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RESPONSE IN THE LIVING
AND NON-LIVING
BY
JAGADIS CHUNDER BOSE, M.A.(CANTAB.), D.Sc.(LOND.)
PROFESSOR, PRESIDENCY COLLEGE, CALCUTTA
WITH ILLUSTRATIONS
LONGMANS, GREEN, AND CO.
39 PATERNOSTER ROW, LONDON
NEW YORK AND BOMBAY
1902
All rights reserved
_'The real is one: wise men call it variously'_
RIG VEDA
_To my Countrymen_
_This Work is Dedicated_
PREFACE
I have in the present work put in a connected and a more complete form
results, some of which have been published in the following Papers:
'De la Généralité des Phénomènes Moléculaires produits par l'Electricité
sur la matière Inorganique et sur la matière Vivante.' (_Travaux du
Congrès International de Physique._ Paris, 1900.)
'On the Similarity of Effect of Electrical Stimulus on Inorganic and
Living Substances.' (_Report, Bradford Meeting British Association,
1900.--Electrician._)
'Response of Inorganic Matter to Stimulus.' (Friday Evening Discourse,
Royal Institution, May 1901.)
'On Electric Response of Inorganic Substances. Preliminary Notice.'
(Royal Society, June 1901.)
'On Electric Response of Ordinary Plants under Mechanical Stimulus.'
(_Journal Linnean Society_, 1902.)
'Sur la Réponse Electrique dans les Métaux, les Tissus Animaux et
Végétaux.' (Société de Physique, Paris, 1902.)
'On the Electro-Motive Wave accompanying Mechanical Disturbance in
Metals in contact with Electrolyte.' (_Proceedings Royal Society_,
vol. 70.)
'On the Strain Theory of Vision and of Photographic Action.' (_Journal
Royal Photographic Society_, vol. xxvi.)
These investigations were commenced in India, and I take this
opportunity to express my grateful acknowledgments to the Managers of
the Royal Institution, for the facilities offered me to complete them at
the Davy-Faraday Laboratory.
J. C. BOSE.
DAVY-FARADAY LABORATORY, ROYAL INSTITUTION,
LONDON: _May 1902._
CONTENTS
CHAPTER I
THE MECHANICAL RESPONSE OF LIVING SUBSTANCES
PAGE
Mechanical response--Different kinds of stimuli--Myograph
--Characteristics of response-curve: period, amplitude,
form--Modification of response-curves 1
CHAPTER II
ELECTRIC RESPONSE
Conditions for obtaining electric response--Method of
injury--Current of injury--Injured end, cuproid: uninjured,
zincoid--Current of response in nerve from more excited to
less excited--Difficulties of present nomenclature--Electric
recorder--Two types of response, positive and
negative--Universal applicability of electric mode of
response--Electric response a measure of physiological
activity--Electric response in plants 5
CHAPTER III
ELECTRIC RESPONSE IN PLANTS--METHOD OF NEGATIVE VARIATION
Negative variation--Response recorder--Photographic
recorder--Compensator--Means of graduating intensity of
stimulus--Spring-tapper and torsional vibrator--Intensity of
stimulus dependent on amplitude of vibration--Effectiveness
of stimulus dependent on rapidity also 17
