The Power of Movement in Plants, Charles Darwin [the reading list book TXT] 📗

they bent all day in a strongly zigzag course. On the following day they continued to bend in the same direction (Fig. 169), but zigzagged much less. The sky, however, became between 12.40 and 2.35 P.M.

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overcast with extraordinarily dark thunder-clouds, and it was interesting to note how plainly the cotyledons circumnutated during this interval.

 

The foregoing observations are of some value, from having been made when we were not attending to heliotropism; and they led us to experiment on several kinds of seedlings, by exposing them to a dim lateral light, so as to observe the gradations between ordinary circumnutation and heliotropism.

Seedlings in pots were placed in front of, and about a yard from, a north-east window; on each side and over the pots black boards were placed; in the rear the pots were open to the diffused light of the room, which had a second north-east and a north-west window. By hanging up one or more blinds before the window where the seedlings stood, it was easy to dim the light, so that very little more entered on this side than on the opposite one, which received the diffused light of the room. Late in the evening the blinds were successively removed, and as the plants had been subjected during the day to a very obscure light, they continued to bend towards the window later in the evening than would otherwise have occurred. Most of the seedlings were selected because they were known to be highly sensitive to light, and some because they were but little sensitive, or had become so from having grown old. The movements were traced in the usual manner on a horizontal glass cover; a fine glass filament with little triangles of paper having been cemented in an upright position to the hypocotyls.

Whenever the stem or hypocotyl became much bowed towards the light, the latter part of its course had to be traced on a vertical glass, parallel to the window, and at right angles to the horizontal glass cover.

Fig. 170. Apios graveolens: heliotropic movement of hypocotyl (.45 of inch in height) towards a moderately bright lateral light, traced on a horizontal glass from 8.30 A.M. to 11.30 A.M. Sept. 18th. Figure reduced to one-third of original scale.

 

Apios graveolens.—The hypocotyl bends in a few hours rectan-

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gularly towards a bright lateral light. In order to ascertain how straight a course it would pursue when fairly well illuminated on one side, seedlings were first placed before a south-west window on a cloudy and rainy morning; and the movement of two hypocotyls were traced for 3 h., during which time they became greatly bowed towards the light. One of these tracings is given on p. 422 (Fig. 170), and the course may be seen to be almost straight. But the amount of light on this occasion was superfluous, for two seedlings were placed before a north-east window, protected by an ordinary linen and two muslin blinds, yet their hypocotyls moved towards this rather dim light in only slightly zigzag lines; but after 4 P.M., as the light waned, the lines became distinctly zigzag. One of these seedlings, moreover, described in the afternoon an ellipse of considerable size, with its longer axis directed towards the window.

 

We now determined that the light should be made dim enough, so we began by exposing several seedlings before a north-east window, protected by one linen blind, three muslin blinds, and a towel. But so little light entered that a pencil cast no perceptible shadow on a white card, and the hypocotyls did not bend at all towards the window. During this time, from 8.15 to 10.50 A.M., the hypocotyls zigzagged or circumnutated near the same spot, as may be seen at A, in Fig. 171. The towel, therefore, was removed at 10.50 A.M., and replaced by two muslin blinds, and now the light passed through one ordinary linen and four muslin blinds. When a pencil was held upright on a card close to the seedlings, it cast a shadow (pointing from the window) which could only just be distinguished. Yet this very slight excess of light on one side sufficed to cause the hypocotyls of all the seedlings immediately to begin bending in zigzag lines towards the window.

The course of one is shown at A (Fig. 171): after moving towards the window from 10.50 A.M. to 12.48 P.M. it bent from the window, and then returned in a nearly parallel line; that is, it almost completed between 12.48 and 2

P.M. a narrow ellipse. Late in the evening, as the light waned, the hypocotyl ceased to bend towards the window, and circumnutated on a small scale round the same spot; during the night it moved considerably backwards, that is, became more upright, through the action of apogeotropism. At B, we have a tracing of the movements of another seedling from the hour (10.50 A.M.) when the towel was removed; and it is in all essential respects

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similar to the previous one. In these two cases there could be no doubt that the ordinary circumnutating movement of the hypocotyl was modified and rendered heliotropic.

 

Fig. 171. Apios graveolens: heliotropic movement and circumnutation of the hypocotyls of two seedlings towards a dim lateral light, traced on a horizontal glass during the day. The broken lines show their return nocturnal courses. Height of hypocotyl of A .5, and of B .55 inch. Figure reduced to one-half of original scale.

 

Brassica oleracea.—The hypocotyl of the cabbage, when not disturbed by a lateral light, circumnutates in a complicated [page 425]

manner over nearly the same space, and a figure formerly given is here reproduced (Fig. 172). If the hypocotyl is exposed to a moderately strong lateral light it moves quickly towards this side, travelling in a straight, or nearly straight, line. But when the lateral light is very dim its course is extremely tortuous, and evidently consists of modified circumnutation.

