Table Of ContentCeskoslovenska akademie ved
Sekce biologicko-lekarska
THE DENERVATED MUSCLE
The Denervated Muse le
Edited by E. Gutmann
Scientific Editors:
Zdenek Servit and Otakar Poupa
Translated by:
Pavel Hnlk
Foreign ianguage Editor:
Joseph Cort
THE DENERVATED MUSCLE
EDITED BY E. GUTMANN
List of Contributors
A. Bass, R. Beranek, Z. Drahota,
E. Gutmann, P. Hnik,
O. Hudlicka, V. Skorpil, L. Vyklicky,
J. Zelena, R. Zak
PRAGUE 1962
Springer Science+Business Media, LLC
ISBN 978-1-4899-4856-4 ISBN 978-1-4899-4854-0 (eBook)
DOI 10.1007/978-1-4899-4854-0
©Springer Science+Business Media New York
Originally published by Nakladatelstvi Ceskoslovenske akademie ved, Praha 1962
THE DENERVATED MUSCLE
Errata
Page 32, Line 2 from below: Ellenberg read Ellenbeck
Page 35, Line 6 from above: Horton and Smith read Horton-Smith
Page 127, Line 9 from above: methodical read 'methodological
Page 208, Line 13 from above: Ben Tor read Burton
Page 273, Line 20 from above: Huscles read Muscles
Page 273, Line 21 from above: mopkins read Hopkin!!
Page 321, Line 3 from above: Chagowets read Chagovet!l
Page 360, Line 11 from below: Nimmi read Nimni
Page 363, Line 15 from below: Waking read Wakim
Page 369, Line 3 from below: Nimmi read Nimni
Page 462, Line 24 from above: influence read influences
CONTENTS
I. Denervation Studies in Research of Neurotrophic Relationships (E. Gutmann, P~ ll'imt)
A. Definition of "trophic processes" and "nervous re6ulation of trophic processes"
("trophic influence of the nervous system") . . . . . . . . . . . . . . . . 13
B. The study of trophic functions from the aspect of cellular metabolism 16
C. The trophic function of the nervous system and its place in the study of the
physiology of neurotrophic relations . . . . . . . . . . . . . . 20
D. The trophic influence of the nervous system-ar historical survey. 25
E. Mechanisms of the trophic influence of the nervous system 31
l. Components of innervation affecting trophic processes 31
a) The trophic function of the motoneuron . . . 31
b) The trophic function of vegetative nerve fibres . . 32
c) The trophic function of afferent nerve fibres . . . 35
2. The role of specific muscle function in regulation of trophic processes 37
3. The existence of an independent trophic mechanisms of the nervous system 43
F. The significance and limitations of denervation studies . . . . . . . . 49
II. Morphological Changes in the Denervated Muscle (E. Gutmann, J. Zelemi)
A. Aims and limitations of morphological studies 57
B. Denervation changes in human muscles . . . . 60
C. Expetimental studies of the denervated muscle. 67
l. Atrophy of muscle fibres . . . . . . . . . . 68
2. Changes in motor end-plates and mtiscle spindles 74
a) End-plate . . . . . . 74
b) Muscle spindles 75
3. Changes in muscle nuclei . 77
4. Degenerative changes in muscle fibres 82
5. Connective and adipose tissues . . . 87
D. Correlation between physiological, biochemical and morphological changes ac-
companying denervation atrophy 90
l. Protein changes . . . . . . . . 95
2. Changes in nuclei . . . . . . . 96
3. Changes in the content of lipids . 97
4. Changes in the content of collagen 98
III. The Effect of Denervation on Muscle Development (J. Zelena)
A. The effect of denervation upon the initial differentiation of muscle fibres . I 03
l. Teratology . . . . . . . . . . . 104
2. Experimental studies in amphibia 105
3. Experimental studies in birds . . . 106
B. The effect of denervation upon further muscle differentiation 107
l. Differentiation of muscle in tissue cultures . . . . . . . 107
2. Muscle differentiation in mammals following denervation at a late stage of
development . . . . . . . . . . . . . . . . . . . . . . . . 108
a) Differentiation of myotubes into muscle fibres in the denervated muscle . 109
b) Arrest of differentiation of myotubes into intrafusal fibres of muscle
spindles in the denervated muscle . . . . . . . . . . . . . . . . . ll2
c) Arrest of differentiation of the postsynaptic membrane of the motor end-
plate after denervation . . . . . . . . . . . . . . . . . . . . . . ll4
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C. The effect of denervation upon muscle growth . . 116
D. The effect of denervation on muscle regeneration 121
IV. Electrophysiology of Denervated Muscle (R. Beranek)
A. Excitability . . . . . . . . . . . . . . . . . 127
B. Membrane potential and spontaneous activity. . 130
C. Passive electrical properties and membrane constants 131