CHAPTER IV
ELECTRIC RESPONSE IN PLANTS--BLOCK METHOD
Method of block--Advantages of block method--Plant response a
physiological phenomenon--Abolition of response by
anæsthetics and poisons--Abolition of response when plant is
killed by hot water 27
CHAPTER V
PLANT RESPONSE--ON THE EFFECTS OF SINGLE STIMULUS AND OF
SUPERPOSED STIMULI
Effect of single stimulus--Superposition of stimuli--Additive
effect--Staircase effect--Fatigue--No fatigue when sufficient
interval between stimuli--Apparent fatigue when stimulation
frequency is increased--Fatigue under continuous stimulation 35
CHAPTER VI
PLANT RESPONSE--ON DIPHASIC VARIATION
Diphasic variation--Positive after-effect and positive
response--Radial E.M. variation 44
CHAPTER VII
PLANT RESPONSE--ON THE RELATION BETWEEN STIMULUS AND RESPONSE
Increased response with increasing stimulus--Apparent diminution
of response with excessively strong stimulus 51
CHAPTER VIII
PLANT RESPONSE--ON THE INFLUENCE OF TEMPERATURE
Effect of very low temperature--Influence of high
temperature--Determination of death-point--Increased response
as after-effect of temperature variation--Death of plant and
abolition of response by the action of steam 59
CHAPTER IX
PLANT RESPONSE--EFFECT OF ANÆSTHETICS AND POISONS
Effect of anæsthetics, a test of vital character of
response--Effect of chloroform--Effect of chloral--Effect of
formalin--Method in which response is unaffected by variation
of resistance--Advantage of block method--Effect of dose 71
CHAPTER X
RESPONSE IN METALS
Is response found in inorganic substances?--Experiment on tin,
block method--Anomalies of existing terminology--Response by
method of depression--Response by method of exaltation 81
CHAPTER XI
INORGANIC RESPONSE--MODIFIED APPARATUS TO EXHIBIT RESPONSE IN
METALS
Conditions of obtaining quantitative measurements--Modification
of the block method--Vibration cell--Application of
stimulus--Graduation of the intensity of
stimulus--Considerations showing that electric response is
due to molecular disturbance--Test experiment--Molecular
voltaic cell 91
CHAPTER XII
INORGANIC RESPONSE--METHOD OF ENSURING CONSISTENT RESULTS
Preparation of wire--Effect of single stimulus 100
CHAPTER XIII
INORGANIC RESPONSE--MOLECULAR MOBILITY: ITS INFLUENCE ON RESPONSE
Effects of molecular inertia--Prolongation of period of recovery
by overstrain--Molecular model--Reduction of molecular
sluggishness attended by quickened recovery and heightened
response--Effect of temperature--Modification of latent
period and period of recovery by the action of chemical
reagents--Diphasic variation 104
CHAPTER XIV
INORGANIC RESPONSE--FATIGUE, STAIRCASE, AND MODIFIED RESPONSE
Fatigue in metals--Fatigue under continuous
stimulation--Staircase effect--Reversed responses due to
molecular modification in nerve and in metal, and their
transformation into normal after continuous
stimulation--Increased response after continuous stimulation 118
