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

The cotyledons of several plants were prepared for trial, but the weather was mild and we succeeded only in a single instance in having seedlings of the proper age on nights which were clear and cold. The cotyledons of 6

seedlings of Mimosa pudica were fastened open on cork and were thus exposed for 1 h. 45 m. to a clear sky, with the temperature on the surrounding ground at 29o F.; of these, 3 were killed. Two other seedlings, after their cotyledons had risen up and had closed together, were bent over and fastened so that they stood horizontally, with the lower surface of one cotyledon fully exposed to the zenith, and both were killed. Therefore of the 8 seedlings thus tried 5, or more than half, were killed. Seven other seedlings with their cotyledons in their normal nocturnal position, viz., vertical and closed, were exposed at the same time, and of these only 2

were killed.* Hence it appears, as far as these few trials tell anything, that the vertical position at night of the cotyledons of Mimosa pudica protects them to a certain degree from the evil effects of radiation and cold.]

Concluding Remarks on the Radiation from Leaves at Night.—We exposed on two occasions during the summer to a clear sky several pinned-open leaflets of Trifolium pratense, which naturally rise at night, and of Oxalis purpurea, which naturally sink at night (the plants growing out of doors), and looked at

* We were surprised that young seedlings of so tropical a plant as Mimosa pudica were able to resist, as well as they did, exposure for 1 hr. 45 m.

to a clear sky, the temperature on the surrounding ground being 29o F. It may be added that seedlings of the Indian ‘Cassia pubescens’ were exposed for 1 h. 30 m. to a clear sky, with the temp. on the surrounding ground at -2o C., and they were not in the least injured.

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them early on several successive mornings, after they had assumed their diurnal positions. The difference in the amount of dew on the pinned-open leaflets and on those which had gone to sleep was generally conspicuous; the latter being sometimes absolutely dry, whilst the leaflets which had been horizontal were coated with large beads of dew. This shows how much cooler the leaflets fully exposed to the zenith must have become, than those which stood almost vertically, either upwards or downwards, during the night.

From the several cases above given, there can be no doubt that the position of the leaves at night affects their temperature through radiation to such a degree, that when exposed to a clear sky during a frost, it is a question of life and death. We may therefore admit as highly probable, seeing that their nocturnal position is so well adapted to lessen radiation, that the object gained by their often complicated sleep movements, is to lessen the degree to which they are chilled at night. It should be kept in mind that it is especially the upper surface which is thus protected, as it is never directed towards the zenith, and is often brought into close contact with the upper surface of an opposite leaf or leaflet.

We failed to obtain sufficient evidence, whether the better protection of the upper surface has been gained from its being more easily injured than the lower surface, or from its injury being a greater evil to the plant.

That there is some difference in constitution between the two surfaces is shown by the following cases. Cassia floribunda was exposed to a clear sky on a sharp frosty night, and several leaflets which had assumed their nocturnal dependent position with their lower surfaces turned outwards so as to be

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exposed obliquely to the zenith, nevertheless had these lower surfaces less blackened than the upper surfaces which were turned inwards and were in close contact with those of the opposite leaflets. Again, a pot full of plants of Trifolium resupinatum, which had been kept in a warm room for three days, was turned out of doors (Sept. 21st) on a clear and almost frosty night. Next morning ten of the terminal leaflets were examined as opaque objects under the microscope. These leaflets, in going to sleep, either turn vertically upwards, or more commonly bend a little over the lateral leaflets, so that their lower surfaces are more exposed to the zenith than their upper surfaces. Nevertheless, six of these ten leaflets were distinctly yellower on the upper than on the lower and more exposed surface. In the remaining four, the result was not so plain, but certainly whatever difference there was leaned to the side of the upper surface having suffered most.

It has been stated that some of the leaflets experimented on were fastened close to the cork, and others at a height of from � to 3/4 of an inch above it; and that whenever, after exposure to a frost, any difference could be detected in their states, the closely pinned ones had suffered most. We attributed this difference to the air, not cooled by radiation, having been prevented from circulating freely beneath the closely pinned leaflets. That there was really a difference in the temperature of leaves treated in these two different methods, was plainly shown on one occasion; for after the exposure of a pot with plants of Melilotus dentata for 2 h. to a clear sky (the temperature on the surrounding grass being -2o C.), it was manifest that more dew had congealed into hoar-frost on the closely pinned leaflets, than on those which stood horizontally

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a little above the cork. Again, the tips of some few leaflets, which had been pinned close to the cork, projected a little beyond the edge, so that the air could circulate freely round them. This occurred with six leaflets of Oxalis acetosella, and their tips certainly suffered rather less then the rest of the same leaflets; for on the following morning they were still slightly green. The same result followed, even still more clearly, in two cases with leaflets of Melilotus officinalis which projected a little beyond the cork; and in two other cases some leaflets which were pinned close to the cork were injured, whilst other free leaflets on the same leaves, which had not space to rotate and assume their proper vertical position, were not at all injured.

