THE PERIPHERAL NERVOUS SYSTEM

Transcription

1 THE PERIPHERAL NERVOUS SYSTEM

2 THE PERIPHERAL NERVOUS SYSTEM Edited by John I. Hubbard Department 0/ Physiology University 0/ Otago Dunedin, New Zealand PLENUM PRESS NEW YORK AND LONDON

3 Library of Congress Cataloging in Publication Data Hubbard, John I The peripheral nervous system. Includes bibliographies. 1. Nerves, Peripheral. I. Title. [DNLM: 1. Peripheral nerves. W1500 H875p 1974] QP365.5.H8 612' Plenum Press, New York Softcover reprint of the hardcover 1 st edition 1974 A Division of Plenum Publishing Corporation 227 West 17th Street, New York, N.Y United Kingdom edition published by Plenum Press, London A Division of Plenum Publishing Company, Ltd. 4a Lower John Street, London WIR 3PD, England ISBN-13: e-isbn-13: DOl: /

4 Contributors Eric A. Barnard Departments of Biochemistry and Biochemical Pharmacology State University of New York Buffalo, New York, USA Christopher Bell Department of Zoology University of Melbourne Victoria, Australia T. J. Biscoe Department of Physiology University of Bristol Bristol, England J. G. Blackman Department of Pharmacology University of Otago Medical School Dunedin, New Zealand Geoffrey Burnstock Department of Zoology University of Melbourne Victoria, Australia Mohyee E. Eldefrawi Section of Neurobiology and Behavior Cornell University Ithaca, New York, USA Lloyd Guth Laboratory of Neurochemistry National Institute of Neurological Diseases and Stroke Public Health Service, Department of Health, Education and Welfare Bethesda, Maryland, USA Arthur Hess Department of Anatomy Rutgers Medical School New Brunswick, New Jersey, USA John I. Hubbard Department of Physiology, Medical School University of Otago Dunedin, New Zealand Carlton C. Hunt Department of Physiology and Biophysics Washington University School of Medicine St. LOllis, Missouri, USA Ainsley Iggo Department of Veterinary Physiology University of Edinburgh Edinburgh, Scotland David M. Jacobowitz Laboratory of Clinical Science National Institute of Mental Health Bethesda, Maryland, USA Margaret R. Matthews Department of Human Anatomy Oxford University Oxford, England P. B. C. Matthews University Laboratory of Physiology Oxford University Oxford, England Syogoro Nishi Neurophysiology Laboratory, Department of Pharmacology Loyola University Medical Center Maywood, Illinois, USA Denis Noble Fellow of Balliol College, Oxford University University Lecturer in Physiology Oxford, England Sidney Ochs Department of Physiology Indiana University Medical Center Indianapolis, Indiana, USA Henry def. Webster Head, Section on Cellular Neuropathology Laboratory of Neuropathology and Neuroanatomical Sciences National Institute of Neurological Diseases and Stroke Bethesda, Maryland, USA J. G. Widdicombe Department of Physiology St. George's Hospital Medical School London, England

5 Preface The peripheral nervous system is usually defined as the cranial nerves, spinal nerves, and peripheral ganglia which lie outside the brain and spinal cord. To describe the structure and function of this system in one book may have been possible last century. Today, only a judicious selection is possible. It may be fairly claimed that the title of this book is not misleading, for in keeping the text within bounds only accounts of olfaction, vision, audition, and vestibular function have been omitted, and as popularly understood these topics fall into the category of special senses. This book contains a comprehensive treatment of the structure and function of peripheral nerves (including axoplasmic flow and trophic functions); junctional regions in the autonomic and somatic divisions of the peripheral nervous system; receptors in skin, tongue, and deeper tissues; and the integrative role of ganglia. It is thus a handbook of the peripheral nervous system as it is usually understood for teaching purposes. The convenience of having this material inside one set of covers is already proven, for my colleagues were borrowing parts of the text even while the book was in manuscript. It is my belief that lecturers will find here the information they need, while graduate students will be able to get a sound yet easily read account of results of research in their area. JOHN 1. HUBBARD vii

6 SECTION I-PERIPHERAL NERVE Chapter 1 Peripheral Nerve Structure Henry def. Webster 1. Introduction. 2. Histology and Development 3. The Axon 3.1. Filaments and Microtubules 3.2. Other Organelles and the Axolemma 4. Sheaths ofaxons 4.1. Schwann Cells Myelin 4.3. Function of Schwann Cells and Their Myelin Sheaths 4.4. Connective Tissue Sheaths. 5. References Chapter 2 The Nerve Impulse Denis Noble 1. Introduction. 2. Passive Electrical Properties 3. Voltage-Clamp Analysis of the Ionic Current 4. Momentary Current-Voltage Relations 5. The Threshold Conditions for Excitation 6. Factors Determining Conduction Velocity 7. References ix

