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

bright sunshine, it seems probable that they have been acquired for the special purpose of avoiding too intense an illumination. As it would have been very troublesome in all the above cases to

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have watched for a fitting opportunity and to have traced the movement of the leaves whilst they were fully exposed to the sunshine, we did not ascertain whether paraheliotropism always consisted of modified circumnutation; but this certainly was the case with the Averrhoa, and probably with the other species, as their leaves were continually circumnutating.

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CHAPTER IX.

 

SENSITIVENESS OF PLANTS TO LIGHT: ITS TRANSMITTED EFFECTS.

 

Uses of heliotropism—Insectivorous and climbing plants not heliotropic—

Same organ heliotropic at one age and not at another—Extraordinary sensitiveness of some plants to light—The effects of light do not correspond with its intensity—Effects of previous illumination—Time required for the action of light—After-effects of light—Apogeotropism acts as soon as light fails—Accuracy with which plants bend to the light—

This dependent on the illumination of one whole side of the part—Localised sensitiveness to light and its transmitted effects—Cotyledons of Phalaris, manner of bending—Results of the exclusion of light from their tips—

Effects transmitted beneath the surface of the ground—Lateral illumination of the tip determines the direction of the curvature of the base—

Cotyledons of Avena, curvature of basal part due to the illumination of upper part—Similar results with the hypocotyls of Brassica and Beta—

Radicles of Sinapis apheliotropic, due to the sensitiveness of their tips—

Concluding remarks and summary of chapter—Means by which circumnutation has been converted into heliotropism or apheliotropism.

 

NO one can look at the plants growing on a bank or on the borders of a thick wood, and doubt that the young stems and leaves place themselves so that the leaves may be well illuminated. They are thus enabled to decompose carbonic acid. But the sheath-like cotyledons of some Gramineae, for instance, those of Phalaris, are not green and contain very little starch; from which fact we may infer that they decompose little or no carbonic acid. Nevertheless, they are extremely heliotropic; and this probably serves them in another way, namely, as a guide from the buried seeds through fissures in the ground or through overlying masses of vegetation, into the light and air. This view

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is strengthened by the fact that with Phalaris and Avena the first true leaf, which is bright green and no doubt decomposes carbonic acid, exhibits hardly a trace of heliotropism. The heliotropic movements of many other seedlings probably aid them in like manner in emerging from the ground; for apogeotropism by itself would blindly guide them upwards, against any overlying obstacle.

 

Heliotropism prevails so extensively among the higher plants, that there are extremely few, of which some part, either the stem, flower-peduncle, petiole, or leaf, does not bend towards a lateral light. Drosera rotundifolia is one of the few plants the leaves of which exhibit no trace of heliotropism. Nor could we see any in Dionaea, though the plants were not so carefully observed. Sir J. Hooker exposed the pitchers of Sarracenia for some time to a lateral light, but they did not bend towards it.* We can understand the reason why these insectivorous plants should not be heliotropic, as they do not live chiefly by decomposing carbonic acid; and it is much more important to them that their leaves should occupy the best position for capturing insects, than that they should be fully exposed to the light.

 

Tendrils, which consist of leaves or of other organs modified, and the stems of twining plants, are, as Mohl long ago remarked, rarely heliotropic; and here again we can see the reason why, for if they had moved towards a lateral light they would have been drawn away from their supports. But some tendrils are apheliotropic, for instance those of Bignonia capreolata

 

* According to F. Kurtz (‘Verhandl. des Bot. Vereins der Provinz Brandenburg,’ Bd. xx. 1878) the leaves or pitchers of Darlingtonia Californica are strongly apheliotropic. We failed to detect this movement in a plant which we possessed for a short time.

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and of Smilax aspera; and the stems of some plants which climb by rootlets, as those of the Ivy and Tecoma radicans, are likewise apheliotropic, and they thus find a support. The leaves, on the other hand, of most climbing plants are heliotropic; but we could detect no signs of any such movement in those of Mutisia clematis.

 

As heliotropism is so widely prevalent, and as twining plants are distributed throughout the whole vascular series, the apparent absence of any tendency in their stems to bend towards the light, seemed to us so remarkable a fact as to deserve further investigation, for it implies that heliotropism can be readily eliminated. When twining plants are exposed to a lateral light, their stems go on revolving or circumnutating about the same spot, without any evident deflection towards the light; but we thought that we might detect some trace of heliotropism by comparing the average rate at which the stems moved to and from the light during their successive revolutions.* Three young plants (about a foot in height) of Ipomoea caerulea and four of I. purpurea, growing in separate pots, were placed on a bright day before a north-east window in a room otherwise darkened, with the tips of their revolving stems fronting the window. When the tip of each plant pointed directly from the window, and when again towards it, the times were recorded. This was continued from 6.45 A.M. till a little after 2 P.M. on June 17th. After a few observations we concluded that we could safely estimate the time

 

* Some erroneous statements are unfortunately given on this subject, in ‘The Movements and Habits of Climbing Plants,’ 1875, pp. 28, 32, 40, and 53. Conclusions were drawn from an insufficient number of observations, for we did not then know at how unequal a rate the stems and tendrils of climbing plants sometimes travel in different parts of the same revolution.

