The Fine Structure of the Nervous System
The Fine Structure of theNervous System                       Neurons and Their Supporting Cells                      Thir...
Oxford University PressOxford New York Toronto Delhi BombayCalcutta Madras Karachi Petaling JayaSingapore Hong Kong Tokyo ...
PrefaceWe wish to thank our many friends and colleagues who encouraged us to un-dertake a third edition of this book on th...
decline in standards is regrettable, it reflects the fact that electron microscopy ofthe nervous system has passed the cla...
DeFeo for providing facilities for one of us (A.P.) to enjoy a session of quietwriting at the University of Hawaii, and Dr...
Contents  List of Illustrations, xv1 General Morphology of the Neuron, 32 The Neuronal Cell Body, 14  THE PERIKARYON, 14  ...
4 The Axon, 101  AXON HILLOCK AND INITIAL AXON SEGMENT, 101  THE AXON BEYOND THE INITIAL SEGMENT, 108    Neurofilaments an...
The Differences Between Peripheral and Central Myelin Sheaths, 262      The Proximity of Adjacent Sheaths, 262      The Th...
9 Choroid Plexus, 328    THE CHOROIDAL EPITHELIUM, 330    THE VASCULARIZED CONNECTIVE TISSUE CORE, 336    FUNCTIONS OF THE...
List of Illustrations1-1    The Neuronal Cell Body, 112-1    The Cell Body of a Pyramidal Cell, 172-2    A Purkinje Cell, ...
3-10    Olfactory Bulb, 933-11    Myelinated Dendrite in Olfactory Bulb, 973-12    Dendrite Growth Cones, 994-1     Axon H...
6-6      Developing Schwann Cell Sheaths, 2296-7      Developing Schwann Cell Sheaths, Later Stage, 2316-8      Diagrammat...
11-1    The Neuropil, Anterior Horn, Spinal Cord, 35911-2    The Neuropil, Cerebellar Cortex, 36111-3    The Neuropil, Cer...
Dedicated to the Memory ofJan Evangelista Purkinje, 1787-1869Louis-Antoine Ranvier, 1835-1922Camillo Golgi, 1843-1926Santi...
The Fine Structure of the Nervous System
1                       General Morphology of                       the NeuronAnyone who has studied the early history of ...
Вы достигли ограничения для предварительного просмотра.Полный текст доступен только для сотрудников лаборатории.   You hav...
1991 peters, palay, webster. the fine structure of the nervous system
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1991 peters, palay, webster. the fine structure of the nervous system

  1. 1. The Fine Structure of the Nervous System
  2. 2. The Fine Structure of theNervous System Neurons and Their Supporting Cells Third EditionALAN PETERSWaterhouse Professor of Anatomy, Department of Anatomy and NeurobiologyBoston University School of Medicine, Boston, MassachusettsSANFORD L. PALAYBullard Professor of Neuroanatomy, EmeritusHarvard Medical School, Boston, MassachusettsHENRY DBF. WEBSTERChief, Laboratory of Experimental NeuropathologyNational Institute of Neurological Diseases and Stroke, Bethesda, MarylandNew York Oxford OXFORD UNIVERSITY PRESS 1991
  3. 3. Oxford University PressOxford New York Toronto Delhi BombayCalcutta Madras Karachi Petaling JayaSingapore Hong Kong Tokyo Nairobi Dar esSalaam Cape Town Melbourne Aucklandand associated companies inBerlin IbadanCopyright © 1970, 1976, 1991 by Alan PetersCopyrighted under the International Copyright Union.Published by Oxford University Press, Inc.,200 Madison Avenue, New York, New York 10016Oxford is a registered trademark of Oxford University PressAll rights reserved. No part of this publication may be reproduced, stored ina retrieval system, or transmitted, in any form or by any means, electronic,mechanical, photocopying, recording, or otherwise, without prior permissionof Oxford University Press.Library of Congress Cataloging-in-Publication DataPeters, Alan, 1929-The fine structure of the nervous system : neurons and their supporting cellsAlan Peters, Sanford L. Palay, Henry deF. Webster. —3rd ed.p. cm. Includes bibliographical references.Includes index.ISBN 0-19-506571-91. Nervous system—Ultrastructure.I. Palay, Sanford L. II. Webster, Henry deF. III. Title.