D. Conclusions . . . . . . . . . . . . . . . . . . . 132
V. Fibrillation Activity in Denervated Muscle (P. Hnik, V. Skorpil)
A. Conditions, onset and duration of denervation fibrillations 136
I. Conditions for appearance of denervation fibrillations 136
2. Time of onset of denervation fibrillations . 136
3. Duration of denervation fibrillations . . . . . . 138
4. Number of fibrillating muscle fibres . . . . . . 139
B. Physiological mechanism of denervation fibrillations 139
C. Factors affecting fibrillation activity 141
I. Pharmacological 141
2. Arrest of circulation . . . . . . 142
3. Physical factors . . . . . . . . 143
a) Mechanical irritation due to insertion 143
b) The effect of muscle stretch . 144
c) Pressure and muscle massage 145
d) Electric current . . . . . . 145
e) The effect of temperature . . 146
D. Relationship of denervation fibrillations to other symptoms of muscle denervation 14 7
1. Relation of fibrillations to muscle atrophy . . . . . . . . . . . . . 147
2. Relation of fibrillations to changes in muscle excitability . . . . . . . 147
3. Relation of increased oxygen consumption to denervation fibrillations 148
VI. Electrolytes in Denervated Muscle (Z. Drahota)
Electrolyte composition of denervated muscle 151
A. Water and extracellula? ions in the denervated muscle 152
1. Changes in water content after denervation . . . . 152
2. Changes in sodium and chloride, and calculations of extra- and intracellular
water . . . . . . . . . . . . . . . . . . . . 153
3. Changes in Na, Cl and water during reinnervation . . 157
B. Intracellular ions in the denervated muscle . . . . . . 159
I. Changes in Mg and Ca ions in the denervated muscle 159
2. Changes in potassium content shortly after denervation 160
3. Potassium changes in the course of denervation atrophy 162
4. Potassium turnover in denervated and reinnervated muscles 164
C. Conclusions . . . . . . . . . . . . . . . . . . . . . . . 168
VII. Vasomotor Mechanism in Genesis of Denervation Atrophy (0. Hudlicka)
A. Blood flow through denervated muscle 175
B. Uptake of substances in the muscle 192
VIII. Energy Metabolism in Denervated Muscle (A. Bass)
A. Energy metabolism in normal muscle. . . . . 203
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I. Introduction . . . . . . . . . . . . . . . . . . . 203
2. Sources of energy in muscle at rest and their utilization. 203
3. Sources of energy in muscle during and after work 207
4. Formation of energy stores in the muscle 210
B. Energy metabolism in denervated muscle . . 212
I. A survey of disturbances in muscle energy metabolism following denervation 212
a) Historical survey . . . . . . . . . . . . . . . . . . . . . . . . . 213
h) Content of metabolites following denervation . . . . . . . . . . . . 215
c) Changes in enzyme activity associated with energy metabolism following
denervation . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
d) The consumption o£ substrates and formation of products of energy meta
bolism in the denervated muscle . . . . . . . . . . . . . . . . . . 223
e) Metabolic changes in the denervated muscle under conditions of "metabolic
loads" 230
aa) Starvation . . . . . . . 231
bb) Food intake . . . . . . 233
cc) Direct muscle stimulation 235
dd) Recovery processes after stimulation 238
ee) Changes in energy metabolism during reduced blood flow 242
ff) Conclusions concerning metabolic loads . . . . . . . . 216
f) The use of labelled compounds . . . . . . . . . . . . . . 246
g) Special methods for investigating energy metabolism in denervated muscle 251
h) Conclusions of the survey . . . . . . . . . . . . . . . . . . 253
2. Character and basis of energy metabolism of muscle after denervation 254
a) Disturbances in substrate supply . . 254
h) Disturbances of substrate utilization . 254
aa) Fibrillation activity . . . . . . . 255
bb) Formation of surplus energy stores 256
cc) Degradation of proteins . . . . . 257
dd) Possible alternate metabolic pathways 257
ee) Increased heat production and disturbances of oxidative phosphoryl-
ations . . . . . . . . . . . . . . . . . . . . 258
ff) Conclusions concerning the utilization of substrates 261
c) Questions ofthe primary link . . . . . . . . . . . 261
C. Main conclusions, problems and aims of research on energy metabolism in the
nervous regulation of muscle metabolism . . . . . . . . . . . . . . . . 264
IX. Proteins in the Denervated Muscle, Changes in their Quality, Properties
and Metabolism (R. ~ak)