Another analogous fact deserves notice: we observed on several occasions that a greater number of free leaves were injured on the branches which had been kept motionless by some of their leaves having been pinned to the corks, than on the other branches. This was conspicuously the case with those of Melilotus Petitpierreana, but the injured leaves in this instance were not actually counted. With Arachis hypogaea, a young plant with 7

stems bore 22 free leaves, and of these 5 were injured by the frost, all of which were on two stems, bearing four leaves pinned to the cork-supports.

With Oxalis carnosa, 7 free leaves were injured, and every one of them belonged to a cluster of leaves, some of which had been pinned to the cork.

We could account for these cases only by supposing that the branches which were quite free had been slightly waved about by the wind, and that their leaves had thus been a little warmed by the surrounding warmer air. If we hold our hands motionless before a hot fire, and then wave them about, we [page 297]

immediately feel relief; and this is evidently an analogous, though reversed, case. These several facts—in relation to leaves pinned close to or a little above the cork-supports—to their tips projecting beyond it—

and to the leaves on branches kept motionless—seem to us curious, as showing how a difference, apparently trifling, may determine the greater or less injury of the leaves. We may even infer as probable that the less or greater destruction during a frost of the leaves on a plant which does not sleep, may often depend on the greater or less degree of flexibility of their petioles and of the branches which bear them.

NYCTITROPIC OR SLEEP MOVEMENTS OF COTYLEDONS.

We now come to the descriptive part of our work, and will begin with cotyledons, passing on to leaves in the next chapter. We have met with only two brief notices of cotyledons sleeping. Hofmeister,* after stating that the cotyledons of all the observed seedlings of the Caryophylleae (Alsineae and Sileneae) bend upwards at night (but to what angle he does not state), remarks that those of Stellaria media rise up so as to touch one another; they may therefore safely be said to sleep. Secondly, according to Ramey**, the cotyledons of Mimosa pudica and of Clianthus Dampieri rise up almost vertically at night and approach each other closely. It has been shown in a previous chapter that the cotyledons of a large number of plants bend a little upwards at night, and we here have to meet the difficult question at what inclination may they be said to sleep? According to the view which we maintain, no movement deserves to be called * ‘Die Lehre von der Pflanzenzelle,’ 1867, p. 327.

** ‘Adansonia,’ March 10th, 1869.

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nyctitropic, unless it has been acquired for the sake of lessening radiation; but this could be discovered only by a long series of experiments, showing that the leaves of each species suffered from this cause, if prevented from sleeping. We must therefore take an arbitrary limit. If a cotyledon or leaf is inclined at 60o above or beneath the horizon, it exposes to the zenith about one-half of its area; consequently the intensity of its radiation will be lessened by about half, compared with what it would have been if the cotyledon or leaf had remained horizontal. This degree of diminution certainly would make a great difference to a plant having a tender constitution. We will therefore speak of a cotyledon and hereafter of a leaf as sleeping, only when it rises at night to an angle of about 60o, or to a still higher angle, above the horizon, or sinks beneath it to the same amount. Not but that a lesser diminution of radiation may be advantageous to a plant, as in the case of Datura stramonium, the cotyledons of which rose from 31o at noon to 55o at night above the horizon. The Swedish turnip may profit by the area of its leaves being reduced at night by about 30 per cent., as estimated by Mr. A.

S. Wilson; though in this case the angle through which the leaves rose was not observed. On the other hand, when the angular rise of cotyledons or of leaves is small, such as less than 30o, the diminution of radiation is so slight that it probably is of no significance to the plant in relation to radiation. For instance, the cotyledons of Geranium Ibericum rose at night to 27o above the horizon, and this would lessen radiation by only 11 per cent.: those of Linum Berendieri rose to 33o, and this would lessen radiation by 16 per cent.

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respect to the sleep of cotyledons. In certain cases, the cotyledons whilst young diverge during the day to only a very moderate extent, so that a small rise at night, which we know occurs with the cotyledons of many plants, would necessarily cause them to assume a vertical or nearly vertical position at night; and in this case it would be rash to infer that the movement was effected for any special purpose. On this account we hesitated long whether we should introduce several Cucurbitaceous plants into the following list; but from reasons, presently to be given, we thought that