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

two ways,* firstly, by means of pulvini which become, as Pfeffer has shown, alternately more turgescent on opposite sides; and secondly, by increased growth along one side of the petiole or midrib, and then on the opposite side, as was first proved by Batalin.** But as it has been shown by De Vries*** that in these latter cases increased growth is preceded by the increased turgescence of the cells, the difference between the above two means of movement is much diminished, and consists chiefly in the turgescence of the cells of a fully developed pulvinus, not being followed by growth. When the movements of leaves or cotyledons, furnished with a pulvinus and destitute of one, are compared, they are seen to be closely similar, and are apparently effected for the same purpose. Therefore, with our object in view, it does not appear advisable to separate the above two sets of cases into two distinct classes. There is, however, one important distinction between them, namely, that movements effected by growth on the alternate sides, are confined to young growing leaves, whilst those effected by means of a pulvinus last for a long time. We have already seen well-marked instances of this latter fact with cotyledons, and so it is with leaves, as has been observed by Pfeffer and by ourselves. The long endurance of the nyctitropic movements when effected by the aid of pulvini indicates, in addition to the evidence already advanced, the functional import-

 

* This distinction was first pointed out (according to Pfeffer, ‘Die Periodischen Bewegungen der Blattorgane,’ 1875, p. 161) by Dassen in 1837.

 

** ‘Flora,’ 1873, p. 433.

 

*** ‘Bot. Zeitung,’ 1879, Dec. 19th, p. 830.

 

[page 284]

ance of such movements to the plant. There is another difference between the two sets of cases, namely, that there is never, or very rarely, any torsion of the leaves, excepting when a pulvinus is present;* but this statement applies only to periodic and nyctitropic movements as may be inferred from other cases given by Frank.**

The fact that the leaves of many plants place themselves at night in widely different positions from what they hold during the day, but with the one point in common, that their upper surfaces avoid facing the zenith, often with the additional fact that they come into close contact with opposite leaves or leaflets, clearly indicates, as it seems to us, that the object gained is the protection of the upper surfaces from being chilled at night by radiation. There is nothing improbable in the upper surface needing protection more than the lower, as the two differ in function and structure. All gardeners know that plants suffer from radiation. It is this and not cold winds which the peasants of Southern Europe fear for their olives.*** Seedlings are often protected from radiation by a very thin covering of straw; and fruit-trees on walls by a few fir-branches, or even by a fishing-net, suspended over them. There is a variety of the gooseberry,**** the flowers of which from being produced before the leaves, are not protected by them from radiation, and consequently often fail to yield fruit. An excellent observer***** has remarked * Pfeffer, ‘Die Period. Beweg. der Blattorgane.’ 1875, p. 159.

 

** ‘Die Nat. Wagerechte Richtung von Pflanzentheilen,’ 1870, p. 52

 

*** Martins in ‘Bull. Soc. Bot. de France,’ tom. xix. 1872. Wells, in his famous ‘Essay on Dew,’ remarks that an exposed thermometer rises as soon as even a fleecy cloud, high in the sky, passes over the zenith.

 

**** ‘Loudon’s Gardener’s Mag.,’ vol. iv. 1828, p. 112.

 

***** Mr. Rivers in ‘Gardener’s Chron.,’ 1866, p. 732.

[page 285]

 

that one variety of the cherry has the petals of its flowers much curled backwards, and after a severe frost all the stigmas were killed; whilst at the same time, in another variety with incurved petals, the stigmas were not in the least injured.

 

This view that the sleep of leaves saves them from being chilled at night by radiation, would no doubt have occurred to Linnaeus, had the principle of radiation been then discovered; for he suggests in many parts of his ‘Somnus Plantarum’ that the position of the leaves at night protects the young stems and buds, and often the young inflorescence, against cold winds. We are far from doubting that an additional advantage may be thus gained; and we have observed with several plants, for instance, Desmodium gyrans, that whilst the blade of the leaf sinks vertically down at night, the petiole rises, so that the blade has to move through a greater angle in order to assume its vertical position than would otherwise have been necessary; but with the result that all the leaves on the same plant are crowded together as if for mutual protection.