7 x Chapter 3 Axoplasmic Transport-Energy Metabolism and Mechanism Sidney Ochs 1. Introduction. 2. Fast Axoplasmic Transport 2.1. Characterization 2.2. Mechanism and Energy Supply 2.3. Transport and Membrane Function 3. Slow Axoplasmic Transport 3.1. Characterization 3.2. Mechanism 4. References SECTION lia-junctlonal TRANSMISSION-STRUCTURE Chapter 4 Neuromuscular Junctions and Electric Organs Arthur Hess 1. Introduction. 2. The Typical Neuromuscular Junction Distribution and Location of Nerve Terminals 2.2. The Axon 2.3. The Synaptic Space Post junctional Muscle Fiber 3. Variations of Motor End Plates 3.1. Variations from Class to Class 3.2. Endings on Slow-Twitch and Rapid-Twitch Fibers 3.3. Endings on Slow Tonic Muscle Fibers 4. Electric Organs 4.1. Electrocytes 4.2. Innervation and Ultrastructure 5. References Chapter 5 The Peripheral Autonomic System David M. Jacobowitz 1. Anatomical Considerations: Sympathetic and Parasympathetic Divisions 87 87

8 2. Morphological Observations 2.1. Preganglionic Neurons 2.2. Postganglionic Neurons 2.3. Adrenal and Extra-Adrenal Chromaffin Cells 3. References xi Chapter 6 Ultrastructure of Ganglionic Junctions Margaret R. Matthews 1. General Considerations 2. Sympathetic Ganglia 2. I. Amphibia 2.2. Reptiles 2.3. Mammals 2.4. Some Effects of Different Fixatives 3. Parasympathetic Ganglia 3.1. Ciliary Ganglion 3.2. Otic Ganglion 3.3. Ganglia of the Enteric Plexuses 3.4. Cardiac Ganglion Cells 4. Summary and Comment 5. References SECTION lib-junctional TRANSMISSION-FUNCTION Chapter 7(i) Neuromuscular Transmission-Presynaptic Factors John l. Hubbard 1. Synthesis, Storage, and Release of Acetylcholine 1.1. Synthesis of ACh 1.2. Storage and Release. 2. The Acceleration of Release by Nerve Impulses 2.1. The Role of the Nerve Impulse The Role of Ca After-Effects of Depolarization-Secretion Coupling 3. References

9 xii Chapter 7 (U) Neuromuscular Transmission-The Transmitter-Receptor Combination 181 M ohyee E. Eldefrawi 1. Introduction Molecular Basis of Chemoelectric Transduction Pharmacology Chemical Nature of the Acetylcholine Receptor Desensitization References 196 Chapter 7 (iii) Neuromuscular Transmission-Enzymatic Destruction of Acetylcholine 201 Eric A. Barnard 1. Location and Measurement of Cholinesterases at the Junction Histochemical Staining Microchemical Methods Assay of External AChE Radioautographic Methods Amounts and Types of Cholinesterase at the Junctions Requirement for AChE in Impulse Transmission Relation of AChE to ACh-Receptors Quantitative Relation of AChE to ACh at the End Plate References 219 Chapter 8 Ganglionic Transmission 225 Syogoro Nishi 1. Introduction Response of Autonomic Ganglia to Preganglionic Volleys Response of Normal Ganglia Response of Curarized Ganglia Slow Ganglionic Responses and Afterdischarges Electrical Constants of Ganglion Cell Membrane Action Potentials of Single Ganglion Cells Response to Antidromic Stimulation Response to Direct Intracellular Stimulation Response to Orthodromic Stimulation Ionic Requirement for Generation of Action Potential 238

