TRANSACTIONS
OF THE
BOSE RESEARCH INSTITUTE, CALCUTTA,
VOL. II, 1919
LIFE MOVEMENTS IN PLANTS
BY
SIR JAGADIS CHUNDER BOSE, Kt., M.A., D.Sc., C.S.I., C.I.E.,
PROFESSOR EMERITUS, PRESIDENCY COLLEGE,
DIRECTOR, BOSE RESEARCH INSTITUTE,
WITH 128 ILLUSTRATIONS
CALCUTTA
BENGAL GOVERNMENT PRESS
1919
PUBLISHED BY
THE BOSE RESEARCH INSTITUTE, CALCUTTA.
WORKS BY THE SAME AUTHOR.
RESPONSE IN THE LIVING AND NON-LIVING. With 117 Illustrations, 8vo. 10s. 6d. 1902
PLANT RESPONSE: AS A MEANS OF PHYSIOLOGICAL INVESTIGATION. With 278 Illustrations, 8vo. 21s. 1906
COMPARATIVE ELECTRO-PHYSIOLOGY. A PHYSICO-PHYSIOLOGICAL STUDY. With 406 Illustrations, 8vo. 15s. 1907
RESEARCHES ON IRRITABILITY OF PLANTS. With 190 Illustrations, 8vo. 10s. 6d. net 1913
LIFE MOVEMENTS IN PLANTS, VOL. I. With 92 Illustrations, 8vo. 10s. 6d. 1918
Longmans, Green & Co.
London, New York, Bombay and Calcutta.
PREFACE TO VOLUME II.
I have in the present volume dealt with the intricate phenomena of different tropisms. The movements in plants under the stimuli of the environment—the twining of tendrils, the effect of temperature, the action of light inducing movements sometimes towards and at other times away from the stimulus, the diametrically opposite responses of the shoot and the root to the same stimulus of gravity, the day and night positions of organs of plants—these, and many others present such diversities that it must have appeared a hopeless endeavour to discover any fundamental reaction applicable in all cases. It has therefore been customary to assume different sensibilities especially evolved for the advantage of the plant. But teleological argument and the use of descriptive phrases, like positive and negative tropism, offer no real explanation of the phenomena. Thus to quote Pfeffer "When we say that an organ curves towards a source of illumination, because of its heliotropic irritability we are simply expressing an ascertained fact in a conveniently abbreviated form, without explaining why such curvature is possible or how it is produced.... Many observers have unfortunately devoted their attention to artificially classifying the phenomenon observed, and have entirely neglected the explanation of causes underlying them." He also adds that in regard to the phenomenon of growth and its variations, an empirical treatment is all that is possible in the present state of our knowledge; but deduction from results of experimental investigation "still remains the ideal of physiology, and only when this ideal has been attained, shall we be able to obtain a comprehensive view of the interacting factors at work in the living organism."
In my previous work on "Plant Response" (1906) I described detailed investigations on irritability of plants which I carried out with highly sensitive recorders. The plant was thus made to tell its own story by means of its self-made records. The results showed that there is no specific difference in physiological reaction of different organs to justify the assumption of positive and negative irritabilities. A generalisation was obtained which gave a complete explanation of diverse movements in plants. The results were fully confirmed by an independent method of inquiry, namely that of electric response, which I have been able to elaborate so as to become a very important means of research.
The investigations described in the present volume not only support the conclusions reached in my earlier works, but have led to important additions. It is evident that the range of our investigation is limited only by our power of recording the rate of plant-movement, that is to say, in the measurement of length and time. In these respects the instruments that I have been able to devise have surpassed my sanguine expectations. The Resonant Recorder traces time-intervals as short as a thousandth part of a second, while my Balanced Crescograph enables us to measure variation of rate of growth as minute as 1⁄1000 millionth of an inch per second, the sensitiveness of this apparatus thus rivals that of the spectroscope. The increasing refinement in our experimental methods cannot but lead to important advances towards a deeper understanding of underlying reactions in the living organism.
I shall here draw attention to only a few of the important results given in the present volume. The tropic effect of light has been shown to have a definite relation to the quantity of incident light. A complete tropic curve has been obtained from sub-minimal to maximal stimulation which shows the inadequacy of Weber's law, for the sub-minimal stimulus induces a qualitative difference in physiological reaction. It has further been shown that the prevalent idea that perception and heliotropic excitation are two distinct phenomena is without any foundation.
With reference to the effect of ether waves on plants, I have given an account of my discovery of the response of all plants to wireless stimulation, the results being similar to that induced by visible light. The perceptive range of the plant is thus infinitely greater than ours; for it not only perceives, but also responds to different rays of the vast ethereal spectrum.
The results obtained by the method of geo-electric response show that the responsive reaction of the root is in no way different from that of the shoot, the opposite movements being due to the fact that in the shoot the stimulation is direct, and in the root it is indirect.
Full description is given of the new method of physiological exploration by means of the electric probe, by which the particular layer which perceives the stimulus of gravity is definitely localised. The method of electric probe is also found to be of extended application in the detection of physiological changes in the interior of an organ.
An important factor of nyctitropic movements, hitherto unsuspected, is the effect of variation of temperature on geotropic curvature. This and other co-operative factors have been fully analysed, and a satisfactory explanation has been offered of various types of diurnal movement.
A generalisation has been obtained which explains all the diverse movements of plants, under all modes of stimulation: it has been shown that direct stimulation induces contraction and retardation of growth, and that indirect stimulation induces an expansion and acceleration of growth.
