The Power of Movement in Plants - Charles Darwin (top reads txt) 📗

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After having seen how the epicotyls moved towards a moderately bright light, seedlings were placed at 7.48 A.M. (Sept. 7th) before a north-east window, covered by a towel, and shortly afterwards by an ordinary linen blind, but the epicotyls still moved towards the window. At 9.13 A.M. two additional muslin blinds were suspended, so that the seedlings received very little more light from the window than from the interior of the room.

The sky varied in brightness, and the seedlings occasionally Fig. 175. Tropaeolum majus: heliotropic movement and circumnutation of the epicotyl of a young seedling towards a dull lateral light, traced on a horizontal glass from 7.48 A.M. to 10.40 P.M. Figure reduced to one-half of the original scale.

received for a short time less light from the window than from the opposite side (as ascertained by the shadow cast), and then one of the blinds was temporarily removed. In the evening the blinds were taken away, one by one.

the course pursued by an epicotyl under these circumstances is shown in Fig. 175. During the whole day, until 6.45 P.M., it plainly bowed itself towards the light; and the tip moved over a considerable space. After 6.45

P.M. it moved backwards, or from the window, till [page 430]

10.40 P.M., when the last dot was made. Here, then, we have a distinct heliotropic movement, effected by means of six elongated figures (which if dots had been made every few minutes would have been more or less elliptic) directed towards the light, with the apex of each successive ellipse nearer to the window than the previous one. Now, if the light had been only a little brighter, the epicotyl would have bowed itself more to the light, as we may safely conclude from the previous trials; there would also have been less lateral movement, and the ellipses or other figures would have been drawn out into a strongly marked zigzag line, with probably one or two small loops still formed. If the light had been much brighter, we should have had a slightly zigzag line, or one quite straight, for there would have been more movement in the direction of the light, and much less from side to side.

Fig. 176. Tropaeolum majus: heliotropic movement and circumnutation of an old internode towards a lateral light, traced on a horizontal glass from 8

A.M. Nov. 2nd to 10.20 A.M. Nov. 4th. Broken lines show the nocturnal course.

Sachs states that the older internodes of this Tropaeolum are apheliotropic; we therefore placed a plant, 11 3/4 inches high, in a box, blackened within, but open on one side in front of a north-east window without any blind. A filament was fixed to the third internode from the summit on one plant, and to the fourth internode of another. These internodes were either not old enough, or the light was not sufficiently bright, to induce apheliotropism, for both plants bent slowly towards, instead of from the window during four days. The course, during two days of the first-mentioned internode, is given in Fig. 176; and we see that it either circumnutated on a small scale, or travelled in a zigzag line towards the light. We have thought this case of feeble heliotropism in one of the older internodes of a plant,

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which, whilst young, is so extremely sensitive to light, worth giving.

Fig. 177. Cassia tora: heliotropic movement and circumnutation of a hypocotyl (1 � inch in height) traced on a horizontal glass from 8 A.M. to 10.10 P.M. Oct. 7th. Also its circumnutation in darkness from 7 A.M. Oct.

8th to 7.45 A.M. Oct. 9th.

Cassia tora.—The cotyledons of this plant are extremely sensitive to light, whilst the hypocotyls are much less sensitive than those of most other seedlings, as we had often observed with surprise. It seemed therefore worth while to trace their movements. They were exposed to a lateral light before a north-east window, which was at first covered merely by a muslin blind, but as the sky grew brighter about 11 A.M., an additional linen blind was suspended. After 4 P.M. one blind and then the other was removed. The seedlings were protected on each side and above, but were open to the diffused light of the room in the rear. Upright filaments were fixed to the hypocotyls of two seedlings, which stood vertically in the morning. The accompanying figure (Fig. 177) shows the course pursued by one of them during two days; but it should be particularly noticed that during the second day the seedlings were kept in darkness, and they then circumnutated round nearly the same small space. On the first day (Oct.

7th) the hypocotyl moved from 8 A.M. to 12.23 P.M., toward the light in a zigzag line, then turned abruptly to the left and afterwards described a small ellipse. Another irregular

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ellipse was completed between 3 P.M. and about 5.30 P.M., the hypocotyl still bending towards the light. The hypocotyl was straight and upright in the morning, but by 6 P.M. its upper half was bowed towards the light, so that the chord of the arc thus formed stood at an angle of 20o with the perpendicular. After 6 P.M. its course was reversed through the action of apogeotropism, and it continued to bend from the window during the night, as shown by the broken line. On the next day it was kept in the dark (excepting when each observation was made by the aid of a taper), and the course followed from 7 A.M. on the 8th to 7.45 A.M. on the 9th is here likewise shown. The difference between the two parts of the figure (177), namely that described during the daytime on the 7th, when exposed to a rather dim lateral light, and that on the 8th in darkness, is striking. The difference consists in the lines during the first day having been drawn out in the direction of the light. The movements of the other seedling, traced under the same circumstances, were closely similar.

