The Power of Movement in Plants. Charles Darwin

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The Power of Movement in Plants - Charles  Darwin

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spot; but they did not rise at night. Nevertheless, on the following night they rose as usual.

      Cytisus fragrans (Leguminosae).—Only a few observations were made on this plant. The hypocotyl circumnutated to a considerable extent, but in a simple manner—namely, for two hours in one direction, and then much more slowly back again in a zigzag course, almost parallel to the first line, and beyond the starting-point. It moved in the same direction all night, but next morning began to return. The cotyledons continually [page 38] move both up and down and laterally; but they do not rise up at night in a conspicuous manner.

      Lupinus luteus (Leguminosae).—Seedlings of this plant were observed because the cotyledons are so thick (about .08 of an inch) that it seemed unlikely that they would move. Our observations were not very successful, as the seedlings are strongly heliotropic, and their circumnutation could not be accurately observed near a north-east window, although they had been kept during the previous day in the same position. A seedling was then placed in darkness with the hypocotyl secured to a stick; both cotyledons rose a little at first, and then fell during the rest of the day; in the evening between 5 and 6 P.m. they moved very slowly; during the night one continued to fall and the other rose, though only a little. The tracing was not much magnified, and as the lines were plainly zigzag, the cotyledons must have moved a little laterally, that is, they must have circumnutated.

      The hypocotyl is rather thick, about .12 of inch; nevertheless it circumnutated in a complex course, though to a small extent. The movement of an old seedling with two true leaves partially developed, was observed in the dark. As the movement was magnified about 100 times it is not trustworthy and is not given; but there could be no doubt that the hypocotyl moved in all directions during the day, changing its course 19 times. The extreme actual distance from side to side through which the upper part of the hypocotyl passed in the course of 14½ hours was only ⅙0 of an inch; it sometimes travelled at the rate of ⅕0 of an inch in an hour.

      Cucurbita ovifera (Cucurbitaceae).—Radicle: a seed which had

      Fig. 26. Cucurbita ovifera: course followed by a radicle in bending geotropically downwards, traced on a horizontal glass, between 11.25 A.m. and 10.25 P.m.; the direction during the night is indicated by the broken line. Movement of bead magnified 14 times.

      germinated on damp sand was fixed so that the slightly curved radicle, which was only .07 inch in length, stood almost vertically [page 39] upwards, in which position geotropism would act at first with little power. A filament was attached near to its base, and projected at about an angle of 45o above the horizon. The general course followed during the 11 hours of observation and during the following night is shown in the accompanying diagram (Fig. 26), and was plainly due to geotropism; but it was also clear that the radicle circumnutated. By the next morning the tip had curved so much downwards that the filament, instead of projecting at 45o above the horizon, was nearly horizontal. Another germinating seed was turned upside down and covered with damp sand; and a filament was fastened to the radicle so as to project at an angle of about 50o above the horizon; this radicle was .35 of an inch in length and a little curved. The course pursued was mainly governed, as in the last case, by geotropism, but the line traced during 12 hours and magnified as before was more strongly zigzag, again showing circumnutation.

      Four radicles were allowed to grow downwards over plates of smoked glass, inclined at 70o to the horizon, under the

      Fig. 27. Cucurbita ovifera: tracks left by tips of radicles in growing downwards over smoked glass-plates, inclined at 70o to the horizon.

      Fig. 28. Cucurbita ovifera: circumnutation of arched hypocotyl at a very early age, traced in darkness on a horizontal glass, from 8 A.m. to 10.20 A.m. on the following day. The movement of the bead magnified 20 times, here reduced to one-half of original scale.

      same conditions as in the cases of Aesculus, Phaseolus, and Vicia. Facsimiles are here given (Fig. 27) of two of these tracks; and a third short one was almost as plainly serpentine as that at A. It was also manifest by a greater or less amount of soot having been swept off the glasses, that the tips had [page 40] pressed alternately with greater and less force on them. There must, therefore, have been movement in at least two planes at right angles to one another. These radicles were so delicate that they rarely had the power to sweep the glasses quite clean. One of them had developed some lateral or secondary rootlets, which projected a few degrees beneath the horizon; and it is an important fact that three of them left distinctly serpentine tracks on the smoked surface, showing beyond doubt that they had circumnutated like the main or primary radicle. But the tracks were so slight that they could not be traced and copied after the smoked surface had been varnished.

      Fig. 29. Cucurbita ovifera: circumnutation of straight and vertical hypocotyl, with filament fastened transversely across its upper end, traced in darkness on a horizontal glass, from 8.30 A.m. to 8.30 P.m. The movement of the terminal bead originally magnified about 18 times, here only 4½ times.

      Hypocotyl.—A seed lying on damp sand was firmly fixed by two crossed wires and by its own growing radicle. The cotyledons were still enclosed within the seed-coats; and the short hypocotyl, between the summit of the radicle and the cotyledons, was as yet only slightly arched. A filament (.85 of inch in length) was attached at an angle of 35o above the horizon to the side of the arch adjoining the cotyledons. This part would ultimately form the upper end of the hypocotyl, after it had grown straight and vertical. Had the seed been properly planted, the hypocotyl at this stage of growth would have been deeply buried beneath the surface. The course followed by the bead of the filament is shown in Fig. 28. The chief lines of movement from left to right in the figure were parallel to the plane of the two united cotyledons and of the flattened seed; and this movement would aid in dragging them out of the seed-coats, which are held down by a special structure hereafter to be described. The movement at right angles to the above lines was due to the arched hypocotyl becoming more arched as it increased in height. The foregoing observations apply to the leg of the arch next to the cotyledons, but [page 41] the other leg adjoining the radicle likewise circumnutated at an equally early age.

      The movement of the same hypocotyl after it had become straight and vertical, but with the cotyledons only partially expanded, is shown in Fig. 29. The course pursued during 12 h. apparently represents four and a half ellipses or ovals, with the longer axis of the first at nearly right angles to that of the others. The longer axes of all were oblique to a line joining the opposite cotyledons. The actual extreme distance from side to side over which the summit of the tall hypocotyl passed in the course of 12 h. was .28 of an inch. The original figure was traced on a large scale, and from the obliquity of the line of view the outer parts of the diagram are much exaggerated.

      Cotyledons.—On two occasions the movements of the cotyledons were traced on a vertical glass, and as the ascending and descending lines did not quite coincide, very narrow ellipses were formed; they therefore circumnutated. Whilst young they rise vertically up at night, but their tips always remain reflexed; on the following morning they sink down again. With a seedling kept in complete darkness they moved in the same manner, for they sank from 8.45 A.m. to 4.30 P.m.; they then began to rise and remained close together until 10 P.m., when they were last observed. At 7 A.m. on the following morning they were as much expanded as at any hour on the previous day. The cotyledons of another young seedling, exposed to the light, were fully open for the first time on a certain day, but were found completely closed at 7 A.m. on the following morning. They soon began to expand again, and continued doing so till about 5 P.m.; they then began to rise, and by 10.30 P.m. stood vertically and were almost closed. At 7 A.m. on the third morning they were nearly vertical, and again expanded during the day; on the fourth morning they were not closed, yet they opened a little in the course of the day and rose a little on the following night. By this time a minute true leaf had become developed. Another seedling, still older, bearing a well-developed leaf, had a sharp rigid filament affixed to one of its cotyledons (85 mm. in length), which recorded its own movements on a revolving drum with smoked paper. The observations were made in the hot-house, where the plant had lived, so that

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