CHAPTER VIII.
MODIFIED CIRCUMNUTATION: MOVEMENTS EXCITED BY LIGHT.
Distinction between heliotropism and the effects of light on the periodicity of the movements of leaves—Heliotropic movements of Beta, Solanum, Zea, and Avena—Heliotropic movements towards an obscure light in Apios, Brassica, Phalaris, Tropaeolum, and Cassia—Apheliotropic movements of tendrils of Bignonia—Of flower-peduncles of Cyclamen—Burying of the pods—Heliotropism and apheliotropism modified forms of circumnutation—Steps by which one movement is converted into the other—Transversal-heliotropismus or diaheliotropism influenced by epinasty, the weight of the part and apogeotropism—Apogeotropism overcome during the middle of the day by diaheliotropism—Effects of the weight of the blades of cotyledons—So called diurnal sleep—Chlorophyll injured by intense light—Movements to avoid intense light
Sachs first clearly pointed out the important difference between the action of light in modifying the periodic movements of leaves, and in causing them to bend towards its source.[[1]] The latter, or heliotropic movements are determined by the direction of the light, whilst periodic movements are affected by changes in its intensity and not by its direction. The periodicity of the circumnutating movement often continues for some time in darkness, as we have seen in the last chapter; whilst heliotropic bending ceases very quickly when the light fails. Nevertheless, plants which have ceased through long-continued darkness to move periodically, if re-exposed to the light are still, according to Sachs, heliotropic.
[1] ‘Physiologie Veg.’ (French Translation), 1868, pp. 42, 517, etc.
Apheliotropism, or, as usually designated, negative heliotropism, implies that a plant, when unequally illuminated on the two sides, bends from the light, instead of, as in the last sub-class of cases, towards it; but apheliotropism is comparatively rare, at least in a well-marked degree. There is a third and large sub-class of cases, namely, those of “transversal-Heliotropismus” of Frank, which we will here call diaheliotropism. Parts of plants, under this influence, place themselves more or less transversely to the direction whence the light proceeds, and are thus fully illuminated. There is a fourth sub-class, as far as the final cause of the movement is concerned; for the leaves of some plants when exposed to an intense and injurious amount of light direct themselves, by rising or sinking or twisting, so as to be less intensely illuminated. Such movements have sometimes been called diurnal sleep. If thought advisable, they might be called paraheliotropic, and this term would correspond with our other terms.
It will be shown in the present chapter that all the movements included in these four sub-classes, consist of modified circumnutation. We do not pretend to say that if a part of a plant, whilst still growing, did not circumnutate—though such a supposition is most improbable—it could not bend towards the light; but, as a matter of fact, heliotropism seems always to consist of modified circumnutation. Any kind of movement in relation to light will obviously be much facilitated by each part circumnutating or bending successively in all directions, so that an already existing movement has only to be increased in some one direction, and to be lessened or stopped in the other directions, in order that it should become heliotropic, apheliotropic, etc., as the case may be. In the next chapter some observations on the sensitiveness of plants to light, their rate of bending towards it, and the accuracy with which they point towards its source, etc., will be given. Afterwards it will be shown—and this seems to us a point of much interest—that sensitiveness to light is sometimes confined to a small part of the plant; and that this part when stimulated by light, transmits an influence to distant parts, exciting them to bend.
Heliotropism.—When a plant which is strongly heliotropic (and species differ much in this respect) is exposed to a bright lateral light, it bends quickly towards it, and the course pursued by the stem is quite or nearly straight. But if the light is much dimmed, or occasionally interrupted, or admitted in only a slightly oblique direction, the course pursued is more or less zigzag; and as we have seen and shall again see, such zigzag movement results from the elongation or drawing out of the ellipses, loops, etc., which the plant would have described, if it had been illuminated from above. On several occasions we were much struck with this fact, whilst observing the circumnutation of highly sensitive seedlings, which were unintentionally illuminated rather obliquely, or only at successive intervals of time.
Fig. 168. Beta vulgaris: circumnutation of hypocotyl, deflected by the light being slightly lateral, traced on a horizontal glass from 8.30 A.M. to 5.30 P.M. Direction of the lighted taper by which it was illuminated shown by a line joining the first and penultimate dots. Figure reduced to one-third of the original scale.
For instance two young seedlings of Beta vulgaris were placed in the middle of a room with north-east windows, and were kept covered up, except during each observation which lasted for only a minute or two; but the result was that their hypocotyls bowed themselves to the side, whence some light occasionally entered, in lines which were only slightly zigzag. Although not a single ellipse was even approximately formed, we inferred from the zigzag lines—and, as it proved, correctly—that their hypocotyls were circumnutating, for on the following day these same seedlings were placed in a completely darkened room, and were observed each time by the aid of a small wax taper held almost directly above them, and their movements were traced on a horizontal glass above; and now their hypocotyls clearly circumnutated (Fig. 168, and Fig. 39, formerly given, p. 52); yet they moved a short distance towards the side where the taper was held up. If we look at these diagrams, and suppose that the taper had been held more on one side, and that the hypocotyls, still circumnutating, had bent themselves within the same time much more towards the light, long zigzag lines would obviously have been the result.
Fig. 169. Avena sativa: heliotropic movement and circumnutation of sheath-like cotyledon (1½ inch in height) traced on horizontal glass from 8 A.M. to 10.25 P.M. Oct. 16th.