[DNLM: 1. Nervous System—ultrastructure. WL 101 P481f]QM575.P45 1990 611.8—dc20 DNLM/DLCfor Library of Congress 90-14201Various aspects of nerve cells in light microscopic preparations.A. A small pyramidal cell from the visual cortex, Golgi method. The axon (a) descends from the cell body.B. A small neuron in the dentate nucleus of the cerebellum, Golgi method. The axon (a) is represented only by its initial segment. c.Protoplasmic (velate) astrocyte in the gray matter, Golgi method.D. Oligodendrocyte in the white matter, Golgi method.E. A motor neuron in the spinal cord showing the Golgi apparatus, osmium tetroxide impregnation.F. A motor neuron in the abducens nucleus showing the distribution of mitochondria, Altmann-Kull method.G. A motor neuron in the spinal cord showing the distribution of neurofibrils, Cajals silver stain. H. A motor neuron in theabducens nucleus showing the disposition of the Nissl bodies, thionin. .i. A dorsal root ganglion cell, showing the axon coiled about the* perikaryon and dividing into central and peripheral fibers, j. Amyelinated peripheral nerve fiber, showing a node of Ranvier, Schmidt-Lanterman clefts, Schwann cell nucleus, and neurofibrils.98765432Printed in the United States of Americaon acid-free paper
  4. 4. PrefaceWe wish to thank our many friends and colleagues who encouraged us to un-dertake a third edition of this book on the fine structure of the nervous system.This revision, like the previous editions (1970 and 1976), aims "to present inwords and pictures an account of the salient features of mammalian neuronsand neuroglial cells." We have thoroughly revised the text in order to bring itup to date, and we have exchanged many of the original micrographs for onesthat we believe better show the characteristics of various structures. Throughthe generosity of our colleagues, we have been able to add new freeze-fracturedmaterial and some deep-etched preparations, as well as examples of variouslabeling techniques. Consequently, the number of figures has increased from 118to 137, and 51 of them are new illustrations. Since the last edition was published there has been not only an informationexplosion in neuroscience, but also a notable improvement in microtomes andelectron microscopes, so that the production of good electron micrographs posesless of a challenge than it did even a decade ago. At the same time, however,some of the "art" of electron microscopy has been lost. In the 1960s and early1970s, when the technical demands of electron microscopy were greater, inves-tigators devoted themselves wholeheartedly to acquiring the skills necessary forproducing electron micrographs that were both informative and esthetically at-tractive. Sharp and clean images of well-fixed material were the aims of everycytologist. Considerable effort was expended in the pursuit of the most completerendition of protoplasmic structure possible. Such images permitted neurocytol-ogists to distinguish and describe all the components of the complex tissue thatthe brain of any animal contains. Today, it is taken for granted that any studythat requires them can be illustrated with electron micrographs. But with theincreasing facility with the elementary techniques has come a decline in the ex-acting criteria both for acceptable electron micrography and for credible inter-pretation of the profiles displayed within them. Good examples of these changesin standards can be found in the identification of synaptic junctions in tissuestaken from tracing experiments or from immunocytochemical studies. While this vii
