The Power of Movement in Plants

by Charles Darwin; Sir Francis Darwin

Vicia faba.—Radicles of this plant were extended horizontally either over water or with their lower surfaces just touching it. Their tips had previously been cut off, in a direction as accurately transverse as could be done, to different lengths, measured from the apex of the root-cap, and which will be specified in each case. Light was always excluded. We had previously tried hundreds of unmutilated radicles under similar circumstances, and found that every one that was healthy became plainly geotropic in under 12 h. In the case of four radicles which had their tips cut off for a length of 1.5 mm., new root caps and new vegetative points were re-formed after an interval of 3 days 20 h.; and these when placed horizontally were acted on by geotropism. On some other occasions this regeneration of the tips and reacquired sensitiveness occurred within a somewhat shorter time. Therefore, radicles having their tips amputated should be observed in from 12 to 48 h. after the operation.

Four radicles were extended horizontally with their lower surfaces touching the water, and with their tips cut off for a length of only 0.5 mm.: after 23 h. three of them were still horizontal; after 47 h. one of the three became fairly geotropic; and after 70 h. the other two showed a trace of this action. The fourth radicle was vertically geotropic after 23 h.; but by an accident the root-cap alone and not the vegetative point was found to have been amputated; so that this case formed no real exception and might have been excluded.

Five radicles were extended horizontally like the last, and had their tips cut off for a length of 1 mm.; after 22–23 h., four of them were still horizontal, and one was slightly geotropic; after 48 h. the latter had become vertical; a second was also somewhat geotropic; two remained approximately horizontal; and the last or fifth had grown in a disordered manner, for it was inclined upwards at an angle of 65° above the horizon.

Fourteen radicles were extended horizontally at a little height over the water with their tips cut off for a length of 1.5 mm.; after 12 h. all were horizontal, whilst five control or standard specimens in the same jar were all bent greatly downwards. After 24 h. several of the amputated radicles remained horizontal, but some showed a trace of geotropism, and one was plainly geotropic, for it was inclined at 40° beneath the horizon.

Seven horizontally extended radicles from which the tips had been cut off for the unusual length of 2 mm. unfortunately were not looked at until 35 h. had elapsed; three were still horizontal, but to our surprise, four were more or less plainly geotropic.

The radicles in the foregoing cases were measured before their tips were amputated, and in the course of 24 h. they had all increased greatly in length; but the measurements are not worth giving. It is of more importance that Sachs found that the rate of growth of the different parts of radicles with amputated tips was the same as with unmutilated ones. Altogether twenty-nine radicles were operated on in the manner above described, and of these only a few showed any geotropic curvature within 24 h.; whereas radicles with unmutilated tips always became, as already stated, much bent down in less than half of this time. The part of the radicle which bends most lies at the distance of from 3 to 6 mm. from the tip, and as the bending part continues to grow after the operation, there does not seem any reason why it should not have been acted on by geotropism, unless its curvature depended on some influence transmitted from the tip. And we have clear evidence of such transmission in Ciesielski’s experiments, which we repeated and extended in the following manner.

Beans were embedded in friable peat with the hilum downwards, and after their radicles had grown perpendicularly down for a length of from ½ to 1 inch, sixteen were selected which were perfectly straight, and these were placed horizontally on the peat, being covered by a thin layer of it. They were thus left for an average period of 1 h. 37 m. The tips were then cut off transversely for a length of 1.5 mm., and immediately afterwards they were embedded vertically in the peat. In this position geotropism would not tend to induce any curvature, but if some influence had already been transmitted from the tip to the part which bends most, we might expect that this part would become curved in the direction in which geotropism had previously acted; for it should be noted that these radicles being now destitute of their sensitive tips, would not be prevented by geotropism from curving in any direction. The result was that of the sixteen vertically embedded radicles, four continued for several days to grow straight downwards, whilst twelve became more or less bowed laterally. In two of the twelve, a trace of curvature was perceptible in 3 h. 30 m., counting from the time when they had first been laid horizontally; and all twelve were plainly bowed in 6 h., and still more plainly in 9 h. In every one of them the curvature was directed towards the side which had been downwards whilst the radicles remained horizontal. The curvature extended for a length of from 5 to, in one instance, 8 mm., measured from the cut-off end. Of the twelve bowed radicles five became permanently bent into a right angle; the other seven were at first much less bent, and their curvature generally decreased after 24 h., but did not wholly disappear. This decrease of curvature would naturally follow, if an exposure of only 1 h. 37 m. to geotropism, served to modify the turgescence of the cells, but not their subsequent growth to the full extent. The five radicles which were rectangularly bent became fixed in this position, and they continued to grow out horizontally in the peat for a length of about 1 inch during from 4 to 6 days. By this time new tips had been formed; and it should be remarked that this regeneration occurred slower in the peat than in water, owing perhaps to the radicles being often looked at and thus disturbed. After the tips had been regenerated, geotropism was able to act on them, so that they now became bowed vertically downwards. An accurate drawing (Fig. 195) is given on the opposite page of one of these five radicles, reduced to half the natural size.

We next tried whether a shorter exposure to geotropism would suffice to produce an after-effect. Seven radicles were extended horizontally for an hour, instead of 1 h. 37 m. as in the former trial; and after their tips (1.5 mm. in length) had been amputated, they were placed vertically in damp peat. Of these, three were not in the least affected and continued for days to grow straight downwards. Four showed after 8 h. 30 m. a mere trace of curvature in the direction in which they had been acted on by geotropism; and in this respect they differed much from those which had been exposed for 1 h. 37 m., for many of the latter were plainly curved in 6 h. The curvature of one of these four radicles almost disappeared after 24 h. In the second, the curvature increased during two days and then decreased. the third radicle became permanently bent, so that its terminal part made an angle of about 45° with its original vertical direction. The fourth radicle became horizontal. These two, latter radicles continued during two more days to grow in the peat in the same directions, that is, at an angle of 45° beneath the horizon and horizontally. By the fourth morning new tips had been re-formed, and now geotropism was able to act on them again, and they became bent perpendicularly downwards, exactly as in the case of the five radicles described in the last paragraph and as is shown in (Fig. 195) here given.

Fig. 195. Vicia faba: radicle, rectangularly bent at A, after the amputation of the tip, due to the previous influence of geotropism. L, side of bean which lay on the peat, whilst geotropism acted on the radicle. A, point of chief curvature of the radicle, whilst standing vertically downwards. B, point of chief curvature after the regeneration of the tip, when geotropism again acted. C, regenerated tip.

Lastly, five other radicles were similarly treated, but were exposed to geotropism during only 45 m. After 8 h. 30 m. only one was doubtfully affected; after 24 h. two were just perceptibly curved towards the side which had been acted on by geotropism; after 48 h. the one first mentioned had a radius of curvature of 60 mm. That this curvature was due to the action of geotropism during the horizontal position of the radicle, was shown after 4 days, when a new tip had been re-formed, for it then grew perpendicularly downwards. We learn from this case that when the tips are amputated after an exposure to geotropism of only 45 m., though a slight influence is sometimes transmitted to the adjoining part of the radicle, yet this seldom suffices, and then only slowly, to induce even moderately well-pronounced curvature.