Another generalisation of still greater importance is the establishment of identical nature of physiological reaction in the plant and the animal, leading to advances in general physiology. Thus the discovery of a method for immediate enhancement or inhibition of nervous impulse in the plant led to my success in the control of nervous impulse in the animal. Another important discovery was the dual nervous impulses in plants, and I have very recently been able to establish, that the nervous impulse generated in the animal nerve by stimulus is not single, but double.
The study of the responsive phenomena in plants must thus form an integral part of physiological investigation into various problems relating to the irritability of all living tissues, and without such study the investigation must in future remain incomplete.
J. C.
October 1919.
CONTENTS.
TROPISM IN PLANTS. PAGE. XXII.—THE BALANCED CRESCOGRAPH. Principle of the Method of Balance—Compensating movement—Growth-scale—Sensitiveness of the Crescographic Balance—Effect of CO2—Effect of anæsthetics 255 XXIII.—ON TROPIC MOVEMENTS. Complexity of the problem—Contradictory nature of responses—Two classes of tropic responses—Longitudinal transmission of effect of stimulus—Transverse transmission of effect of stimulus—Modification of tropic curvature by conducting power of tissues and differential excitability of the organ 268 XXIV.—TROPIC CURVATURE WITH LONGITUDINAL TRANSMISSION OF EFFECT OF STIMULUS. Dual impulses, positive and negative, caused by stimulus—Direct and Indirect stimulus—Tropic effect of Indirect stimulation 271 XXV.—TROPIC CURVATURE WITH TRANSVERSE TRANSMISSION OF EFFECT OF STIMULUS. Turgor variation under transverse transmission of stimulus-effect—Tropic responses of pulvinated and growing organs to unilateral stimulation—Direct unilateral stimulation—Indirect unilateral stimulation—Difference of effects induced by Direct and Indirect stimulation—Laws of tropic curvature 279 XXVI.—MECHANOTROPISM: TWINING OF TENDRILS. Anomalies of mechanotropism—Effects of indirect and direct electric stimulation on growth of tendril—Effect of direct and indirect mechanical stimulus—Immediate and after-effect of stimulus—Inhibitory action of stimulus—Response of less excitable side of the tendril—Relative intensities of responses of upper and under sides of tendril of Passiflora—Negative curvature of tendril 288 XXVII.—ON GALVANOTROPISM. Polar effects of electric current on growth—Effect of anode and cathode on growth 301 XXVIII.—ON THERMONASTIC PHENOMENA. Effect of temperature—Different thermonastic organs—Two types of response: Positive and Negative—Effect of rise of temperature and of stimulus on thermonastic organs—Law of thermonastic reaction 305 XXIX.—ON PHOTOTROPISM. Complexity of problem of phototropic reaction—Action of light—Positive phototropic curvature of pulvinated organs—Positive phototropic curvature of growing organs—Phenomenon of recovery—Immediate and after-effect of light on growth—Latent period of phototropic reaction—Growth variation induced by flash of light from a single spark—Maximum positive curvature under continued action of light 313 XXX.—DIA-PHOTOTROPISM AND NEGATIVE PHOTOTROPISM. Differential excitability of two halves of pulvinus of Mimosa—Transformation of positive to negative curvature—Tropic effect under sunlight—Negative phototropism of shoot and root 328 XXXI.—RELATION BETWEEN THE QUANTITY OF LIGHT AND THE INDUCED PHOTOTROPIC CURVATURE. Effect of increasing intensity of light on pulvinated and growing organs—Effect of increasing angle—Effect of duration of exposure 338 XXXII.—THE PHOTOTROPIC CURVE AND ITS CHARACTERISTICS. Summation of stimulus—General consideration—The general characteristic curve—Characteristics of simple phototropic curve—Variation of susceptibility for excitation in different parts of the curve—Effect of sub-minimal stimulus—The complete phototropic curves of pulvinated and growing organs—Limitation of Weber's law 346 XXXIII.—TRANSMITTED EFFECT OF PHOTIC STIMULATION. Effect of light applied on tip of Setaria—Response of growing region to unilateral stimulus—Effect of simultaneous stimulation of the tip and the hypocotyl—Algebraical summation of effects of direct and indirect stimuli 362 XXXIV.—ON PHOTONASTIC CURVATURES. Phototropic response of anisotropic organs—Positive para-heliotropism—Negative para-heliotropism—Responses of pulvinated and growing organs to light 378 XXXV.—EFFECT OF TEMPERATURE ON PHOTOTROPIC CURVATURE. Effect of temperature on excitability—Effect of temperature on conduction—Phototropic response of tendrils—Seasonal variation of phototropic curvature—Antagonistic effects of light and of rise of
NIGHT AND DAY MOVEMENTS IN PLANTS. XLVI.—DIURNAL MOVEMENTS IN PLANTS. Complexity of the problem—The different factors involved—Autonomous movements—Epinasty and hyponasty—Positive and negative thermonasty—Thermo-geotropism—Classification of diurnal movements—Discriminating tests for classification—Diurnal variation of light and of temperature 523 XLVII.—DIURNAL MOVEMENT DUE TO ALTERNATION OF LIGHT AND DARKNESS. Experimental arrangements—The Quadruplex Nyctitropic Recorder—Diurnal movement of the leaflet of Cassia alata—Effect of variation of temperature—Effect of variation of light—Diurnal movement of the terminal leaflet of Desmodium gyrans—The 'midday sleep' 535 XLVIII.—DIURNAL MOVEMENT DUE TO VARIATION OF TEMPERATURE AFFECTING GROWTH. Tropic and nastic movements—Distinction between thermonastic and