10 xiii 5. Nature and Electrogenesis of Postsynaptic Potentials The" Fast" Excitatory Postsynaptic Potential The" Slow" Excitatory Postsynaptic Potential The" Late Slow" Excitatory Postsynaptic Potential The" Slow" Inhibitory Postsynaptic Potential Cholinergic and Adrenergic Receptors at Preganglionic Nerve Terminals Cholinergic Receptor Site Adrenergic Receptor Site References 250 Chapter 9 Function of Autonomic Ganglia 257 J. G. Blackman 1. Introduction Ganglia as Coordinating Centers The Relay Hypothesis of Ganglionic Function Development of a Stochastic Hypothesis Experimental Evidence Observed Patterns of Innervation Ganglionic Activity and Factors Influencing It Relative Autonomy of Ganglia Conclusions References 274 Chapter 10 Peripheral Autonomic Transmission 277 Geoffrey Burnstock and Christopher Bell 1. Introduction Definition of the Autonomic Neuromuscular Junction Relation of Nerve Fibers to Muscle Effector Bundles Relation of Nerve Fibers to Individual Smooth Muscle Cells Adrenergic Transmission Introduction Structure of Adrenergic Neurons and Storage of Noradrenaline Electrophysiology of Adrenergic Transmission Ionic Basis of the Action of Catecholamines on the Postjunctional Membrane Summary. 298

11 xiv 4. Cholinergic Transmission Introduction Localization of Acetylcholinesterase Electrophysiology of Cholinergic Transmission Ionic Basis of the Action of ACh on the Post junctional Membrane Summary Purinergic Transmission Introduction Electrophysiology of Purinergic Transmission Summary Conclusions References 317 Chapter 11 " Trophic" Functions 329 Lloyd Guth 1. Introduction Regulation of Taste Buds Regulation of Amphibian Limb Regeneration Regulation of Physiological and Metabolic Properties of Muscle Resting Membrane Potential Acetylcholine Sensitivity Cholinesterase Activity The Role of ACh Release The Dynamic Nature of the Muscle Fiber Plasticity of the Motor Unit Mechanisms of Neural Regulation References 342 SECTION ill-receptors-structure AND FUNCTION Chapter 12 Cutaneous Receptors 347 Ainsley /ggo 1. Introduction Morphology of Cutaneous Nerves Uniformity of Cutaneous Axons Relative Numbers of Myelinated and Nonmyelinated Axons 351

12 xv 3. Morphology of Cutaneous Receptors 3.1. Encapsulated Receptors 3.2. Unencapsulated Corpuscular Receptors 3.3. Noncorpuscular Receptors. 4. Physiology of Cutaneous Receptors 4.1. Cutaneous Mechanoreceptors 4.2. Cutaneous Thermoreceptors 4.3. Nociceptors 5. References Chapter 13 The Pacinian Corpuscle. Carlton C. Hunt 1. Introduction. 2. Morphology 3. Afferent Responses to Mechanical Stimuli 4. Mechanical Properties of the Corpuscle 5. Receptor Potentials. 6. Impulse Activity in the Nerve Terminal 7. Distribution of Pacinian Corpuscles. 8. Central Effects of Impulses from Pacinian Corpuscles 9. References Chapter 14 Receptors in Muscles and Joints P. B. C. Matthews 1. Introduction. 2. Joint Receptors 3. Tendon Organs 4. Muscle Spindles 4.1. Reptiles Amphibia 4.3. Birds 4.4. Mammals 5. Uncertain Origin of Adaptation 6. References

13 xvi Chapter 15 Enteroceptors J. G. Widdicombe 1. Introduction. 2. Methods Histology Physiology 3. Cardiovascular Receptors Systemic Arterial Baroreceptors 3.2. Pulmonary Arterial Baroreceptors 3.3. Ventricular Receptors 3.4. Atriovenous Receptors 4. Respiratory System Receptors 4.1. Cough and Irritant Receptors 4.2. Pulmonary Stretch Receptors 4.3. Type J Receptors 4.4. Other Receptors 5. Alimentary System Receptors 5.1. Muscular Receptors Serosal Receptors 5.3. Muscularis Mucosae Receptors 5.4. Chemoreceptors 5.5 Hepatic Osmoreceptors 6. Urinary Tract Receptors 6.1. Bladder 6.2. Urethra. 7. Other Enteroreceptors 8. References Chapter 16 Arterial Chemoreceptors 487 T. J. Biscoe 1. Introduction Structure Light Microscopy Electron Microscopy Degeneration Studies Function Types of Activity in the Nerve Supply to the Receptor Complex The Type I Cell 494

14 xvii 4. The Identity of the Receptor The Received View A New Hypothesis References 500 Chapter 17 Taste Receptors 507 M asayasu Sato 1. Introduction Gustatory Nerve Fiber Response to Chemical Stimuli Multiple Sensitivity of Single Chorda Tympani Fibers Neural Code for Quality of Taste and" Across-Fiber Pattern" Theory Electrical Responses of Gustatory Cells to Chemical Stimuli Innervation and Structure of Taste Bud How Do Gustatory Cells Respond to Chemical Stimuli? References 515 Index 517