  5. 5. decline in standards is regrettable, it reflects the fact that electron microscopy ofthe nervous system has passed the classic stage of exploration. Electron micro-scopy is now being used to examine specific issues, such as the interconnectionsamong neurons and the locations of specific proteins or neuroactive substances.Fifteen years ago we were using degeneration techniques and tracers, such ashorseradish peroxidase or radioactively labeled amino acids, in order to under-stand how the nervous system is constructed. Although important informationwas obtained through the use of these methods and they are still useful, theirplace at the forefront has largely been ceded to intracellular filling techniquesand combined Golgi-electron microscopy. But the modern explosion in the neu-romorphological sciences has been brought about by the use of antibodies toidentify the chemical signatures of neural pathways and individual neurons andsynaptic terminals. For all of these new approaches the appreciation of finestructure is more pertinent than ever. We have rewritten this book in the light of the information obtained throughthe use of these newer methods because they have led to a much better under-standing of the relationships between neuronal circuits and their functions. Con-sequently we have extensively revised all of the chapters and added many newreferences to the bibliography. We have, however, retained those references thatreflect the foundations upon which our new information is based. A reader fa-miliar with the previous edition of this book will certainly recognize paragraphsand descriptions that have not changed appreciably because no significant newknowledge has come to our notice in that area. Other chapters, such as thechapters on axons, synapses, sheaths, and the neuropil, have been almost en-tirely rewritten. In the chapter on the neuropil we have tried to show the pos-sibilities and limitations of the various techniques, so that this chapter has be-come a vehicle for giving an account of the methods available. To a large extentthis strategy has allowed us to eliminate details of techniques from the otherchapters. We hope that in this version of the book we have succeeded in correlatingstructure and function and in providing a reference source of electron micro-graphs and literature, in which both experienced neuroscientists and studentsinterested in the fine structure of the nervous system can find information be-yond the scope of their immediate interests. Although most of the illustrations come from our own collections, we haverelied on the generosity of many colleagues for illustrations of structures andtechniques that we have not explored ourselves. We gratefully acknowledge thecontributions of figures from J. Anders, D. J. Allen, Dennis Bray, Milton Bright-man, Mary B. Bunge, Victoria Chan-Palay, M. W. Cloyd, Edward V. Famig-lietti, Martin L. Feldman, James E. Hamos, C. K. Henrikson, John E. Heuser,J. Hirokawa, James Kerns, Frank N. Low, Douglas L. Meinecke, Enrico Mug-naini, Elio Raviola, Thomas S. Reese, Bruce Schnapp, Constantino Sotelo, Deb-orah W. Vaughan, James E. Vaughn, Bruce W. Warr, and Raymond B. Wuer-ker. We are also grateful to Janet Harry, Mary Alba, Lilian Galloway and Joyce Resil for typing the several versions of the manuscript and references, and to Katherine Harriman, Karen Josephson and Claire Sethares for their expert tech- nical assistance. In addition we wish to thank Dr. R. Hammer and Dr. V. J.viii PREFACE
  6. 6. DeFeo for providing facilities for one of us (A.P.) to enjoy a session of quietwriting at the University of Hawaii, and Dr. P. Hashimoto of Osaka Universityfor providing facilities for another of us (S.L.P.) during an extended visit. By no means of least importance, we wish to pay tribute to our wives. With-out their patience, understanding and support, we could not have completedthis revision.Boston, Massachusetts Alan PetersConcord, Massachusetts Sanford L. PalayBethesda, Maryland Henry deF. WebsterFebruary 1990 PREFACE ix
