Jagadis Chandra Bose

“ It will thus be seen, that external stimulus of electric shock induces a reaction which is of opposite sign to the normal growth elongation or expansion. We may conveniently describe this effect as ‘incipient’ contraction; for under increasing intensity of stimulus, the contractile reaction, opposing growth elongation, becomes more and more pronounced; at an intermediate stage this results in an arrest of growth; at the further stage, it culminates in an actual shortening of the organ. ”

“ Taking first the case of the pulvinus of Mimosa, we find that it responds to mechanical stimulation, to constant electric current, to induction shock, to the action of chemical agents, to light, and to warmth as differentiated from thermal radiation. The reactions induced by these agents may be similar or dissimilar. An identical agent, again, may give rise to movements which are not merely different, but sometimes even of diametrically opposite characters. Certain organs, for example, direct themselves towards light, others away from it. ”

“ In the second, or MOVING PLATE METHOD, a curve is obtained, the ordinate representing growth elongation, and the abscissa, time. A stimulating agent causes an upward flexure of the normal curve; a depressing agent, on the other hand, lessens the slope of the curve.
The action of external stimulus induces a variation of the rate of growth, the time relations of which are found from the automatic record of the growth. The latent period is shortened with the intensity of the stimulus. A responsive variation of growth is induced by an intensity of stimulus which is below human perception. ”

“ From the facts given above it will be seen that the diurnal movement is not brought about by variation in transpiration. I now turn to another phenomenon which appeared at first to have some connection with the movement of the tree. Kraus found that the tissue tensions of a shoot exhibit a daily periodicity. He, however, found that between 10°C. and 30°C., variation of temperature had no effect on the daily period. But as regards the diurnal movement of the tree, it is the temperature which is the principal factor. ”

“ As an example of this may be cited the periodic variation of growth. Here the daily periodicity exhibited by a plant is not only different in varying seasons, but it also differs in diverse species of plants. The complexity of the problem is very great, for not only are the direct effects of the changing environment to be taken into consideration but also their unknown after-effects. Even in the case of direct effect, different factors, such as light, temperature, turgor, and so on, are undergoing independent variations ”

“ I shall in the course of the following series of Papers, describe the different apparatus by which the movement of pulvinated organ and its time-relations are automatically recorded. In a growing organ the induced movement under stimulus is brought about by the change in its rate of growth. That the change is solely due to the particular stimulus can only be assured by strict maintenance of constancy of external conditions, during the period of experiment ”

“ It has been shown that stimulus induces simultaneously both A- and D-effects, with the attendant positive and negative responsive reactions, alike in pulvinated and in growing organs. A tissue, in an optimum condition, exhibits only the resultant negative response; the comparatively feeble positive is imperceptible, being masked by the predominant negative; but with the decline of its tone ex­cit­abil­ity diminishes, with it the D-effect, and we get the A-effect unmasked, resulting response then becomes diphasic. In extreme sub-tonic condition, it exhibits only the positive. ”

“ There may be a difficulty on account of the diurnal movement of Mimosa; the leaf, generally speaking, has a movement in a downward direction from morning till noon, after which there is a comparative state of rest. It is better to choose the time of noon for experiment. In any case the response to stimulus is very abrupt and in strong contrast with the slow diurnal movement. A horizontal pencil of light was thrown upwards by means of a small mirror and made to fall on the lower half of a pulvinus of the Mimosa leaf. ”

“ Experiment 26.—It is obvious that the mechanical moment exerted by the weight of the leaf must help its responsive fall under excitation. But the relative importance of the factors of active contraction of the lower half of the pulvinus and of the weight, in the rate of the responsive down-movement, still remains to be determined. A satisfactory way of solving the problem would lie in the study of the characteristics of response-records taken under three different conditions: (1) When the leaf is helped in its fall by its weight; (2) when the action of the weight is eliminated ”

“ In a tissue whose conduct­ing power is not great, the excitatory impulse is weakened, even to extinction in trans­mission through a long distance. Thus the negative impulse may fail to reach the responding organ, when the stimulus is feeble or the intervening distance long or semi-conduct­ing. Hence, under the above conditions, stimulus applied at a distance will give rise only to a positive response. ”