CHAPTER XV
INORGANIC RESPONSE--RELATION BETWEEN STIMULUS AND
RESPONSE--SUPERPOSITION OF STIMULI
Relation between stimulus and response--Magnetic
analogue--Increase of response with increasing
stimulus--Threshold of response--Superposition of
stimuli--Hysteresis 131
CHAPTER XVI
INORGANIC RESPONSE--EFFECT OF CHEMICAL REAGENTS
Action of chemical reagents--Action of stimulants on
metals--Action of depressants on metals--Effect of 'poisons'
on metals--Opposite effect of large and small doses 139
CHAPTER XVII
ON THE STIMULUS OF LIGHT AND RETINAL CURRENTS
Visual impulse: (1) chemical theory; (2) electrical
theory--Retinal currents--Normal response positive--Inorganic
response under stimulus of light--Typical experiment on the
electrical effect induced by light 148
CHAPTER XVIII
INORGANIC RESPONSE--INFLUENCE OF VARIOUS CONDITIONS ON THE
RESPONSE TO STIMULUS OF LIGHT
Effect of temperature--Effect of increasing length of
exposure--Relation between intensity of light and magnitude
of response--After-oscillation--Abnormal effects: (1)
preliminary negative twitch; (2) reversal of response; (3)
transient positive twitch on cessation of light; (4) decline
and reversal--Résumé 158
CHAPTER XIX
VISUAL ANALOGUES
Effect of light of short duration--After-oscillation--Positive
and negative after-images--Binocular alternation of
vision--Period of alternation modified by physical
condition--After-images and their revival--Unconscious visual
impression. 170
CHAPTER XX
GENERAL SURVEY AND CONCLUSION 181
INDEX 193
ILLUSTRATIONS
FIG. PAGE
1. MECHANICAL LEVER RECORDER 3
2. ELECTRIC METHOD OF DETECTING NERVE RESPONSE 6
3. DIAGRAM SHOWING INJURED END OF NERVE CORRESPONDS TO 8
COPPER IN A VOLTAIC ELEMENT
4. ELECTRIC RECORDER 11
5. SIMULTANEOUS RECORD OF MECHANICAL AND ELECTRICAL 13
RESPONSES
6. NEGATIVE VARIATION IN PLANTS 19
7. PHOTOGRAPHIC RECORD OF NEGATIVE VARIATION IN PLANTS 20
8. RESPONSE RECORDER 21
9. THE COMPENSATOR 22
10. THE SPRING-TAPPER 23
11. THE TORSIONAL VIBRATOR 24
12. RESPONSE IN PLANT TO MECHANICAL TAP OR VIBRATION 25
13. INFLUENCE OF SUDDENNESS ON THE EFFICIENCY OF 26
STIMULUS
14. THE METHOD OF BLOCK 28
15. RESPONSE IN PLANT COMPLETELY IMMERSED UNDER WATER 29
16. UNIFORM RESPONSES IN PLANT 36
17. FUSION OF EFFECT UNDER RAPIDLY SUCCEEDING STIMULI IN 36
MUSCLE AND IN PLANT
18. ADDITIVE EFFECT OF SINGLY INEFFECTIVE STIMULI ON 37
PLANT
19. 'STAIRCASE EFFECT' IN PLANT 37
20. APPEARANCE OF FATIGUE IN PLANT UNDER SHORTENED 39
PERIOD OF REST
21. FATIGUE IN CELERY 40
22. FATIGUE IN CAULIFLOWER-STALK 41
23. FATIGUE FROM PREVIOUS OVERSTRAIN 41
24. FATIGUE UNDER CONTINUOUS STIMULATION IN CELERY 42
25. EFFECT OF REST IN REMOVAL OF FATIGUE IN PLANT 43
26. DIPHASIC VARIATION IN PLANT 46
27, 28. ABNORMAL POSITIVE RESPONSES IN STALE PLANT 48, 49
TRANSFORMED INTO NORMAL NEGATIVE UNDER STRONG
STIMULATION
29. RADIAL E.M. VARIATION 50