Seedlings were placed before a north-east window, protected by a linen and muslin blind and by a towel. The sky was cloudy, and whenever the clouds grew a little lighter an additional muslin blind was temporarily suspended.

The light from the window was

 

Fig. 172. Brassica oleracea: ordinary circumnutating movement of the hypocotyl of a seedling plant.

 

thus so much obscured that, judging by the unassisted eye, the seedlings appeared to receive more light from the interior of the room than from the window; but this was not really the case, as was shown by a very faint shadow cast by a pencil on a card. Nevertheless, this extremely small excess of light on one side caused the hypocotyls, which in the morning had stood upright, to bend at right angles towards the window, so that in the evening (after 4.23 P.M.) their course had to be traced on a vertical glass parallel to the window. It should be stated that at 3.30 P.M., by which time the sky had become darker, the towel was removed and replaced by an additional muslin blind, which itself was removed at 4 P.M., the other two [page 426]

blinds being left suspended. In Fig. 173 the course pursued, between 8.9

A.M. and 7.10 P.M., by one of the hypocotyls thus Fig. 173. Brassica oleracea: heliotropic movement and circumnutation of a hypocotyl towards a very dim lateral light, traced during 11 hours, on a horizontal glass in the morning, and on a vertical glass in the evening.

Figure reduced to one-third of the original scale.

 

exposed is shown. It may be observed that during the first 16 m. the hypocotyl moved obliquely from the light, and this, [page 427]

no doubt, was due to its then circumnutating in this direction. Similar cases were repeatedly observed, and a dim light rarely or never produced any effect until from a quarter to three-quarters of an hour had elapsed.

After 5.15 P.M., by which time the light had become obscure, the hypocotyl began to circumnutate about the same spot. The contrast between the two figures (172 and 173) would have been more striking, if they had been originally drawn on the same scale, and had been equally reduced. But the movements shown in Fig. 172 were at first more magnified, and have been reduced to only one-half of the original scale; whereas those in Fig. 173

were at first less magnified, and have been reduced to a one-third scale. A tracing made at the same time with the last of the movements of a second hypocotyl, presented a closely analogous appearance; but it did not bend quite so much towards the light, and it circumnutated rather more plainly.

 

Fig. 174. Phalaris Canariensis: heliotropic movement and circumnutation of a rather old cotyledon, towards a dull lateral light, traced on a horizontal glass from 8.15 A.M. Sept. 16th to 7.45 A.M. 17th. Figure reduced to one-third of original scale.

 

Phalaris Canariensis.—The sheath-like cotyledons of this monocotyledonous plant were selected for trial, because they are very sensitive to light and circumnutate well, as formerly shown (see Fig. 49, p. 63). Although we felt no doubt about the result, some seedlings were first placed before a south-west window on a moderately bright morning, and the movements of one were traced. As is so common, it moved

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for the first 45 m. in a zigzag line; it then felt the full influence of the light, and travelled towards it for the next 2 h. 30 m. in an almost straight line. The tracing has not been given, as it was almost identical with that of Apios under similar circumstances (Fig. 170). By noon it had bowed itself to its full extent; it then circumnutated about the same spot and described two ellipses; by 5 P.M. it had retreated considerably from the light, through the action of apogeotropism. After some preliminary trials for ascertaining the right degree of obscurity, some seedlings were placed (Sept. 16th) before a north-east window, and light was admitted through an ordinary linen and three muslin blinds. A pencil held close by the pot now cast a very faint shadow on a white card, pointing from the window. In the evening, at 4.30 and again at 6 P.M., some of the blinds were removed. In Fig. 174 we see the course pursued under these circumstances by a rather old and not very sensitive cotyledon, 1.9 inch in height, which became much bowed, but was never rectangularly bent towards the light. From 11 A.M., when the sky became rather duller, until 6.30

P.M., the zigzagging was conspicuous, and evidently consisted of drawn-out ellipses. After 6.30 P.M. and during the night, it retreated in a crooked line from the window. Another and younger seedling moved during the same time much more quickly and to a much greater distance, in an only slightly zigzag line towards the light; by 11 A.M. it was bent almost rectangularly in this direction, and now circumnutated about the same place.

 

Tropaeolum majus.—Some very young seedlings, bearing only two leaves, and therefore not as yet arrived at the climbing stage of growth, were first tried before a north-east window without any blind. The epicotyls bowed themselves towards the light so rapidly that in little more than 3 h. their tips pointed rectangularly towards it. The lines traced were either nearly straight or slightly zigzag; and in this latter case we see that a trace of circumnutation was retained even under the influence of a moderately bright light. Twice whilst these epicotyls were bending towards the window, dots were made every 5 or 6 minutes, in order to detect any trace of lateral movement, but there was hardly any; and the lines formed by their junction were nearly straight, or only very slightly zigzag, as in the other parts of the figures. After the epicotyls had bowed themselves to the full extent towards the light, ellipses of considerable size