“ Certain agents induce excitation in living tissues, the excitatory change being detected by contraction, or by electromotive variation, or by change of electric resistance, and in growing organs by the retardation of the rate of growth. ”

“ In Mimosa, the responsive movement is brought about by a sudden diminution of turgor in the pulvinus. But very little is definitely known about the responsive reaction in growing organs. Thus in a tendril, one-sided contraction causes a shortening of the concave side and a sudden increase of growth on the convex. No explanation of this difference has hitherto been forthcoming. Under the action of light of different intensities a growing organ may approach the source of light, or place itself at right angles or move away from it. ”

“ The sensitiveness of Mimosa to electric stimulation is very great; the plant often responds to a shock which is quite imperceptible to a human subject. By the employment of a sliding induction coil, the intensity of the shock can be regulated with great accuracy; the secondary if gradually brought nearer the primary till a stimulus is found which is minimally effective. The intensity of stimulus actually employed is slightly higher than this, but within the sub-maximal range. ”

“ Millardet observed a daily periodicity of tension in Mimosa pudica. He found that maximum tension occurs before dawn; the petiole becomes erected, the movement being upwards or towards the tip of the stem. Tension decreases during the day, and reaches a minimum early in the evening; in correspondence with this is the fall of the petiole, the movement being away from the tip of the stem. [C] If the plant were placed upside down the periodic movement of the petiole in relation to the stem will evidently remain the same, but become reversed in space. ”

“ A very troublesome complication of pulsations in growth, arises at high temperatures, which render further record extremely difficult. Growth is undoubtedly a pulsatory phenomenon; but under favourable circumstances, these merge practically into a continuous average rate of elongation. At a high temperature the effect of certain disturbing factors comes into prominence. ”

“ A pot-specimen of Mimosa produces relatively few leaves, but one grown in the open ground is extremely luxuriant. As an instance in point, I may state that for the last five months I have taken from a plant grown in a field about 20 leaves a day for experiment, without making any impression on it. A large box containing soil would be practically as good as the open ground, and the slower rate of growth in a colder climate could be easily made up by planting half a dozen specimens. ”

“ Looked at from above, the opening outwards during rise of temperature is a movement downwards, and therefore belongs to the negative type. In such cases the changed rate of growth by variation of temperature is the most important factor in the movement. It may be asked whether all thermonastic movements must necessarily belong to the negative type, where rise of temperature is attended by a movement downwards. ”

“ I also determined the relative values of the minimal intensity of stimulus, effective in causing trans­mission of excitation in the three cases. Adopting as before the intensity of electric stimulus which causes bare perception in a human being as the unit, I find that the effective stimulus for a fully grown young petiole is 0.3 unit, while the very young required 2.5 units, and the very old 5 units. Hence it may be said that the conduct­ing power of a very young is an eighth, and of the very old one-sixteenth of the conductivity of the fully grown young specimen. ”

“ A given plant-tissue may exist in widely different conditions of tonicity. Let us take two extreme conditions, the optimum and the minimum. The tonic level will be at its lowest at the minimum, where growth will be at a standstill. The range between the optimum and minimum will be very extended; hence strong and long continued stimulation will be necessary to raise the tissue from the tonic minimum to the optimum level. There are innumerable grades of tonicity between the optimum and minimum. Within this wide range the char­ac­ter­is­tic response will be the abnormal positive. ”

“ As we approach the optimum, the range for positive response will become circumscribed, and the intensity and duration of stimulus necessary to convert the positive to negative will be feebler and shorter. It will be very seldom that a plant is likely to be found at the optimum. Hence plants in general may be expected to give a feeble positive response under sub-minimal stimulus. ”

“ A given stimulus will bring about an intense excitation, and the ‘down’ D-change will therefore be very much greater than the A-change. Let us now consider the case at the opposite extreme where, owing to unfavourable condition, the ex­cit­abil­ity is at its lowest. Under stimulus the excitatory D-change will now be relatively feeble compared to the A-change, by which the potential energy of the system becomes increased. In such a case successive stimuli will increase the functional activity of the tissue, and bring about staircase response. ”