30. CURVES SHOWING THE RELATION BETWEEN INTENSITY OF 52
STIMULUS AND RESPONSE IN MUSCLE AND NERVE
31. INCREASING RESPONSES TO INCREASING STIMULI (TAPS) IN 52
PLANTS
32. INCREASING RESPONSES TO INCREASING VIBRATIONAL 53
STIMULI IN PLANTS
33. RESPONSES TO INCREASING STIMULI IN FRESH AND STALE 54
SPECIMENS OF PLANTS
34. APPARENT DIMINUTION OF RESPONSE CAUSED BY FATIGUE 57
UNDER STRONG STIMULATION
35. DIMINUTION OF RESPONSE IN EUCHARIS LILY AT LOW 61
TEMPERATURE
36. RECORDS SHOWING THE DIFFERENCE IN THE EFFECTS OF LOW 62
TEMPERATURE ON IVY, HOLLY, AND EUCHARIS LILY
37. PLANT CHAMBER FOR STUDYING THE EFFECT OF TEMPERATURE 64
AND ANÆSTHETICS
38. EFFECT OF HIGH TEMPERATURE ON PLANT RESPONSE 64
39. AFTER-EFFECT ON THE RESPONSE DUE TO TEMPERATURE 66
VARIATION
40. RECORDS OF RESPONSES IN EUCHARIS LILY DURING RISE 67
AND FALL OF TEMPERATURE
41. CURVE SHOWING VARIATION OF SENSITIVENESS DURING A 68
CYCLE OF TEMPERATURE VARIATION
42. RECORD OF EFFECT OF STEAM IN ABOLITION OF RESPONSE 69
AT DEATH OF PLANT
43. EFFECT OF CHLOROFORM ON NERVE RESPONSE 72
44. EFFECT OF CHLOROFORM ON THE RESPONSES OF CARROT 74
45. ACTION OF CHLORAL HYDRATE ON PLANT RESPONSES 75
46. ACTION OF FORMALIN ON RADISH 75
47. ACTION OF SODIUM HYDRATE IN ABOLISHING THE RESPONSE 78
IN PLANT
48. STIMULATING ACTION OF POISON IN SMALL DOSES IN 79
PLANTS
49. THE POISONOUS EFFECT OF STRONGER DOSE OF KOH 79
50. BLOCK METHOD FOR OBTAINING RESPONSE IN TIN 83
51. RESPONSE TO MECHANICAL STIMULATION IN A Zn-Cu COUPLE 85
52. ELECTRIC RESPONSE IN METAL BY THE METHOD OF RELATIVE 88
DEPRESSION (NEGATIVE VARIATION)
53. METHOD OF RELATIVE EXALTATION 89
54. VARIOUS CASES OF POSITIVE AND NEGATIVE VARIATION 90
55. MODIFICATIONS OF THE BLOCK METHOD FOR EXHIBITING 93
ELECTRIC RESPONSE IN METALS
56. EQUAL AND OPPOSITE RESPONSES GIVEN BY TWO ENDS OF 95
THE WIRE
57. TOP VIEW OF THE VIBRATION CELL 96
58. INFLUENCE OF ANNEALING IN THE ENHANCEMENT OF 101
RESPONSE IN METALS
59. UNIFORM ELECTRIC RESPONSES IN METALS 102
60. PERSISTENCE OF AFTER-EFFECT 105
61. PROLONGATION OF PERIOD OF RECOVERY AFTER OVERSTRAIN 106
62. MOLECULAR MODEL 107
63, 64. EFFECTS OF REMOVAL OF MOLECULAR SLUGGISHNESS IN 109, 110
QUICKENED RECOVERY AND HEIGHTENED RESPONSE IN METALS
65. EFFECT OF TEMPERATURE ON RESPONSE IN METALS 111
66. DIPHASIC VARIATION IN METALS 113
67. NEGATIVE, DIPHASIC, AND POSITIVE RESULTANT RESPONSE 115
IN METALS
68. CONTINUOUS TRANSFORMATION FROM NEGATIVE TO POSITIVE 116
THROUGH INTERMEDIATE DIPHASIC RESPONSE
69. FATIGUE IN MUSCLE 118
70. FATIGUE IN PLATINUM 118
71. FATIGUE IN TIN 119
72. APPEARANCE OF FATIGUE DUE TO SHORTENING THE PERIOD 120
OF RECOVERY
73. FATIGUE IN METAL UNDER CONTINUOUS STIMULATION 121
74. 'STAIRCASE' RESPONSE IN MUSCLE AND IN METAL 122
75. ABNORMAL RESPONSE IN NERVE CONVERTED INTO NORMAL 124
UNDER CONTINUED STIMULATION
76, 77. ABNORMAL RESPONSE IN TIN AND PLATINUM CONVERTED INTO 125
NORMAL UNDER CONTINUED STIMULATION
