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

directly opposite one. Thus, in three genera of Phaseoleae the leaflets moved vertically downwards at night, and the petioles rose in two of them, whilst in the third they sank. Species in the same genus often differ widely in the movements of their petioles. Even on the same plant of Lupinus pubescens some of the petioles rose 30o, others only 6o, and others sank 4o at night. The leaflets of Cassia Barclayana moved so little at night that they could not be said to sleep, yet the petioles of some young leaves rose as much as 34o. These several facts apparently indicate that the movements

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of the petioles are not performed for any special purpose; though a conclusion of this kind is generally rash. When the leaflets sink vertically down at night and the petioles rise, as often occurs, it is certain that the upward movement of the latter does not aid the leaflets in placing themselves in their proper position at night, for they have to move through a greater angular space than would otherwise have been necessary.

 

Notwithstanding what has just been said, it may be strongly suspected that in some cases the rising of the petioles, when considerable, does beneficially serve the plant by greatly reducing the surface exposed to radiation at night. If the reader will compare the two drawings (Fig. 155, p. 371) of Cassia pubescens, copied from photographs, he will see that the diameter of the plant at night is about one-third of what it is by day, and therefore the surface exposed to radiation is nearly nine times less. A similar conclusion may be deduced from the drawings (Fig. 149, p. 358) of a branch awake and asleep of Desmodium gyrans. So it was in a very striking manner with young plants of Bauhinia, and with Oxalis Ortegesii.

 

We are led to an analogous conclusion with respect to the movements of the secondary petioles of certain pinnate leaves. The pinnae of Mimosa pudica converge at night; and thus the imbricated and closed leaflets on each separate pinna are all brought close together into a single bundle, and mutually protect one another, with a somewhat smaller surface exposed to radiation. With Albizzia lophantha the pinnae close together in the same manner. Although the pinnae of Acacia Farnesiana do not converge much, they sink downwards. Those of Neptunia oleracea likewise [page 403]

move downwards, as well as backwards, towards the base of the leaf, whilst the main petiole rises. With Schrankia, again, the pinnae are depressed at night. Now in these three latter cases, though the pinnae do not mutually protect one another at night, yet after having sunk down they expose, as does a dependent sleeping leaf, much less surface to the zenith and to radiation than if they had remained horizontal.

 

Any one who had never observed continuously a sleeping plant, would naturally suppose that the leaves moved only in the evening when going to sleep, and in the morning when awaking; but he would be quite mistaken, for we have found no exception to the rule that leaves which sleep continue to move during the whole twenty-four hours; they move, however, more quickly when going to sleep and when awaking than at other times. That they are not stationary during the day is shown by all the diagrams given, and by the many more which were traced. It is troublesome to observe the movements of leaves in the middle of the night, but this was done in a few cases; and tracings were made during the early part of the night of the movements in the case of Oxalis, Amphicarpaea, two species of Erythrina, a Cassia, Passiflora, Euphorbia and Marsilea; and the leaves after they had gone to sleep, were found to be in constant movement. When, however, opposite leaflets come into close contact with one another or with the stem at night, they are, as we believe, mechanically prevented from moving, but this point was not sufficiently investigated.

 

When the movements of sleeping leaves are traced during twenty-four hours, the ascending and descending lines do not coincide, except occasionally and by accident for a short space; so that with many plants a [page 404]

single large ellipse is described during each twenty-four hours. Such ellipses are generally narrow and vertically directed, for the amount of lateral movement is small. That there is some lateral movement is shown by the ascending and descending lines not coinciding, and occasionally, as with Desmodium gyrans and Thalia dealbata, it was strongly marked. In the case of Melilotus the ellipses described by the terminal leaflet during the day are laterally extended, instead of vertically, as is usual; and this fact evidently stands in relation with the terminal leaflet moving laterally when it goes to sleep. With the majority of sleeping plants the leaves oscillate more than once up and down in the twenty-four hours; so that frequently two ellipses, one of moderate size, and one of very large size which includes the nocturnal movement, are described within the twenty-four hours. For instance, a leaf which stands vertically up during the night will sink in the morning, then rise considerably, again sink in the afternoon, and in the evening reascend and assume its vertical nocturnal position. It will thus describe, in the course of the twenty-four hours, two ellipses of unequal sizes. Other plants describe within the same time, three, four, or five ellipses. Occasionally the longer axes of the several ellipses extend in different directions, of which Acacia Farnesiana offered a good instance. The following cases will give an idea of the rate of movement: Oxalis acetosella completed two ellipses at the rate of 1 h.

