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Sensory Coding in the mammalian nervous system PDF

388 Pages·1972·10.175 MB·English
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SENSORY CODING in the mammalian nervous system Neuroscience Series EDITOR: Arnold Towe, University of Washington PREVIOUS TITLES: Physiological Basis of the Alpha Rhythm by Per Andersen (University of Oslo) and Sven Andersson (University of Gothenburg) The Bisected Brain by Michael S. Gazzaniga (New York University) Hibernation and the Hypothalilmus by Nicholas Mrosovky (University of Toronto) SENSORY CODING in the mammalian nervous system GEORGE SOMJEN Duke University APPLETON-CENTURY-CROFTS EDUCATIONAL DIVISION MEREDITH CORPORATION ISBN 978-1-4684-8192-1 ISBN 978-1-4684-8190-7 (eBook) DOI 10.1007/978-1-4684-8190-7 Copyright © 1972 by Softcover reprint of the hardcover 1st edition 1972 MEREDITH CORPORATION All rights reserved This book, or parts thereof, must not be used or reproduced in any manner without written permission. For information address the publisher, Appleton-Century-Crofts, Educational Division, Meredith Corporation, 440 Park Av enue South, New York, N. Y. 10016. 73747576/1098765432 Library of Congress Card Number: 72-78258 390-F -83013-5 Acknowledgments Figure 6-F, p. 148. From W. R. Lowenstein, Excitation and the inactivation in a receptor membrane, Annals of the N. Y. Academy of Sciences, 94, 2: Fig. 8, p. 516. © 1961 by The New York Academy of Sciences. Reprinted by permission. Figure 16-A, p. 166. From C. C. Hunt, On the nature of vibration receptors in the hind limb of the cat, The Journal of Physiology, ISS (1961): Figure 3, p. 178. Reprinted by permission. Figure 16-B, p. 166. From M. Sato, Response of pacinian corpuscles to sinusoidal vibrations, The Journal of Physiology, 159 (1961): Figure 4, p. 159. Reprinted by permission. Figure 16-C, p. 166. From V. B. Mountacastle et al., Neural basis of the sense of flutter vibration, Science, ISS (February 3, 1967): Figure 2, pp. 597-600. Copyright 1967 by the American Association for the Advancement of Science. Reprinted by permission. Figures 17-A and B, p. 167. From S. Skoglund, Anatomical and physiological studies of knee joint innervation in the cat,Acta Physiologica Scandinavica, Supp. 124: Figures 17 and 18. Reprinted by permission. vi Figure 19-A, p. 171. From H. Hensel and D. R. Kenshalo, Warm receptors in the nasal region of cats, The Journal of Physiology, 206 (1969): 99-112. Reprinted by permission. Figure 19-D, p. 171. From H. Hensel, A. Igoo, I Witt. A quantitative study of sensitive cutaneous thermoceptors with C afferent fibres, The Journal of Physiology, 153 (1960): Figure 5, pp. 113-126. Reprinted by permission. Figure 23-A, p. 177. From H. Davis et al., Acoustic trauma in the guinea pig, Journal of the Acoustical Society ofA merica, 25 (1953): 1180-1189. Reprinted by permission. Figure 23-B, p. 177. From Hearing and Deafness, third edition, edited by Halowell Davis and S. Richard Silverman. Copyright (1947), © 1960, 1970 by Holt, Rinehart and Winston, Inc. Reprinted by permission of Holt, Rinehart and Winston, Inc. Figure 23-C, p. 177. From C. Bredberg et al., Scanning electron microscopy of the organ of corti, Science, 170 (November 20, 1970): 861-863. Copyright 1970 by the American Association for the Advancement of Science. Reprinted by permission. Figure 23-D, p. 177. From Cold Spring Harbor Laboratory. Reprinted by permission. Figure 24-D, p. 179. From G. V. Bekesy, The variation of phase along the basilar membrane with sinusoidal vibration, Journal of the Acoustical Society of America, 19 (1947): 1180-1189. Reprinted by permission. Figure 25-B, p. 181. From S. D. Erulkar, P. G. Nelson, and J. S. Bryan, Experimental and theoretical approaches to neural processing in the cential auditory pathway in Contributions to Sensory Physiology, edited by W. D. Neff, Vol. 2 (New York: Academic Press, 1968). Figure 