 

We doubted at first whether radiation would affect in any important manner objects so thin as are many cotyledons and leaves, and more especially affect differently their upper and lower surfaces; for although the temperature of their upper surfaces would undoubtedly fall when freely exposed to a clear sky, yet we thought that they would so quickly acquire by conduction the temperature of the surrounding air, that it could hardly make any sensible difference to them, whether they stood horizontally and radiated into the open sky, or vertically and radiated chiefly in a lateral direction towards neighbouring plants and other objects. We endeavoured, therefore, to ascertain something on this head by preventing the leaves [page 286]

of several plants from going to sleep, and by exposing to a clear sky when the temperature was beneath the freezing-point, these, as well as the other leaves on the same plants which had already assumed their nocturnal vertical position. Our experiments show that leaves thus compelled to remain horizontal at night, suffered much more injury from frost than those which were allowed to assume their normal vertical position. It may, however, be said that conclusions drawn from such observations are not applicable to sleeping plants, the inhabitants of countries where frosts do not occur. But in every country, and at all seasons, leaves must be exposed to nocturnal chills through radiation, which might be in some degree injurious to them, and which they would escape by assuming a vertical position.

 

In our experiments, leaves were prevented from assuming their nyctitropic position, generally by being fastened with the finest entomological pins (which did not sensibly injure them) to thin sheets of cork supported on sticks. But in some instances they were fastened down by narrow strips of card, and in others by their petioles being passed through slits in the cork. The leaves were at first fastened close to the cork, for as this is a bad conductor, and as the leaves were not exposed for long periods, we thought that the cork, which had been kept in the house, would very slightly warm them; so that if they were injured by the frost in a greater degree than the free vertical leaves, the evidence would be so much the stronger that the horizontal position was injurious. But we found that when there was any slight difference in the result, which could be detected only occasionally, the leaves which had been fastened closely down suffered rather more than those fastened with very long and [page 287]

thin pins, so as to stand from � to 3/4 inch above the cork. This difference in the result, which is in itself curious as showing what a very slight difference in the conditions influences the amount of injury inflicted, may be attributed, as we believe, to the surrounding warmer air not circulating freely beneath the closely pinned leaves and thus slightly warming them. This conclusion is supported by some analogous facts hereafter to be given.

 

We will now describe in detail the experiments which were tried. These were troublesome from our not being able to predict how much cold the leaves of the several species could endure. Many plants had every leaf killed, both those which were secured in a horizontal position and those which were allowed to sleep—that is, to rise up or sink down vertically. Others again had not a single leaf in the least injured, and these had to be re-exposed either for a longer time or to a lower temperature.

 

[Oxalis acetosella.—A very large pot, thickly covered with between 300 and 400 leaves, had been kept all winter in the greenhouse. Seven leaves were pinned horizontally open, and were exposed on March 16th for 2 h. to a clear sky, the temperature on the surrounding grass being -4o C. (24o to 25o F.). Next morning all seven leaves were found quite killed, so were many of the free ones which had previously gone to sleep, and about 100 of them, either dead or browned and injured were picked off. Some leaves showed that they had been slightly injured by not expanding during the whole of the next day, though they afterwards recovered. As all the leaves which were pinned open were killed, and only about a third or fourth of the others were either killed or injured, we had some little evidence that those which were prevented from assuming their vertically dependent position suffered most.

 

The following night (17th) was clear and almost equally cold (-3o to -4o C.

on the grass), and the pot was again exposed, but this time for only 30 m.

Eight leaves had been pinned out,

[page 288]

and in the morning two of them were dead, whilst not a single other leaf on the many plants was even injured.

 

On the 23rd the pot was exposed for 1 h. 30 m., the temperature on the grass being only -2o C., and not one leaf was injured: the pinned open leaves, however, all stood from � to 3/4 of an inch above the cork.

 

On the 24th the pot was again placed on the ground and exposed to a clear sky for between 35 m. and 40 m. By a mistake the thermometer was left on an adjoining sun-dial 3 feet high, instead of being placed on the grass; it recorded 25o to 26o F. (-3.3o to -3.8o C.), but when looked at after 1 h.

had fallen to 22o F. (-5.5o C.); so that the pot was perhaps exposed to rather a lower temperature than on the two first occasions. Eight leaves had been pinned out, some close to the cork and some above it, and on the following morning five of them (i.e. 63 per cent.) were found killed. By counting a portion of the leaves we estimated that about 250 had been allowed to go to sleep, and of these about 20 were killed (i.e. only 8 per cent.), and about 30 injured.

 

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 -2o 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., -3o to -4o 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