Table Of ContentSENSORY 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
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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
