The Power of Movement in Plants. Charles Darwin

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СКАЧАТЬ [page 14] age to circumnutate irregularly. On the first day the greater movement (from right to left in the figure) was not in the plane of the vertical and arched hypocotyl, but at right angles to it, or in the plane of the two cotyledons, which were still in close contact. The basal leg of the arch at the time when the filament was affixed to it, was already bowed considerably backwards, or from the cotyledons; had the filament been affixed before this bowing occurred, the chief movement would have been at right angles to that shown in the figure. A filament was attached to another buried hypocotyl of the same age, and it moved in a similar general manner, but the line traced was not so complex. This hypocotyl became almost straight, and the cotyledons were dragged from beneath the ground on the evening of the second day.

      Fig. 4. Brassica oleracea: circumnutating movement of buried and arched hypocotyl, with the two legs of the arch tied together, traced on horizontal glass during 33½ hours. Movement of the bead of filament magnified about 26 times, and here reduced to one-half original scale.

      Before the above observations were made, some arched hypocotyls buried at the depth of a quarter of an inch were uncovered; and in order to prevent the two legs of the arch from beginning to separate at once, they were tied together with fine silk. This was done partly because we wished to ascertain how long the hypocotyl, in its arched condition, would continue to move, and whether the movement when not masked and disturbed by the straightening process, indicated circumnutation. Firstly a filament was fixed to the basal leg of an arched hypocotyl close above the summit of the radicle. The cotyledons were still partially enclosed within the seed-coats. The movement was traced (Fig. 4) from 9.20 A.m. on Dec. [page 15] 23rd to 6.45 A.m. on Dec. 25th. No doubt the natural movement was much disturbed by the two legs having been tied together; but we see that it was distinctly zigzag, first in one direction and then in an almost opposite one. After 3 P.m. on the 24th the arched hypocotyl sometimes remained stationary for a considerable time, and when moving, moved far slower than before. Therefore, on the morning of the 25th, the glass filament was removed from the base of the basal leg, and was fixed horizontally on the summit of the arch, which, from the legs having been tied, had grown broad and almost flat. The movement was now traced during 23 hours (Fig. 5), and we

      Fig. 5. Brassica oleracea: circumnutating movement of the crown of a buried and arched hypocotyl, with the two legs tied together, traced on a horizontal glass during 23 hours. Movement of the bead of the filament magnified about 58 times, and here reduced to one-half original scale.

      see that the course was still zigzag, which indicates a tendency to circumnutation. The base of the basal leg by this time had almost completely ceased to move.

      As soon as the cotyledons have been naturally dragged from beneath the ground, and the hypocotyl has straightened itself by growth along the inner or concave surface, there is nothing to interfere with the free movements of the parts; and the circumnutation now becomes much more regular and clearly displayed, as shown in the following cases:—A seedling was placed in front and near a north-east window with a line joining the [page 16] two cotyledons parallel to the window. It was thus left the whole day so as to accommodate itself to the light. On the following morning a filament was fixed to the midrib of the larger and taller cotyledon (which enfolds the other and smaller one, whilst still within the seed), and a mark being placed close behind, the movement of the whole plant, that is, of the hypocotyl and cotyledon, was traced greatly magnified on a vertical glass. At first the plant bent so much towards the light that it was useless to attempt to trace the movement; but at 10 A.m. heliotropism almost wholly ceased and the first dot was

      Fig. 6. Brassica oleracea: conjoint circumnutation of the hypocotyl and cotyledons during 10 hours 45 minutes. Figure here reduced to one-half original scale.

      made on the glass. The last was made at 8.45 P.m.; seventeen dots being altogether made in this interval of 10 h. 45 m. (see Fig. 6). It should be noticed that when I looked shortly after 4 P.m. the bead was pointing off the glass, but it came on again at 5.30 P.m., and the course during this interval of 1 h. 30 m. has been filled up by imagination, but cannot be far from correct. The bead moved seven times from side to side, and thus described 3½ ellipses in 10¾ h.; each being completed on an average in 3 h. 4 m.

      On the previous day another seedling had been observed under similar conditions, excepting that the plant was so [page 17] placed that a line joining the two cotyledons pointed towards the window; and the filament was attached to the smaller cotyledon on the side furthest from the window. Moreover the plant was now for the first time placed in this position. The cotyledons bowed themselves greatly towards the light from 8 to 10.50 A.m., when the first dot was made (Fig. 7). During the

      Fig. 7. Brassica oleracea: conjoint circumnutation of the hypocotyl and cotyledons, from 10.50 A.m. to 8 A.m. on the following morning. Tracing made on a vertical glass.

      next 12 hours the bead swept obliquely up and down 8 times and described 4 figures representing ellipses; so that it travelled at nearly the same rate as in the previous case. during the night it moved upwards, owing to the sleep-movement of the cotyledons, and continued to move in the same direction till 9 A.m. on the following morning; but this latter movement would not have occurred with seedlings under their natural conditions fully exposed to the light.

      By 9.25 A.m. on this second day the same cotyledon had [page 18] begun to fall, and a dot was made on a fresh glass. The movement was traced until 5.30 P.m. as shown in (Fig. 8), which is given, because the course followed was much more irregular than on the two previous occasions. During these 8 hours the bead changed its course greatly 10 times. The upward movement of the cotyledon during the afternoon and early part of the night is here plainly shown.

      Fig. 8. Brassica oleracea: conjoint circumnutation of the hypocotyl and cotyledons during 8 hours. Figure here reduced to one-third of the original scale, as traced on a vertical glass.

      As the filaments were fixed in the three last cases to one of the cotyledons, and as the hypocotyl was left free, the tracings show the movement of both organs conjoined; and we now wished to ascertain whether both circumnutated. Filaments were therefore fixed horizontally to two hypocotyls close beneath the petioles of their cotyledons. These seedlings had stood for two days in the same position before a north-east window. In the morning, up to about 11 A.m., they moved in zigzag lines towards the light; and at night they again became almost upright through apogeotropism. After about 11 A.m. they moved a little back from the light, often crossing and recrossing their former path in zigzag lines. the sky on this day varied much in brightness, and these observations merely proved that the hypocotyls were continually moving in a manner resembling circumnutation. On a previous day which was uniformly cloudy, a hypocotyl was firmly secured to a little stick, and a filament was fixed to the larger of the two cotyledons, and its movement was traced on a vertical glass. It fell greatly from 8.52 A.m., when the first dot was made, till 10.55 A.m.; it then rose greatly until 12.17 P.m. Afterwards it fell a little and made a loop, but by 2.22 P.m. it had risen a little and continued rising till 9.23 P.m., when it made another loop, and at 10.30 P.m. was again rising. These observations show that the cotyledons move [page 19] vertically up and down all day long, and as there was some slight lateral movement, they circumnutated.

      Fig. 9. Brassica oleracea: circumnutation of hypocotyl, in darkness, traced on a horizontal glass, by means of a filament with a bead fixed across its summit, between 9.15 A.m. and 8.30 A.m. on the following morning. Figure here reduced to one-half of original scale.

      The cabbage was one of the first plants, the seedlings of which were observed by us, and we did not then know how far the circumnutation of the different parts was affected by light. Young seedlings were therefore kept in complete darkness except for a minute or two during each observation, when they were illuminated by a small wax taper held almost vertically above them. During the first day the hypocotyl of one changed its course 13 times (see Fig. 9); and it deserves notice that the longer axes of the figures described often cross one another at right or nearly right angles. Another seedling was observed in the same manner, СКАЧАТЬ