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taken by each semicircle, within a limit of error of at most 5 minutes.

Although the rate of movement in different parts of the same revolution varied greatly, yet 22 semicircles to the light were completed, each on an average in 73.95 minutes; and 22 semicircles from the light each in 73.5

minutes. It may, therefore, be said that they travelled to and from the light at exactly the same average rate; though probably the accuracy of the result was in part accidental. In the evening the stems were not in the least deflected towards the window. Nevertheless, there appears to exist a vestige of heliotropism, for with 6 out of the 7 plants, the first semicircle from the light, described in the early morning after they had been subjected to darkness during the night and thus probably rendered more sensitive, required rather more time, and the first semicircle to the light considerably less time, than the average. Thus with all 7 plants, taken together, the mean time of the first semicircle in the morning from the light, was 76.8 minutes, instead of 73.5 minutes, which is the mean of all the semicircles during the day from the light; and the mean of the first semicircle to the light was only 63.1, instead of 73.95 minutes, which was the mean of all the semicircles during the day to the light.

 

Similar observations were made on Wistaria Sinensis, and the mean of 9

semicircles from the light was 117 minutes, and of 7 semicircles to the light 122 minutes, and this difference does not exceed the probable limit of error. During the three days of exposure, the shoot did not become at all bent towards the window before which it stood. In this case the first semicircle from the light in the early morning of each day, required rather less time for its performance than did the first semicircle to the light; and this result,

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if not accidental, appears to indicate that the shoots retain a trace of an original apheliotropic tendency. With Lonicera brachypoda the semicircles from and to the light differed considerably in time; for 5 semicircles from the light required on a mean 202.4 minutes, and 4 to the light, 229.5

minutes; but the shoot moved very irregularly, and under these circumstances the observations were much too few.

 

It is remarkable that the same part on the same plant may be affected by light in a widely different manner at different ages, and as it appears at different seasons. The hypocotyledonous stems of Ipomoea caerulea and purpurea are extremely heliotropic, whilst the stems of older plants, only about a foot in height, are, as we have just seen, almost wholly insensible to light. Sachs states (and we have observed the same fact) that the hypocotyls of the Ivy (Hedera helix) are slightly heliotropic; whereas the stems of plants grown to a few inches in height become so strongly apheliotropic, that they bend at right angles away from the light.

Nevertheless, some young plants which had behaved in this manner early in the summer again became distinctly heliotropic in the beginning of September; and the zigzag courses of their stems, as they slowly curved towards a north-east window, were traced during 10 days. The stems of very young plants of Tropaeolum majus are highly heliotropic, whilst those of older plants, according to Sachs, are slightly apheliotropic. In all these cases the heliotropism of the very young stems serves to expose the cotyledons, or when the cotyledons are hypogean the first true leaves, fully to the light; and the loss of this power by the older stems, or their becoming apheliotropic, is connected with their habit of climbing.

 

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it is no doubt a great advantage to them in their struggle for life to expose their cotyledons to the light as quickly and as fully as possible, for the sake of obtaining carbon. It has been shown in the first chapter that the greater number of seedlings circumnutate largely and rapidly; and as heliotropism consists of modified circumnutation, we are tempted to look at the high development of these two powers in seedlings as intimately connected. Whether there are any plants which circumnutate slowly and to a small extent, and yet are highly heliotropic, we do not know; but there are several, and there is nothing surprising in this fact, which circumnutate largely and are not at all, or only slightly, heliotropic. Of such cases Drosera rotundifolia offers an excellent instance. The stolons of the strawberry circumnutate almost like the stems of climbing plants, and they are not at all affected by a moderate light; but when exposed late in the summer to a somewhat brighter light they were slightly heliotropic; in sunlight, according to De Vries, they are apheliotropic. Climbing plants circumnutate much more widely than any other plants, yet they are not at all heliotropic.

 

Although the stems of most seedling plants are strongly heliotropic, some few are but slightly heliotropic, without our being able to assign any reason. This is the case with the hypocotyl of Cassia tora, and we were struck with the same fact with some other seedlings, for instance, those of Reseda odorata. With respect to the degree of sensitiveness of the more sensitive kinds, it was shown in the last chapter that seedlings of several species, placed before a north-east window protected by several blinds, and exposed in the rear to the diffused light of the room, moved with unerring certainty towards the window, although

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it was impossible to judge, excepting by the shadow cast by an upright pencil on a white card, on which side most light entered, so that the excess on one side must have been extremely small.

 

A pot with seedlings of Phalaris Canariensis,