The Power of Movement in Plants by Charles Darwin (book club books TXT) 📖

the free seedlings; and the remaining ten were slightly curved, although the basal part of several of them, which normally becomes more curved than any other part, exhibited hardly a trace of curvature. These 12 seedlings taken together differed greatly in their degree of curvature from all the many other seedlings in the same pots.

 

Better evidence of the efficiency of the blackened tubes was incidentally afforded by some experiments hereafter to be given, [page 473]

in which the upper halves of 14 cotyledons were enclosed in tubes from which an extremely narrow stripe of the black varnish had been scraped off.

These cleared stripes were not directed towards the window, but obliquely to one side of the room, so that only a very little light could act on the upper halves of the cotyledons. These 14 seedlings remained during eight hours of exposure before a south-west window on a hazy day quite upright; whereas all the other many free seedlings in the same pots became greatly bowed towards the light.

 

We will now turn to the trials with caps made of very thin tin-foil. These were placed at different times on the summits of 24 cotyledons, and they extended down for a length of between .15 and .2 of an inch. The seedlings were exposed to a lateral light for periods varying between 6 h. 30 m. and 7 h. 45 m., which sufficed to cause all the other seedlings in the same pots to become almost rectangularly bent towards the light. They varied in height from only .04 to 1.15 inch, but the greater number were about .75

inch. Of the 24 cotyledons with their summits thus protected, 3 became much bent, but not in the direction of the light, and as they did not straighten themselves through apogeotropism during the following night, either the caps were too heavy or the plants themselves were in a weak condition; and these three cases may be excluded. There are left for consideration 21

cotyledons; of these 17 remained all the time quite upright; the other 4

became slightly inclined to the light, but not in a degree comparable with that of the many free seedlings in the same pots. As the glass-tubes, when unpainted, did not prevent the cotyledons from becoming greatly bowed, it cannot be supposed that the caps of very thin tin-foil did so, except through the exclusion of the light. To prove that the plants had not been injured, the caps were removed from 6 of the upright seedlings, and these were exposed before a paraffin lamp for the same length of time as before, and they now all became greatly curved towards the light.

 

As caps between .15 and .2 of an inch in depth were thus proved to be highly efficient in preventing the cotyledons from bending towards the light, 8 other cotyledons were protected with caps between only .06 and .12

in depth. Of these, two remained vertical, one was considerably and five slightly curved towards the light, but far less so than the free seedlings in the same pots.

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Another trial was made in a different manner, namely, by bandaging with strips of tin-foil, about .2 in breadth, the upper part, but not the actual summit, of eight moderately young seedlings a little over half an inch in height. The summits and the basal parts were thus left fully exposed to a lateral light during 8 h.; an upper intermediate zone being protected. With four of these seedlings the summits were exposed for a length of .05 inch, and in two of them this part became curved towards the light, but the whole lower part remained quite upright; whereas the entire length of the other two seedlings became slightly curved towards the light. The summits of the four other seedlings were exposed for a length of .04 inch, and of these one remained almost upright, whilst the other three became considerably curved towards the light. The many free seedlings in the same pots were all greatly curved towards the light.

 

From these several sets of experiments, including those with the glass-tubes, and those when the tips were cut off, we may infer that the exclusion of light from the upper part of the cotyledons of Phalaris prevents the lower part, though fully exposed to a lateral light, from becoming curved. The summit for a length of .04 or .05 of an inch, though it is itself sensitive and curves towards the light, has only a slight power of causing the lower part to bend. Nor has the exclusion of light from the summit for a length of .1 of an inch a strong influence on the curvature of the lower part. On the other hand, an exclusion for a length of between .15 and .2 of an inch, or of the whole upper half, plainly prevents the lower and fully illuminated part from becoming curved in the manner (see Fig. 181) which invariably occurs when a free cotyledon is exposed to a lateral light. With very young seedlings the sensitive zone seems to extend rather lower down relatively to their height than in older seedlings. We must therefore conclude that when seedlings are freely exposed to a lateral light some influence is transmitted from the upper to the lower part, causing the latter to bend.