2a. Copyright © 1968, by Academic Press. Reprinted by permission. Figure 25-C, p. 181. From J. E. Hind et al., Coding of information pertaining to paived low-frequency tones in single auditory nerve fibers of the squirrel monkey, Journal of Neurophysiology, 30 (1967): Figure 10, F-1. Reprinted by permission. Figure 26-A, p. 183. From W. D. Neff, Vocalization and lateralization of sound in space in Hearing Mechanisms in Vertebrates, (Ciba Foundation Symposium, 1967), edited by A. V. S. Reush and J. Knight, (London: J. & A Churchill, 1968): Figure 3, p. 212. Reprinted by permission. Figure 26-B, p. 183. From I. C. Whitfield, The Auditory Pathway, Monographs of the Physiological Society, No. 17 (London: Edward Arnold, Ltd., 1967): Figure 15. Reprinted by permission. Figure 27-A, p. 184. From Y. Katsuki, Integrative organization in thalamic and cortical auditory centers in The Thalamus, edited by D. P. Purpura and M. D. Yahr, (New York: Columbia University Press, 1966): p. 349. Reprinted by permission. Figure 27-B, p. 184. From J. C. Boudreau and C. Tsuchitani, Cat superior olive s-segment cell discharge to tinal stimulation in Contributions to Sensory Physiology, edited by W. D. Neff, vol. 4 (New York: Academic Press, 1968): Figure 13. Copyright © 1968 by Academic Press. Reprinted by permission. Figure 27-C, p. 184. From S. Oonishi and Y. Katsuki, Functional organization and integrative mechanism of the auditory cortex of the cat, Japanese Journal ofP hysiology, 15 (1965): Figure 1, p. 345. Reprinted by permission. Figure 27-D, p. 184. From S. D. Eruklar, P. G. Nelson, and J. S. Bryan, Experimental and theoretical approaches to neural processing in the central auditory pathway in Contributions to Sensory PhYSiology, edited by W. D. Neff, vol. 2 (New York: Academic Press, 1968): Figure 4. Copyright © 1968 by Academic Press. Reprinted by permission. Figure 30, p. 188. From R. L. DeValois, Behavioral and electrophysiological studies of primate vision in Contributions to Sensory Physiology, edited by W. D. Neff, vol. 1 (New York: Academic Press, 1965): Figure 8. Copyright © 1965 by Academic Press Inc. Reprinted by permission. Figure 32, p. 192. From J. E. Dowling and B. B. Boycott, Organization of the primate retina, Proceedings of the Royal Society, Ser. B 166 (1966): Figure 23, p. 104. Reprinted by permission. vii Contents Preface xvii PART I: INTRODUCTION 1 Chapter 1: Assumptions 3 Chapter 2: Methods 9 Critique 9 Design 10 Electrical recording 10 Probes 11 Experimental animals 12 Stimulation 14 Chapter 3: Signalling in the Nervous System 17 Neurons 17 The nerve impulses: Physical nature 18 The nerve impulses: Conduction in myelinated and nonmyelinated fibers 22 Generator potentials and receptor potentials 23 Transmission between neurons 27 PART II: THE FIRST-ORDER CODE 31 Chapter 4: Variables of the Sensory Code 33 The diversity of stimuli and of sensory signalling 33 The coding of quality 34 Intensity 38 Input-output functions of individual afferent fibers 39 Input-output functions of populations 43 Time 45 Velocity: Receptor adaptation 46 The coding of size, shape, and location 48 ix x Contents Chapter 5: Direct Contact with the World 51 The skin as a sense organ 52 The so-called 'sensory spots', and the specificity of cutaneous afferents 54 Corpuscles of Pacini 57 Partially and slowly adapting mechanoreceptors 62 Receptors of hair follicles 62 Temperature receptors 63 Temperature sensitivity of other receptors 66 Pain 69 The shape of things touched 73 Chapter 6: The Inner Senses 75 Proprioception 1: Feedback signals of movement 75 Proprioception 2: Sense organs of joints 77 Visceral receptors 80 Receptor cells sought within the brain 82 Chapter 7: The External Chemical Senses 83 Taste: The stimuli 83 Taste: The receptors 85 Taste: The code 88 The receptors and their nerves 90 Smell 92 The common problem of the codes of taste and of smell 96 Common chemical sense, and how