78. GRADUAL TRANSITION FROM ABNORMAL TO NORMAL RESPONSE 126
IN PLATINUM
79. INCREASE OF RESPONSE IN NERVE AFTER CONTINUOUS 127
STIMULATION
80, 81. RESPONSE IN TIN AND PLATINUM ENHANCED AFTER 127, 128
CONTINUOUS STIMULATION
82. MAGNETIC ANALOGUE 132
83, 84. RECORDS OF RESPONSES TO INCREASING STIMULI IN TIN 134, 135
85. INEFFECTIVE STIMULUS BECOMING EFFECTIVE BY 135
SUPERPOSITION
86. INCOMPLETE AND COMPLETE FUSION OF EFFECTS 136
87. CYCLIC CURVE FOR MAXIMUM EFFECTS SHOWING HYSTERESIS 137
88. ACTION OF POISON IN ABOLISHING RESPONSE IN NERVE 139
89. ACTION OF STIMULANT ON TIN 141
90. ACTION OF STIMULANT ON PLATINUM 142
91. DEPRESSING EFFECT OF KBr ON TIN 143
92. ABOLITION OF RESPONSE IN METALS BY 'POISON' 143
93. 'MOLECULAR ARREST' BY THE ACTION OF 'POISON' 145
94. OPPOSITE EFFECTS OF SMALL AND LARGE DOSES ON THE 146
RESPONSE IN METALS
95. RETINAL RESPONSE TO LIGHT 150
96. RESPONSE OF SENSITIVE CELL TO LIGHT 152
97. TYPICAL EXPERIMENT ON THE E.M. VARIATION PRODUCED BY 154
LIGHT
98. MODIFICATION OF THE PHOTO-SENSITIVE CELL 155
99. RESPONSES IN FROG'S RETINA 156
100. RESPONSES IN SENSITIVE PHOTO-CELL 157
101. EFFECT OF TEMPERATURE ON THE RESPONSE TO LIGHT 159
STIMULUS
102. EFFECT OF DURATION OF EXPOSURE ON THE RESPONSE 159
103. RESPONSES OF SENSITIVE CELL TO INCREASING 161
INTENSITIES OF LIGHT
104. RELATION BETWEEN THE INTENSITY OF LIGHT AND 162
MAGNITUDE OF RESPONSE
105. AFTER-OSCILLATION 163
106. TRANSIENT POSITIVE INCREASE OF RESPONSE IN THE 164
FROG'S RETINA ON THE CESSATION OF LIGHT
107. TRANSIENT POSITIVE INCREASE OF RESPONSE IN THE 165
SENSITIVE CELL
108. DECLINE UNDER THE CONTINUOUS ACTION OF LIGHT 166
109. CERTAIN AFTER-EFFECTS OF LIGHT 168
110. AFTER-EFFECT OF LIGHT OF SHORT DURATION 172
111. STEREOSCOPIC DESIGN FOR THE EXHIBITION OF BINOCULAR 176
ALTERNATION OF VISION
112. UNIFORM RESPONSES IN NERVE, PLANT, AND METAL 184
113. FATIGUE IN MUSCLE, PLANT, AND METAL 185
114. 'STAIRCASE' EFFECT IN MUSCLE, PLANT, AND METAL 186
115. INCREASE OF RESPONSE AFTER CONTINUOUS STIMULATION IN 186
NERVE AND METAL
116. MODIFIED ABNORMAL RESPONSE IN NERVE AND METAL 187
TRANSFORMED INTO NORMAL RESPONSE AFTER CONTINUOUS
STIMULATION
117. ACTION OF THE SAME 'POISON' IN THE ABOLITION OF 189
RESPONSE IN NERVE, PLANT, AND METAL
RESPONSE
IN THE
LIVING AND NON-LIVING
CHAPTER I
THE MECHANICAL RESPONSE OF LIVING SUBSTANCES
Mechanical response--Different kinds of
stimuli--Myograph--Characteristics of response-curve: period,
amplitude, form--Modification of response-curves.
One of the most striking effects of external disturbance on certain
types of living substance is a visible change of form. Thus, a piece of
muscle when pinched contracts. The external disturbance which produced
this change is called the stimulus. The body which is thus capable of
responding is said to be irritable or excitable. A stimulus thus
produces a state of excitability which may sometimes be expressed by
change of form.