  7. 7. Contents List of Illustrations, xv1 General Morphology of the Neuron, 32 The Neuronal Cell Body, 14 THE PERIKARYON, 14 The Nissl Substance, 14 The Agranular Reticulum, 22 The Golgi Apparatus, 26 Multivesicular Bodies, 33 Lysosomes, 33 Peroxisomes, 34 Lipofuscin Granules, 34 Mitochondria, 38 Microtubules and Neurofilaments, 40 Cilia and Centrioles, 41 Cytoplasmic Inclusions, 42 THE NUCLEUS, 48 General Morphology, 48 The Nuclear Envelope, 52 The Karyoplasm, 58 The Nucleolus, 59 Nuclear Inclusions, 60 THE PLASMA MEMBRANE, 643 Dendrites, 70 GENERAL MORPHOLOGY, 70 THE CYTOPLASM OF DENDRITES, 76 THE DENDRITIC SPINES, 82 MYELINATED DENDRITES, 96 GROWING TIPS OF DENDRITES, 98 xi
  8. 8. 4 The Axon, 101 AXON HILLOCK AND INITIAL AXON SEGMENT, 101 THE AXON BEYOND THE INITIAL SEGMENT, 108 Neurofilaments and Microtubules, 110 Membranous Components, 119 Cytoskeleton, 122 The Axonal Membrane, 124 AXOPLASMIC FLOW, 126 THE AXON GROWTH CONE, 132 THE IDENTIFICATION OF SMALL AXONS AND DENDRITES, 1375 Synapses, 138 THE NEUROMUSCULAR SYNAPSE, 138 INTERNEURONAL CHEMICAL SYNAPSES, 147 The Synaptic Junction, 150 The Presynaptic Grid, 154 The Synaptic Cleft, 159 Potsysnaptic Densities, 160 Freeze-Cleavage, 166 Nonsynaptic Junctions Between Neurons, 168 Synaptic Vesicles With Clear Centers, 169 Shapes and Sizes of Vesicles, 169 Correlation Between Vesicle Shape and Function of Chemical Synapses, 176 Granular Vesicles, 178 Neurosecretory Vesicles, 184 Other Presynaptic Organelles, 186 Other Postsynaptic Organelles, 188 Synaptic Relations, 190 Axo-Dendritic Synapses, 190 Axo-Somatic Synapses, 191 Axo-Axonal Synapses, 192 Dendro-Dendritic Synapses, 195 Somato-Dendritic, Dendro-Somatic and Somato-Somatic Synapses, 196 Somato-Axonic Synapses, 198 Dendro-Axonic Synapses, 198 Synaptic Glomeruli, 199 ELECTROTONIC SYNAPSES, 203 MIXED SYNAPSES, 207 "SYNAPSES" INVOLVING NEUROGLIAL CELLS, 2106 The Cellular Sheaths of Neurons, 212 THE SHEATHS OF UNMYELINATED GANGLION CELLS, 213 THE SHEATHS OF UNMYELINATED NERVE FIBERS, 218 THE SHEATHS OF MYELINATED FIBERS, 222 Internodal Peripheral Myelin, 224 The Formation of the Peripheral Myelin Sheath, 226 Internodal Central Myelin, 232 The Formation of the Central Myelin Sheath, 234 Identification of the Myelin-forming Cell of the Central Nervous System, 242 The Mechanism of Myelin Formation, 246 The Node of Ranvier, 250 The Schmidt-Lanterman Incisures, 261xii CONTENTS
  9. 9. The Differences Between Peripheral and Central Myelin Sheaths, 262 The Proximity of Adjacent Sheaths, 262 The Thickness of Myelin Lamellae, 263 The Radial Component of the Central Sheath, 264 THE SHEATHS OF MYELINATED GANGLION CELLS, 265 MYELIN SHEATHS OF DENDRITES IN THE CENTRAL NERVOUS SYSTEM, 266 FUNCTIONS OF SATELLITE AND SCHWANN CELLS, 266 Early Development, 266 Axon Ensheathment and Myelin Formation, 267 Biochemical Relationships, 269 Breakdown of Myelin, 271 Other Functions, 2727 The Neuroglial Cells, 273 THE DEVELOPMENT OF NEUROGLIA, 274 ASTROCYTES, 276 Fibrous Astrocytes, 277 Protoplasmic Astrocytes, 281 Functions of Astrocytes, 284 Structural Support, 284 Guidance for Neuroblast Migration and Axon Growth, 286 Graft Survival and Function, 288 Isolation of Receptive and Nodal Surfaces of Neurons, 288 Interactions with Oligodendroglia: Role in Myelination, 290 Blood-Brain Barrier, 290 Interactions with the Immune System, 293 Repair, 294 OLIGODENDROCYTES, 295 General Morphology, 295 Functions of Oligodendrocytes, 298 NEUROGLIAL CELLS INTERMEDIATE BETWEEN ASTROCYTES AND OLIGODENDROCYTES, 302 MICROGLIA, 304 General Morphology, 306 Functions, 308 Discussion, 3088 The Ependyma, 312 THE MORPHOLOGY OF EPENDYMAL CELLS, 313 THE MORPHOLOGY OF TANYCYTES, 318 INTRAVENTRICULAR NERVE ENDINGS, 322 THE SUBEPENDYMA, 324 FUNCTIONS OF CELLS IN THE EPENDYMA, 325 Movements of Cerebrospinal Fluid, 325 Capture of Materials Present in the Cerebrospinal Fluid, 325 Proliferation, 325 Support, 326 Sensory Function, 326 Secretion, 326 Transport of Substances, 327 CONTENTS xiii