it relates to pain 97 Chapter 8: The Inner Ear 99 The transducers 99 Semicircular canals 102 The utricle 103 The saccule 104 The sound stimulus 104 The cochlea 105 Cochlear potentials and the stimulation of the receptors 107 Analysis of frequencies 110 The neural code 112 Alternatives to the Bekesy /Davis/Tasaki/Whitfield model: The pulse-frequency code 117 Alternatives to Davis' 'carbon microphone' model 119 Some loose ends in auditory theory 122 Chapter 9: The Photoreceptors of the Retina 125 The receptors 126 Visual pigments 127 The stimulation of photoreceptors 128 Photoreceptor potentials 131 Three receptors to see so many tints, hues and shades 133 Contents xi PART III: CODING IN THE CENTER 201 Chapter 10: Approaches to Brain Function 203 Lessons learned from electrical recording and from ablations of the brain 207 Firing patterns of central neurons 209 Chapter 11: Sensory Synaptic Cascades 215 Place and identity of relay sites 215 Transmission and transformation at relay synapses 217 How and where active processing 218 Amplifiers, attenuators, and linear operators 220 Lateral inhibition: The enhancement of contrast 223 Self-inhibition and automatic gain control 225 Relationships of transformations in the domains of time and space 225 Changes of the rules of coding: Abstractions and invariances. Transcriptions? 226 Parallel channels, redundancy, and the possible significance of fiber size 227 The cortex: Blueprint and performance 228 The cortex: On topographic representation 232 Chapter 12: Central Coding in the Somatic Senses 237 One system, two, or several? 237 A paradox resolved? 240 Neurons in somatic relays: 'Lernniscal' and 'anterolateral' types 244 Input from the face 246 Neurons of the nuclei of the dorsal columns 246 Neurons of the ventrobasal region of the thalamus 248 Cells of the first somatic receiving area (S I) of the cerebral cortex 251 Neurons of the dorsal horns of the spinal cord 253 Connections of the dorsal horns with the brain 256 More on Melzack and Wall: Support 257 Even more on Melzack and Wall: Doubts 259 The posterior group of nuclei of the thalamus 260 Spinothalamic contribution to the ventrobasal thalamus 262 Coding for skin temperature by thalamic neurons 262 Skin temperature and neurons in the somatic cortex 263 Chapter 13: The Central Code of Hearing 265 Components of the central auditory system 267 Centrifugal control in the auditory system 268 Tonotopic organization 269 Discharges of cells in the auditory pathway 271 Neural correlates of directional hearing 274 Chapter 14: The Central Code of Sight 275 xii Contents The organization of the retina 276 What excites ganglion cells: Shape of the receptive fields 277 Colored stimuli: Effects on ganglion cells and on cells of the lateral geniculate nucleus 279 Retinal ganglion cells: Adaptation to light and to darkness 283 Synaptic mechanisms of the retina 286 Detectors of movement and of direction 287 Beyond the retina 288 The visual thalamus: The lateral geniculate 288 Neurons of the visual cortex 290 Blueprints for the cortex: In series processing or parallel channels? 292 Seeing in depth 297 Visual function of the roof of the midbrain 298 Chapter 15: The Central Code of the Chemical Senses 301 Neurons in the central pathway of taste 301 The olfactory bulb 302 PART IV: POSTSCRIPT 305 In praise of redundancy 305 The hierarchies of input revisited 309 Relevancies and irrelevancies for sensory physiology in psychophysics and in information theory 310 Last words 312 PART V: LITERATURE 315 Key Titles 317 Works of historic importance and works concerned with history 317 Brain theory 318 Psychology of sensation and perception 318 Coding and information theory 319 Sensory physiology (general works) 319 Sensory receptors 319 Somatic senses 319 Chemical senses 320 Hearing 321 Vision 321 Contents xiii References 323 Name Index 373 Subject Index 381

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