#Mechanical response to different kinds of stimuli.#--This reaction under
stimulus is seen even in the lowest organisms; in some of the amoeboid
rhizopods, for instance. These lumpy protoplasmic bodies, usually
elongated while creeping, if mechanically jarred, contract into a
spherical form. If, instead of mechanical disturbance, we apply salt
solution, they again contract, in the same way as before. Similar
effects are produced by sudden illumination, or by rise of temperature,
or by electric shock. A living substance may thus be put into an
excitatory state by either mechanical, chemical, thermal, electrical, or
light stimulus. Not only does the point stimulated show the effect of
stimulus, but that effect may sometimes be conducted even to a
considerable distance. This power of conducting stimulus, though common
to all living substances, is present in very different degrees. While in
some forms of animal tissue irritation spreads, at a very slow rate,
only to points in close neighbourhood, in other forms, as for example in
nerves, conduction is very rapid and reaches far.
The visible mode of response by change of form may perhaps be best
studied in a piece of muscle. When this is pinched, or an electrical
shock is sent through it, it becomes shorter and broader. A responsive
twitch is thus produced. The excitatory state then disappears, and the
muscle is seen to relax into its normal form.
#Mechanical lever recorder.#--In the case of contraction of muscle, the
effect is very quick, the twitch takes place in too short a time for
detailed observation by ordinary means. A myographic apparatus is
therefore used, by means of which the changes in the muscle are
self-recorded. Thus we obtain a history of its change and recovery from
the change. The muscle is connected to one end of a writing lever. When
the muscle contracts, the tracing point is pulled up in one direction,
say to the right. The extent of this pull depends on the amount of
contraction. A band of paper or a revolving drum-surface moves at a
uniform speed at right angles to the direction of motion of the writing
lever. When the muscle recovers from the stimulus, it relaxes into its
original form, and the writing point traces the recovery as it moves now
to the left, regaining its first position. A curve is thus described,
the rising portion of which is due to contraction, and the falling
portion to relaxation or recovery. The ordinate of the curve represents
the intensity of response, and the abscissa the time (fig. 1).
[Illustration: FIG. 1.--MECHANICAL LEVER RECORDER
The muscle M with the attached bone is securely held at one end, the
other end being connected with the writing lever. Under the action
of stimulus the contracting muscle pulls the lever and moves the
tracing point to the right over the travelling recording surface P.
When the muscle recovers from contraction, the tracing point returns
to its original position. See on P the record of muscle curve.]
#Characteristics of the response-curve: (1) Period, (2) Amplitude, (3)
Form.#--Just as a wave of sound is characterised by its (1) period, (2)
amplitude, and (3) form, so may these response-curves be distinguished
from each other. As regards the period, there is an enormous variation,
corresponding to the functional activity of the muscle. For instance, in
tortoise it may be as high as a second, whereas in the wing-muscles of
many insects it is as small as 1/300 part of a second. 'It is probable
that a continuous graduated scale might, as suggested by Hermann, be
drawn up in the animal kingdom, from the excessively rapid contraction
of insects to those of tortoises and hibernating dormice.'[1]
Differences in form and amplitude of curve are well illustrated by
various muscles of the tortoise. The curve for the muscle of the neck,
used for rapid withdrawal of the head on approach of danger, is quite
different from that of the pectoral muscle of the same animal, used for
its sluggish movements.
Again, progressive changes in the same muscle are well seen in the
modifications of form which consecutive muscle-curves gradually undergo.
In a dying muscle, for example, the amplitude of succeeding curves is
continuously diminished, and the curves themselves are elongated.
Numerous illustrations will be seen later, of the effect, in changing
the form of the curve, of the increased excitation or depression
produced by various agencies.
Thus these response records give us a means of studying the effect of
stimulus, and the modification of response, under varying external
conditions, advantage being taken of the mechanical contraction produced
in the tissue by the stimulus. But there are other kinds of tissue where
the excitation produced by stimulus is not exhibited in a visible form.
In order to study these we have to use an altogether independent method,
the method of electric response.
FOOTNOTES:
[1] Biedermann, _Electro-physiology_, p. 59.