
Voltage-Gated Sodium Channels 105
Cell Size 106
Co-localizations 107
13 Subunits 107
Development 107
Inflammation 107
Peripheral Axotomy 108
Nerve Growth Factor (NGF) 108
x Contents
Peripheral Sensory Axons 108
Non-voltage Gated Sodium Channels 108
Enzymes 109
Alkaline Phosphatase 109
Carbonic Anhydrase (CA) 109
Cholinergic Enzymes 110
ChAT 110
ACHE 110
Cytochrome Oxidase 110
Fluoride-resistant Acid Phosphatase (FRAP) 111
Monoamine Oxidase 112
Nitric Oxide (NO), Nitric Oxide Synthase (NOS) and NADPHd 112
Ca+ + Calmodulin Kinase II 113
Other Kinases 113
Hormones 113
Corticotrophin Releasing Factor (CRF) 113
Insulin 113
Peptides 113
Angiotensin 113
Atrial and Brain Natriuretic Peptide 114
Bombesin 114
Cholycystokinin (CCK) 114
Calcitonin Gene-related Peptide (CGRP) 114
Classification 116
Age 116
Development 116
Co-localizations 116
SP 116
GAL 117
SOM 117
CCK 117
VIP 117
Bombesin, ENK, DYN, VIP, NT, and NPY 117
FRAP 117
Cysteine Proteases 117
Amino Acids 117
IB4 117
Peripherin 118
BDNF 118
GDNF 118
TrkA 118
P75 118
Experimental Manipulations or Disease 118
Dorsal Rhizotomy 118
Peripheral Nerve Lesions 118
Inflammation 119
Diabetes 119
Hypoxia 120
Contents xi
Hereditary Sensory Neuropathy 120
Capsaicin 120
Axonal Transport Blockers 120
NGF Treatment 120
Female Steroid Hormones 120
Sympathectomy 121
Dorsal Roots 121
Periphery 121
Somatic Innervation 121
Visceral Innervation 122
Endothelin (ET) 122
Galanin 122
Nerve Lesions 123
Inflammation 124
Resiniferatoxin 124
Axonal Transport Blockers 124
Co-localizations 124
CGRP 124
SP 124
VIP 124
NPY 124
5-HT 124
Glucocorticoid Receptors 125
Dorsal Roots 125
Neuropeptide FF (NPFF) 125
Neuropeptide Y (NPY), Polypeptide Y (PPY), and Peptide YY 125
Peripheral Axotomy or Compression 125
Trophic Factors 125
Development 126
Co-localizations 126
Peripheral Innervation 126
Neurotensin (NT) 126
Opioids 126
Dynorphin (DYN) 126
Endomorphin 127
Endorphin 127
Enkephalin 127
Nociceptin 128
Oxytocin and Vasopressin 128
Secretoneurin and the Chromagranins 128
Somatostatin 128
Co-localizations 129
GDNF and Peripheral Axotomy 129
Development 129
Periphery 129
Substance P (SP) 130
Segmental Levels 130
Age and Development 130
xii Contents
Co-localizations 130
Tachykinins 130
CGRP 130
GAL 131
SOM 131
CCK 131
ENKand DYN 131
Bombesin, VIP, NT, and NPY 131
FRAP 131
NOS 131
Glutamate 131
IB-4 131
200 KD Neurofilament Protein 132
Peripherin 132
GDNF 132
Opioid Receptors 132
Experimental Manipulations 132
Dorsal Rhizotomy 132
Peripheral Nerve Lesions 132
Inflammation 133
Trophic Factors 134
Adrenalectomy 134
Resiniferatoxin 134
Periphery 134
Somatic Innervation 134
Visceral Innervation 135
VIP 135
Proteins 136
Calcium Binding Proteins 136
Glutamate Transporters 136
Reg-2 137
Bone Matrix Proteins 137
l3-arrestin 137
Receptors 137
Aminergic Receptors 137
Adrenergic Receptors 137
Dopamine Receptors 138
Histamine Receptors 138
5-HT Receptors 138
5-HT Receptors 138
IA
5-HTJB Receptors 138
5-HT Ie Receptors 138
5-HT 1D Receptors 139
5-HT IE Receptors 139
5-HT Receptors 139
IF
5-HT Receptors 139
2
5-HT3 Receptors 139
5-HT4, 5-HTs, and 5-HT7 Receptors 139