 

This conclusion is supported by what may be seen to occur on a small scale, especially with young cotyledons, without any artificial exclusion of the light; for they bend beneath the earth where no light can enter. Seeds of Phalaris were covered with a layer one-fourth of an inch in thickness of very fine sand, consisting of extremely minute grains of silex coated with [page 475]

oxide of iron. A layer of this sand, moistened to the same degree as that over the seeds, was spread over a glass-plate; and when the layer was .05

of an inch in thickness (carefully measured) no light from a bright sky could be seen to pass through it, unless it was viewed through a long blackened tube, and then a trace of light could be detected, but probably much too little to affect any plant. A layer .1 of an inch in thickness was quite impermeable to light, as judged by the eye aided by the tube. It may be worth adding that the layer, when dried, remained equally impermeable to light. This sand yielded to very slight pressure whilst kept moist, and in this state did not contract or crack in the least. In a first trial, cotyledons which had grown to a moderate height were exposed for 8 h.

before a paraffin lamp, and they became greatly bowed. At their bases on the shaded side opposite to the light, well-defined, crescentic, open furrows were formed, which (measured under a microscope with a micrometer) were from .02 to .03 of an inch in breadth, and these had evidently been left by the bending of the buried bases of the cotyledons towards the light. On the side of the light the cotyledons were in close contact with the sand, which was a very little heaped up. By removing with a sharp knife the sand on one side of the cotyledons in the line of the light, the bent portion and the open furrows were found to extend down to a depth of about .1 of an inch, where no light could enter. The chords of the short buried arcs formed in four cases angles of 11o, 13o, 15o, and 18o, with the perpendicular. By the following morning these short bowed portions had straightened themselves through apogeotropism.

 

In the next trial much younger cotyledons were similarly treated, but were exposed to a rather obscure lateral light. After some hours, a bowed cotyledon, .3 inch in height, had an open furrow on the shaded side .04

inch in breadth; another cotyledon, only .13 inch in height, had left a furrow .02 inch in breadth. But the most curious case was that of a cotyledon which had just protruded above the ground and was only .03 inch in height, and this was found to be bowed in the direction of the light to a depth of .2 of an inch beneath the surface. From what we know of the impermeability of this sand to light, the upper illuminated part in these several cases must have determined the curvature of the lower buried portions. But an apparent cause of doubt may be suggested: as the cotyledons are continually circumnutating, they tend to form a minute [page 476]

crack or furrow all round their bases, which would admit a little light on all sides; but this would not happen when they were illuminated laterally, for we know that they quickly bend towards a lateral light, and they then press so firmly against the sand on the illuminated side as to furrow it, and this would effectually exclude light on this side. Any light admitted on the opposite and shaded side, where an open furrow is formed, would tend to counteract the curvature towards the lamp or other source of the light.

It may be added, that the use of fine moist sand, which yields easily to pressure, was indispensable in the above experiments; for seedlings raised in common soil, not kept especially damp, and exposed for 9 h. 30 m. to a strong lateral light, did not form an open furrow at their bases on the shaded side, and were not bowed beneath the surface.

Perhaps the most striking proof of the action of the upper on the lower part of the cotyledons of Phalaris, when laterally illuminated, was afforded by the blackened glass-tubes (before alluded to) with very narrow stripes of the varnish scraped off on one side, through which a little light was admitted. The breadth of these stripes or slits varied between .01 and .02 inch (.25 and .51 mm.). Cotyledons with their upper halves enclosed in such tubes were placed before a south-west window, in such a position, that the scraped stripes did not directly face the window, but obliquely to one side. The seedlings were left exposed for 8 h., before the close of which time the many free seedlings in the same pots had become greatly bowed towards the window. Under these circumstances, the whole lower halves of the cotyledons, which had their summits enclosed in the tubes, were fully exposed to the light of the sky, whilst their upper halves received exclusively or chiefly diffused light from the room, and this only through a very narrow slit on one side. Now, if the curvature of the lower part had been determined by the illumination of this part, all the cotyledons assuredly would have become curved towards the window; but this was far from being the case. Tubes of the kind just described were placed on several occasions over the upper halves of 27 cotyledons; 14 of them remained all the time quite vertical; so that sufficient diffused light did not enter through the narrow slits to produce any effect whatever; and they behaved in the same manner as if their upper halves had been enclosed in completely blackened tubes. The lower halves of the 13

other cotyledons became bowed

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not directly in the line of the window, but obliquely towards it; one pointed at an angle of only 18o, but the remaining 12 at angles varying between 45o and 62o from the line of the window. At the commencement of the experiment, pins had been laid on the earth in the direction towards which the slits in the varnish faced; and in this direction alone a small amount of diffused light entered. At the close of the experiment, 7 of the bowed cotyledons pointed exactly in the line of the pins, and 6 of them in a line between that of the pins and that of the