Apheliotropism.—We succeeded in observing only two cases of apheliotropism, for these are somewhat rare; and the movements are generally so slow that they would have been very troublesome to trace.

Fig. 178. Bignonia capreolata: apheliotropic movement of a tendril, traced on a horizontal glass from 6.45 A.M. July 19th to 10 A.M. 20th. Movements as originally traced, little magnified, here reduced to two-thirds of the original scale.

Bignonia capreolata.—No organ of any plant, as far as we have seen, bends away so quickly from the light as do the tendrils of this Bignonia. They are also remarkable from circumnutating much less regularly than most other tendrils, often remaining stationary; they depend on apheliotropism for coming into

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contact with the trunks of trees.* The stem of a young plant was tied to a stick at the base of a pair of fine tendrils, which projected almost vertically upwards; and it was placed in front of a north-east window, being protected on all other sides from the light. The first dot was made at 6.45 A.M., and by 7.35 A.M. both tendrils felt the full influence of the light, for they moved straight away from it until 9.20 A.M., when they circumnutated for a time, still moving, but only a little, from the light (see Fig. 178 of the left-hand tendril). After 3 P.M. they again moved rapidly away from the light in zigzag lines. By a late hour in the evening both had moved so far, that they pointed in a direct line from the light.

During the night they returned a little in a nearly opposite direction. On the following morning they again moved from the light and converged, so that by the evening they had become interlocked, still pointing from the light. The right-hand tendril, whilst converging, zigzagged much more than the one figured. Both tracings showed that the apheliotropic movement was a modified form of circumnutation.

Cyclamen Persicum.—Whilst this plant is in flower the peduncles stand upright, but their uppermost part is hooked so that the flower itself hangs downwards. As soon as the pods begin to swell, the peduncles increase much in length and slowly curve downwards, but the short, upper, hooked part straightens itself. Ultimately the pods reach the ground, and if this is covered with moss or dead leaves, they bury themselves. We have often seen saucer-like depressions formed by the pods in damp sand or sawdust; and one pod (.3 of inch in diameter) buried itself in sawdust for three-quarters of its length.** We shall have occasion hereafter to consider the object gained by this burying process. The peduncles can change the direction of their curvature, for if a pot, with plants having their peduncles already bowed downwards, be placed horizontally, they slowly bend at right angles to their former direction towards the centre of the earth. We therefore at first attributed the movement to geotropism; but a pot which had lain horizontally with the pods

* ‘The Movements and Habits of Climbing Plants,’ 1875, p. 97.

** The peduncles of several other species of Cyclamen twist themselves into a spire, and according to Erasmus Darwin (‘Botanic Garden,’ Canto., iii. p.

126), the pods forcibly penetrate the earth. See also Grenier and Godron, ‘Flore de France,’ tom. ii. p. 459.

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all pointing to the ground, was reversed, being still kept horizontal, so that the pods now pointed directly upwards; it was then placed in a dark cupboard, but the pods still pointed upwards after four days and nights.

The pot, in the same position, was next brought back into the light, and after two days there was some bending downwards of the peduncles, and on the fourth day two of them pointed to the centre of the earth, as did the others after an additional day or two. Another plant, in a pot which had always stood upright, was left in the dark cupboard for six days; it bore 3

peduncles, and only one became within this Fig. 179. Cyclamen Persicum: downward apheliotropic movement of a flower-peduncle, greatly magnified (about 47 times?), traced on a horizontal glass from 1 P.M. Feb. 18th to 8 A.M. 21st.

time at all bowed downwards, and that doubtfully. The weight, therefore, of the pods is not the cause of the bending down. This pot was then brought back into the light, and after three days the peduncles were considerably bowed downwards. We are thus led to infer that the downward curvature is due to apheliotropism; though more trials ought to have been made.

In order to observe the nature of this movement, a peduncle bearing a large pod which had reached and rested on the ground, was lifted a little up and secured to a stick. A filament was fixed across the pod with a mark beneath, and its move-

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ment, greatly magnified, was traced on a horizontal glass during 67 h. The plant was illuminated during the day from above. A copy of the tracing is given on p. 434 (Fig. 179); and there can be no doubt that the descending movement is one of modified circumnutation, but on an extremely small scale. The observation was repeated on another pod, which had partially buried itself in sawdust, and which was lifted up a quarter of an inch above the surface; it described three very small circles in 24 h.

Considering the great length and thinness of the peduncles and the lightness of the pods, we may conclude that they would not be able to excavate saucer-like depressions in sand or sawdust, or bury themselves in moss, etc., unless they were aided by their continued rocking or circumnutating movement.]

Relation between Circumnutation and Heliotropism.—Any one who will look at the foregoing diagrams, showing the movements of the stems of