A. Changes in protein composition of the denervated muscle 274
I. Basic indicators . .
a) Net changes . . 275
b) Relative changes 278
2. Specific proteins 281
a) Structural proteins 282
b) Sarcoplasmatic proteins 287
c) Interstitial muscle phase 291
3. Denervation hypertrophy 294
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B. Changes in the physico-chemical properties of denervated muscle proteins . 296
1. Solubility and extractihility 296
2. Double refraction of flow (DRF) 297
3. Viscosity . . . . . . . . . . . 298
4. Changes in the electron microscope 299
5. Enzymatic properties . . . . . . 299
C. Disturbances of protein metabolism in the denervated muscle 302
1. Synthesis and degradation of proteins in the normal muscle . 303
a) Proteosynthesis. . . . . . . . . . . . . . . . . . . 304
h) Protein degradation . . . . . . . . . . . . . . . . 308
2. Functional activity and muscle protein metabolism . . . . . . . . . . . 311
a) Changes of physico-chemical properties of muscle proteins 311
h) Protein degradation and muscle function . . . . . . . 313
c) Protein resynthesis following muscle function. . . . . . 323
3. Balance between protein degradation and synthesis in the denervated muscle 326
a) Protein degradation . . . . . . . . 326
h) Incorporation studies . . . . . . . . . . . . . . . . . . . 327
c) Measurements of net protein synthesis . . . . . . . . . . . 330
d) The relationship between synthesis of proteins and nucleic acids 332
X. Rate of Denervation Muscle Atrophy (P. Hnik)
A. Biometric problems associated with the evaluation of muscle atrophy . 341
1. Inaccuracies of currently used criteria for evaluating the rate of muscle atrophy 342
2. Investigation of atrophic gradients. . . . . . . . . . . . . . . . . 345
B. Differences in the rate of denervation atrophy in different animal species.. 352
C. Changes in the rate of denervation atrophy in various muscles . 353
I. Differences between normal muscles 354
a) Functional differences . . 354
h) Morphological differences . . . 354
c) Biochemical differences . . . . 355
d) Differences in the reactivity of muscles to humoral agents .. 355
e) Vascular supply . . . . . . . . . 356
f) Differences in innervation . . . . . 356
2. Differences between denervated muscles 358
a) Differences in colour . 358
h) Functional differences . 358
c) Biochemical changes . 358
d) Rate of atrophy . . . 359
D. Methods affecting the rate of denervation atrophy 360
l. Methods affecting muscle specific function 361
a) Passive movements and massage . . 361
h) Electrotherapy . . . . . . . . . . 362
2. Methods influencing muscle metabolism 168
a) Methods increasing blood flow . . . J68
h) Hormones accelerating muscle atrophy ~68
c) Hormones retarding muscle atrophy. E5 9
d) ATP and other substances . . . . . . 370
XI. Metabolic Reactibility of the Denervated Muscle (E. Gutmann)
A. Denervation studies in relation to problems of nervous regulation of metabolism 377
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