By the Same Author RESPONSE IN THE LIVING AND NON-LAVING, PLANT RESPONSE AS'A MKANS OF PLLYSIO- TORICAL INVESFIGATION, ‘COMPARATIVE. SST SS ee 8 RESEARCHES ON THE IRRITALILILY OK PEA 90 Mose. $9, 7. ot “THE FRYSIOLOGY OF THE ASCENY OF SAP. ‘THR PHYSIOLOGY OF PHOvosyNxinusis. FB MOVEMENTS 1M PLANTS. ‘THe MECHANISM OF PLANT RESVONSE, ‘THE LIFE AND WORK OF SLX JAGADIS 6. BOSE, pigment Bay, Une Ca Ei uaky ot amtay” Wiis Fone Sad anes, LONGMANS, GREEN AND Co. LTD, ‘THE i NERVOUS MECHANISM OF PLANTS SIR JAGADIS CHUNDER BOS WITH 82 ILLUSTRATIONS PO. Box 220 + Bayside, California 95524 USA. LONGMANS, GREEN AND CO, LTD. 39 PATEMNOSIER ROW, LONDON, EC. 4 NEW YORK, TORONTO BOMBAY, CALCUTTA, AND MADRAS 1926PREFACE ‘My investigations on plant-response date from the discovery of the electric response of non-living matter, such as metals, to stimulus, published in 1900 by the International Congress of Science, Paris. The response, like that of living matter, ‘was shown to exhibit fatigue under continuous stimulation, enhancement under chemical stimulants, and permanent abolition under poisons. These results indicated that the response of the more complex and unstable living matter is ultimately the expression of physico-chemical reactions. 1 next tried to find whether ordinary plants, meaning those usually regarded as insensitive, exhibit the characteristic elec- tric response already known in ‘sensitive’ plants. Ordinary plants were regarded at the time as inexcitable, because they did not respond to stimulation by an obvious movement. In my Friday Evening Discourse before the Royal Institution in May 1901, I was, however, able to show that every plant, and even each organ of every plant, is excitable, and responds to stimulus by electric response of galvanometric negativity, the response being abolished at the death of the plant. A more detailed account of the results was published in my work on ‘ Response of the Living and Non-Living’ (x902). My next investigation was directed towards obtaining evidence of responsive mechanical movement in these plants renderéd conspicuous by various devices of magnification (‘Plant Response,’ 1906). The effects of various environ- mental stimuli on different plant-organs were thus demon- strated by automatic records given by the plant. The mostviii PREFACE important fact established in plant-response was the nervous character of the impulse transmitted to a distance. My discovery of the excitatory polar action of an electriccurrent and its transmission to a distance, proved that theconduction of excitation in the plant is fundamentally the same as that in the nerve of the animal, Further corroboration was found in the arrest of transmission by the application of electrotonic and other physiological blocks in the path of ‘conduction. In my ‘Comparative Electro-Physiology * (r907) I ém- ployed the independent method of electric response and was able fully to confirm and extend the results which I had obtained by the method of mechanical response. Experi- ments are described showing that the response of isolated plant-nerveisindistinguishablefrom that of the animal nerve throughout a long series of parallel variations of condition, So complete, indeed, is the similarity, that the discovery of a responsive characteristic in a given case has proved a sure guide to its observation in the other. As an example of this may be mentioned the control of the nervous impulse in the animal by a homodromous or a heterodromous electric: current, the discovery being due to the success which attended my attempt to control the nervous impulse in the plant by the directive action of electric currents. Accurate measurement of the normal velocity of transmission of nervous impulse, and its induced variations, has been rendered possible by my device of Resonant Recorder, described in ' Irritability of Plants’ (r913), by which time. intervals as short as .00$ second can be automatically recorded. ‘The great advance in animal physiology has been due to very sensitive and accurate methods in quantitative measure- ment, on.which alone can any sound theory be based. In plant-physiology, unfortunately, no such methods had been Previously available; this accounts for the unfounded speculations that had paralysed advance of knowledge in plant-physiology. As an instance of this may be mentioned PREFACE ix the far-fetched theory recently advanced that the trans- mission of excitation in the sensitive plant, Mimosa pudica, is due to the movement of sap in the transpiration current. Simultaneous measurements of the rate of trans- mission of excitation and of the ascent of sap show that the former is far greater than the latter. In Mimosa the velocity of nervous impulse in thin petioles is as high as 400 mm. per second, while the movement of sap is about 200 times slower. No demonstration of the unfounded character of the transpiration-current theory could be more simple and convincing than the observation of the effect of application of a drop of acid to the tip of the uppermost leaf ‘of Mimosa described in Chapter II (p, x9). My recent discovery of the transformation of the afferent or sensory into an efferent or motor impulse in the reflex arc inthe pulvinus of Mimosa will, it ishoped, materially advance our knowledge of nervous impulse in general. I have, in the present work, given not only a connected account of my previous results which came out in scattered publications, but also a mass of new material that has been accumulated since the discovery of separate nerves for the conduction of sensory and motor impulses. The results of the investigations which I have carried out for the last quarter of a century establish the generalisa- tion that the physiological mechanism of the plant is identical with that of the animal. For there is hardly any phenomenon of irritability observed in the animal Which is not also discoverable in the plant. In the multicellular animal organism as higher complexity was attained, it was accompanied by the gradual evolution of a nervous system, by which the different organs are put in intimate connection with each other and their various activities co-ordinated for ensuring the common good of the organism. Such connecting nervous links had not been suspected in the plant, commonly regarded as distinctly lower in the scale of evolution. The researches described in the present work show that not only has a nervous system been evolved in