The Power of Movement in Plants

by Charles Darwin; Sir Francis Darwin

Considering these cases, there can be no doubt that the leaves of this Oxalis, when allowed to assume their normal vertically dependent position at night, suffer much less from frost than those (23 in number) which had their upper surfaces exposed to the zenith.

Oxalis carnosa.—A plant of this Chilian species was exposed for 30 m. to a clear sky, the thermometer on the grass standing at –2° C., with some of its leaves pinned open, and not one leaf on the whole bushy plant was in the least injured. On the 16th of March another plant was similarly exposed for 30 m., when the temperature on the grass was only a little lower, viz., –3° to –4° C. Six of the leaves had been pinned open, and next morning five of them were found much browned. The plant was a large one, and none of the free leaves, which were asleep and depended vertically, were browned, excepting four very young ones. But three other leaves, though not browned, were in a rather flaccid condition, and retained their nocturnal position during the whole of the following day. In this case it was obvious that the leaves which were exposed horizontally to the zenith suffered most. This same pot was afterwards exposed for 35–40 m. on a slightly colder night, and every leaf, both the pinned open and the free ones, was killed. It may be added that two pots of O. corniculata (var. Atro-purpurea) were exposed for 2 h. and 3 h. to a clear sky with the temp. on grass –2° C., and none of the leaves, whether free or pinned open, were at all injured.

Arachis hypogoea.—Some plants in a pot were exposed at night for 30 m. to a clear sky, the temperature on the surrounding grass being –2° C., and on two nights afterwards they were again exposed to the same temperature, but this time during 1 h. 30 m. On neither occasion was a single leaf, whether pinned open or free, injured; and this surprised us much, considering its native tropical African home. Two plants were next exposed (March 16th) for 30 m. to a clear sky, the temperature of the surrounding grass being now lower, viz., between –3° and –4° C., and all four pinned-open leaves were killed and blackened. These two plants bore 22 other and free leaves (excluding some very young bud-like ones) and only two of these were killed and three somewhat injured; that is, 23 per cent. were either killed or injured, whereas all four pinned-open leaves were utterly killed.

On another night two pots with several plants were exposed for between 35 m. and 40 m. to a clear sky, and perhaps to a rather lower temperature, for a thermometer on a dial, 3 feet high, close by stood at –3.3° to –3.8° C. In one pot three leaves were pinned open, and all were badly injured; of the 44 free leaves, 26 were injured, that is, 59 per cent. In the other pot 3 leaves were pinned open and all were killed; four other leaves were prevented from sleeping by narrow strips of stiff paper gummed across them, and all were killed; of 24 free leaves, 10 were killed, 2 much injured, and 12 unhurt; that is, 50 per cent. of the free leaves were either killed or much injured. Taking the two pots together, we may say that rather more than half of the free leaves, which were asleep, were either killed or injured, whilst all the ten horizontally extended leaves, which had been prevented from going to sleep, were either killed or much injured.

Cassia floribunda.—A bush was exposed at night for 40 m. to a clear sky, the temperature on the surrounding grass being –2° C., and not a leaf was injured.[^[11]] It was again exposed on another night for 1 h., when the temperature of the grass was –4° C.; and now all the leaves on a large bush, whether pinned flat open or free, were killed, blackened, and shrivelled, with the exception of those on one small branch, low down, which was very slightly protected by the leaves on the branches above. Another tall bush, with four of its large compound leaves pinned out horizontally, was afterwards exposed (temp. of surrounding grass exactly the same, viz., –4° C.), but only for 30 m. On the following morning every single leaflet on these four leaves was dead, with both their upper and lower surfaces completely blackened. Of the many free leaves on the bush, only seven were blackened, and of these only a single one (which was a younger and more tender leaf than any of the pinned ones) had both surfaces of the leaflets blackened. The contrast in this latter respect was well shown by a free leaf, which stood between two pinned-open ones; for these latter had the lower surfaces of their leaflets as black as ink, whilst the intermediate free leaf, though badly injured, still retained a plain tinge of green on the lower surface of the leaflets. This bush exhibited in a striking manner the evil effects of the leaves not being allowed to assume at night their normal dependent position; for had they all been prevented from doing so, assuredly every single leaf on the bush would have been utterly killed by this exposure of only 30 m. The leaves whilst sinking downwards in the evening twist round, so that the upper surface is turned inwards, and is thus better protected than the outwardly turned lower surface. Nevertheless, it was always the upper surface which was more blackened than the lower, whenever any difference could be perceived between them; but whether this was due to the cells near the upper surface being more tender, or merely to their containing more chlorophyll, we do not know.

[11] Cassia laevigata was exposed to a clear sky for 35 m., and C. calliantha (a Guiana species) for 60 m., the temperature on the surrounding grass being –2° C., and neither was in the least injured. But when C. laevigata was exposed for 1 h., the temp. on the surrounding grass being between –3° and –4° C., every leaf was killed.

Melilotus officinalis.—A large pot with many plants, which had been kept during the winter in the greenhouse, was exposed during 5 h. at night to a slight frost and clear sky. Four leaves had been pinned out, and these died after a few days; but so did many of the free leaves. Therefore nothing certain could be inferred from this trial, though it indicated that the horizontally extended leaves suffered most. Another large pot with many plants was next exposed for 1 h., the temperature on the surrounding grass being lower, viz., -3° to –4° C. Ten leaves had been pinned out, and the result was striking, for on the following morning all these were found much injured or killed, and none of the many free leaves on the several plants were at all injured, with the doubtful exception of two or three very young ones.

Melilotus Italica.—Six leaves were pinned out horizontally, three with their upper and three with their lower surfaces turned to the zenith. The plants were exposed for 5 h. to a clear sky, the temperature on ground being about –1° C. Next morning the six pinned-open leaves seemed more injured even than the younger and more tender free ones on the same branches. The exposure, however, had been too long, for after an interval of some days many of the free leaves seemed in almost as bad a condition as the pinned-out ones. It was not possible to decide whether the leaves with their upper or those with their lower surfaces turned to the zenith had suffered most.

Melilotus suaveolens.—Some plants with 8 leaves pinned out were exposed to a clear sky during 2 h., the temperature on the surrounding grass being –2° C. Next morning 6 out of these 8 leaves were in a flaccid condition. There were about 150 free leaves on the plant, and none of these were injured, except 2 or 3 very young ones. But after two days, the plants having been brought back into the greenhouse, the 6 pinned-out leaves all recovered.

Melilotus Taurica.—Several plants were exposed for 5 h. during two nights to a clear sky and slight frost, accompanied by some wind; and 5 leaves which had been pinned out suffered more than those both above and below on the same branches which had gone to sleep. Another pot, which had likewise been kept in the greenhouse, was exposed for 35–40 m. to a clear sky, the temperature of the surrounding grass being between –3° and –4° C. Nine leaves had been pinned out, and all of these were killed. On the same plants there were 210 free leaves, which had been allowed to go to sleep, and of these about 80 were killed, i.e. only 38 per cent.