  10. 10. 9 Choroid Plexus, 328 THE CHOROIDAL EPITHELIUM, 330 THE VASCULARIZED CONNECTIVE TISSUE CORE, 336 FUNCTIONS OF THE CHOROID PLEXUS, 33810 Blood Vessels, 344 CAPILLARIES, 344 ARTERIES AND ARTERIOLES, 350 VEINS, 35411 The Neuropil, 356 THE IDENTIFICATION OF PROFILES IN THE NEUROPIL, 356 THE ORGANIZATION OF THE NEUROPIL AND SYNAPTIC CONNECTIONS, 364 Golgi—Electron Microscope Technique, 366 Intracellularly Injected Markers, 368 Reconstruction of Neurons and Their Processes, 370 Experimental Degeneration, 372 Intracellular Transport of Radioisotopes, 375 Antibodies to Neurotransmitters, 375 Antibodies to Neuropeptides, 380 Techniques Using Two Antibodies, 381 Combined Techniques, 38212 Connective Tissue Sheaths of Peripheral Nerves, 384 EPINEURIUM, 384 PERINEURIUM, 385 ENDONEURIUM, 388 FUNCTIONS OF CONNECTIVE TISSUE SHEATHS, 39213 The Meninges, 395 DURA MATER, 396 ARACHNOID MATER, 398 PIA MATER, 400 ENTRY OF PERIPHERAL NERVES INTO THE CENTRAL NERVOUS SYSTEM, 402 ARACHNOID VILLI, 404 References, 407 Index, 487xiv CONTENTS
  11. 11. List of Illustrations1-1 The Neuronal Cell Body, 112-1 The Cell Body of a Pyramidal Cell, 172-2 A Purkinje Cell, 192-3 Pyramidal Neuron, 212-4 Granule Cells of the Cerebellum, 232-5 The Cytoplasm of a Purkinje Cell, 252-6 The Cytoplasm of a Dorsal Root Ganglion Cell, 292-7 The Cytoplasm of a Dorsal Root Ganglion Cell, 312-8 Nissl Bodies in an Anterior Horn Cell, 352-9 The Golgi Apparatus and the Nissl Substance of a Purkinje Cell, 372-10 Golgi Apparatus of the Purkinje Cell, 392-11 Two Views of the Golgi Apparatus in a Freeze-Fractured Preparation, 432-12 Golgi Apparatus, Lysosomes, Nematosomes, and Fibrillary Inclusions, 452-13 Lipofuscin Granules, Cilia, and Centrioles, 472-14 Laminated Inclusion Body, 492-15 The Nuclear Envelope, Nissl Bodies, and Golgi Apparatus, 552-16 Nuclear Pores, 572-17 The Nucleolus, 612-18 Intranuclear Inclusions, 632-19 Diagram of Freeze Fracturing, 652-20 The Edge of a Purkinje Cell, Freeze-Fractured Preparation, 673-1 Pyramidal Neuron in Cerebral Cortex, 733-2 The Apical Dendrites of Pyramidal Cells, 753-3 Dendrite of a Purkinje Cell, 793-4 Dendrite of a Purkinje Cell, 813-5 Dendrites in Longitudinal and Transverse Section, 833-6 Dendrites in the Neuropil of the Anterior Horn: Transverse Section, 85 3-7 Dendrites in the Neuropil of the Cerebral Cortex, 87 3-8 Dendrites in Cerebellar and Cerebral Cortex, 89 3-9 A Spiny Branchlet of a Purkinje Cell Dendrite, 91 xv
  12. 12. 3-10 Olfactory Bulb, 933-11 Myelinated Dendrite in Olfactory Bulb, 973-12 Dendrite Growth Cones, 994-1 Axon Hillock and the Initial Axon Segment, 1034-2 Axon Hillock and the Initial Axon Segment, 1054-3 The Initial Axon Segment, Longitudinal Section, 1074-4 The Initial Segment, Transverse Section, 1094-5 The Initial Axon Segment and the Node of Ranvier Compared, 1114-6 Axon Hillock and Initial Segment of a Trigeminal Ganglion Cell, 1134-7 The Initial Segment of a Trigeminal Ganglion Cell, 1154-8 Microtubules, Neurofilaments, and Neuroglial Filaments, 1174-9 Axoplasmic Organelles, 1214-10 Quick-frozen and Deep-etched Axoplasm, 1234-11 Quick-frozen and Deep-etched Axoplasm, 1254-12 Growth Cone from a Sympathetic Neuron in Tissue Culture, 1294-13 Growth Cone from a Sympathetic Neuron in Tissue Culture, 1314-14 Small Axons in the Molecular Layer of the Cerebellum, 1334-15 Unmyelinated Axons Entering the Olfactory Bulb, 1355-1 Motor End Plate, 1415-2 Motor End Plate, 1435-3 Freeze-Fractured Motor End Plates to Show Vesicle Release, 1455-4 Puncta Adhaerentia, 1495-5 Axon Terminal Emerging from the Myelin Sheath, 1515-6 Synapses in the Cerebral Cortex, 1535-7 