25 m. for each; Marsilea quadrifoliata, at the rate of 2 h.; Trifolium subterraneum, one in 3 h. 30 m.; and Arachis hypogaea, in 4 h. 50 m. But the number of ellipses described within a given time depends largely on the state of the plant and on the conditions to which it is exposed. It often happens that a single ellipse may be described during one [page 405]

day, and two on the next. Erythrina corallodendron made four ellipses on the first day of observation and only a single one on the third, apparently owing to having been kept not sufficiently illuminated and perhaps not warm enough. But there seems likewise to be an innate tendency in different species of the same genus to make a different number of ellipses in the twenty-four hours: the leaflets of Trifolium repens made only one; those of T. resupinatum two, and those of T. subterraneum three in this time. Again, the leaflets of Oxalis Plumierii made a single ellipse; those of O.

bupleurifolia, two; those of O. Valdiviana, two or three; and those of O.

acetosella, at least five in the twenty-four hours.

 

The line followed by the apex of a leaf or leaflet, whilst describing one or more ellipses during the day, is often zigzag, either throughout its whole course or only during the morning or evening: Robinia offered an instance of zigzagging confined to the morning, and a similar movement in the evening is shown in the diagram (Fig. 126) given under Sida. The amount of the zigzag movement depends largely on the plant being placed under highly favourable conditions. But even under such favourable conditions, if the dots which mark the position of the apex are made at considerable intervals of time, and the dots are then joined, the course pursued will still appear comparatively simple, although the number of the ellipses will be increased; but if dots are made every two or three minutes and these are joined, the result often is that all the lines are strongly zigzag, many small loops, triangles, and other figures being also formed. This fact is shown in two parts of the diagram (Fig. 150) of the movements of Desmodium gyrans. Strephium floribundum, observed under a high temperature, [page 406]

made several little triangles at the rate of 43 m. for each. Mimosa pudica, similarly observed, described three little ellipses in 67 m.; and the apex of a leaflet crossed 1/500 of an inch in a second, or 0.12 inch in a minute. The leaflets of Averrhoa made a countless number of little oscillations when the temperature was high and the sun shining. The zigzag movement may in all cases be considered as an attempt to form small loops, which are drawn out by a prevailing movement in some one direction. The rapid gyrations of the little lateral leaflets of Desmodium belong to the same class of movements, somewhat exaggerated in rapidity and amplitude.

The jerking movements, with a small advance and still smaller retreat, apparently not exactly in the same line, of the hypocotyl of the cabbage and of the leaves of Dionaea, as seen under the microscope, all probably come under this same head. We may suspect that we here see the energy which is freed during the incessant chemical changes in progress in the tissues, converted into motion. Finally, it should be noted that leaflets and probably some leaves, whilst describing their ellipses, often rotate slightly on their axes; so that the plane of the leaf is directed first to one and then to another side. This was plainly seen to be the case with the large terminal leaflets of Desmodium, Erythrina and Amphicarpaea, and is probably common to all leaflets provided with a pulvinus.

 

With respect to the periodicity of the movements of sleeping leaves, Pfeffer* has so clearly shown that this depends on the daily alternations of light and darkness, that nothing farther need be said on this * ‘Die Periodischen Bewegungen der Blattorgane,’ 1875, p. 30, et passim.

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head. But we may recall the behaviour of Mimosa in the North, where the sun does not set, and the complete inversion of the daily movements by artificial light and darkness. It has also been shown by us, that although leaves subjected to darkness for a moderately long time continue to circumnutate, yet the periodicity of their movements is soon greatly disturbed, or quite annulled. The presence of light or its absence cannot be supposed to be the direct cause of the movements, for these are wonderfully diversified even with the leaflets of the same leaf, although all have of course been similarly exposed. The movements depend on innate causes, and are of an adaptive nature. The alternations of light and darkness merely give notice to the leaves that the period has arrived for them to move in a certain manner. We may infer from the fact of several plants (Tropaeolum, Lupinus, etc.) not sleeping unless they have been well illuminated during the day, that it is not the actual decrease of light in the evening, but the contrast between the amount at this hour and during the early part of the day, which excites the leaves to modify their ordinary mode of circumnutation.

 

As the leaves of most plants assume their proper diurnal position in the morning, although light be excluded, and as the leaves of some plants continue to move in the normal manner in darkness during at least a whole day, we may conclude that the periodicity of their movements is to a certain extent inherited.* The strength of such inheritance differs * Pfeffer denies such inheritance; he attributes (‘Die Period. Bewegungen,’

pp. 30-56) the periodicity when prolonged for a day or two in darkness, to “Nachwirkung,” or the after-effects of light and darkness. But we are unable to follow his train of reasoning. There does not seem to be any more reason for

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attributing such movements to this cause than, for instance, the inherited habit of winter and summer wheat to grow best at different seasons; for this habit is lost after a few years, like the movements of leaves in darkness after a few days. No doubt some effect