Asymmetric Synapses, Cerebral Cortex, 1555-8 Presynaptic Grid, 1575-9 Synapses in the Cerebellum, 1615-10 The Synaptic Junction Between an Axon and a Dendritic Thorn, 1635-11 Asymmetric and Symmetric Synapses, 1655-12 Synapses in the Anterior Horn of Spinal Cord, 167 5-13 Anterior Horn of Spinal Cord, 171 5-14 The Glomerulus, Cerebellar Cortex, 173 5-15 The Presynaptic Membrane, P face, 175 5-16 The Presynaptic Membrane, E face, 181 5-17 A Variety of Synapses, 183 5-18 Axo-axonic Synapse and Dense-cored Vesicles, 185 5-19 Dendro-dendritic and Somato-dendritic Synapses, 189 5-20 The Glomerulus, Lateral Geniculate Nucleus, 197 5-21 Electrotonic Synapses, 205 5-22 A Mixed Synapse, 209 6-1 The Sheath Surrounding a Dorsal Root Ganglion Cell, 215 6-2 The Sheath Surrounding a Trigeminal Ganglion Cell, 217 6-3 Unmyelinated Axons, Adult Peripheral Nerve, 219 6-4 Unmyelinated Axons, Adult Peripheral Nerve, 221 6-5 Myelinated Axon, Adult Peripheral Nerve, 227xvi ILLUSTRATIONS
  13. 13. 6-6 Developing Schwann Cell Sheaths, 2296-7 Developing Schwann Cell Sheaths, Later Stage, 2316-8 Diagrammatic Representation of the Formation of Peripheral Myelin Sheaths, 2336-9 Myelinated Nerve Fibers, Central Nervous System, 2356-10 Myelin Sheaths: Central Nervous System, 2376-11 Developing Myelin Sheaths, Central Nervous System, 2396-12 Developing Myelin Sheaths, Central Nervous System, 2416-13 Diagrammatic Representation of the Formation of Myelin in the Central Nervous System, 2436-14 The Myelin Forming Cell, Central Nervous System, 2456-15 The Node of Ranvier, Peripheral Nervous System, 2496-16 The Node of Ranvier, Central Nervous System, 2516-17 The Node and the Paranode, Central Nervous System, 2536-18 Freeze-Fractured Myelin Sheaths, 2556-19 Freeze-Fractured Myelin Sheaths, 2576-20 Freeze-Fractured Myelin Sheaths, 2596-21 Diagram of Membrane Particle Distribution at the Paranode, 260 7-1 Fibrous Astrocytes, 279 7-2 Protoplasmic Astrocytes, 283 7-3 Protoplasmic Astrocytes, 285 7-4 Protoplasmic Astrocyte, 287 7-5 Glial Limiting Membrane; Cerebral Cortex, 289 7-6 Orthogonal Assemblies and Gap Junctions of Astrocytes in Freeze-Fracture Preparations, 291 7-7 Perineuronal Oligodendrocytes, 297 7-8 An Oligodendrocyte, 299 7-9 Interfascicular Oligodendrocyte, 301 7-10 A Perineuronal Microglial Cell, 303 7-11 A Microglial Cell in a Senile Plaque, 307 8-1 The Ependyma, 315 8-2 Ependymal Surface, 317 8-3 The Cilia of Ependymal Cells, 319 8-4 Ependymal Cell Cytoplasm, 321 8-5 Ependymal Cell Junctions, 323 9-1 Scanning Electron Micrograph of the Choroid Plexus, 329 9-2 Epithelium and Stroma of the Choroid Plexus, 331 9-3 The Choroid Plexus, 333 9-4 Choroid Plexus, Intercellular Junctions, 335 9-5 Choroid Plexus, Intercellular Junctions, 337 9-6 Choroid Plexus, Surface Structures, 339 9-7 The Basal Ends of Choroidal Cells, 341 9-8 Kolmer Cells, 343 10-1 Capillary and Pericyte, 347 10-2 Capillaries, 349 10-3 Small Blood Vessel, 351 10-4 Intracerebral Arterioles, 353 10-5 An Arteriole, 355 ILLUSTRATIONS xvii
  14. 14. 11-1 The Neuropil, Anterior Horn, Spinal Cord, 35911-2 The Neuropil, Cerebellar Cortex, 36111-3 The Neuropil, Cerebral Cortex, 36311-4 Lateral Geniculate Body Glomerulus, 36511-5 Degenerating Boutons, 36711-6 Filamentous Degeneration and Horseradish Peroxidase-labeled Neurons, 36911-7 Golgi-Electron Microscope Technique, 37111-8 Intracellular Horseradish Peroxidase Injection, 37311-9 Glutamic Acid Decarboxylase Immunoreactive Axon Terminals, 37711-10 Vasoactive Intestinal Polypeptide in the Cerebral Cortex, 37912-1 Connective Tissue Sheaths of Nerves, 38712-2 Epineurial and Perineurial Sheaths, 38912-3 Perineurium and Endoneurium of Peripheral Nerve, 39113-1 Meninges by Scanning Electron Microscopy, 39713-2 Dura Mater, 39913-3 Arachnoid Mater, 40113-4 Pia Mater and Glia Limitans, 403xviii ILLUSTRATIONS
  15. 15. Dedicated to the Memory ofJan Evangelista Purkinje, 1787-1869Louis-Antoine Ranvier, 1835-1922Camillo Golgi, 1843-1926Santiago Ramon у Cajal, 1852-1934
  16. 16. The Fine Structure of the Nervous System
  17. 17. 1 General Morphology of the NeuronAnyone who has studied the early history of cy- nervous system lay in the shape of the nerve celltology cannot fail to be impressed by the slow itself and to some extent in its size. The medusa-development of the concept of the nerve cell. Most like nerve cell, with its corona of seemingly endlesstypes of cells do not have a history. Once the idea processes, was bizarre. Other cells had relativelywas grasped, in the theory of Schleiden (1838) simple shapes—globular, cylindrical, squamous,and Schwann (1839), that cells are the architec- fusiform, and so on. Some fitted one into the othertonic units of living things, it was fairly quick like pieces of a jigsaw puzzle to form an epithe-work to recognize them in the various tissues and lium; others lay free and definable in the tissueto proceed to the study of their contents, their fluids. Many, such as cartilage or certain epithelialinterrelations, and their functions. But the nerve cells, were clearly circumscribed by walls. Onlycell was more perplexing. It occasioned so much pigment cells, astrocytes, myoepithelial cells, anddifficulty for its students that almost a century a few others had shapes even roughly approxi-passed before they could agree upon its shape. At mating those of nerve cells. But aside from thefirst it was thought to be an independent globular fact that some of these examples were unknowncorpuscle suspended among nerve fibers, which in the early days, such cells could be easily encom-looped and coiled about it and which it somehow passed in a single field or at least in a singlenourished (Valentine, 1836). Later, when the con- preparation under the microscope. The multipolartinuity between the perikaryon and the nerve fi- nerve cell, however, with its meter-long axon didbers was finally established (Remak, 1838, 1841; not fit into a single section and could not be easilyHelmholtz, 1842; Hannover, 1844; Kolliker, 1844; plucked from its context or distinguished from itsBidder, 1847; Wagner, 1847), then the nerve cell neighbors by the methods used for other cells.appeared to have no definite boundaries and seemed New methods had to be developed. And so a trueendless. Except for the fibers attached to organs cell theory of the nervous system did not emergein the periphery, the processes of all nerve cells until the discovery and exploitation of specialseemed to be equivalent and to be confluent with techniques that had the merit of bringing intoone another. The nerve cells seemed to be only view entire nerve cells as if dissected or isolatednodal points in an enormously intricate reticulum from the central nervous system.pervading the nervous system (Gerlach, 1858, Actually, the first successful method was mi-1872). It appeared that the cell theory did not crodissection of whole nerve cells from hardenedreally apply to the nervous system; one had rather specimens of brain and spinal cord. On the basisto speak of cell territories or spheres of influence of experience with such isolated cells, Deiters (1865)surrounding nucleated centers. was able to distinguish between the numerous It seems clear that one of the major obstacles branching processes that we now call dendrites to the appreciation of the cellular nature of the and the single process that slips into a myelin 3
  18. 18. Вы достигли ограничения для предварительного просмотра.Полный текст доступен только для сотрудников лаборатории. You have reached restrictions for preliminary viewing. The full text is accessible only to employees of laboratory.

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