The Power of Movement in Plants - Charles Darwin (top reads txt) 📗

Those on the opposite sides of the same petiole do not come into close contact at night, as occurs with the leaflets of so many Leguminosae but are separated by an open furrow; nor could they exactly coincide, as they stand alternately with respect to one another.

 

The circumnutation of the petiole of a leaf 3/4 of an inch in length, on an upright branch, was observed during 36h., and is shown in the preceding diagram (Fig. 136). On the first morning, the leaf fell a little and then rose until 1 P.M., and this was probably due to its being now illuminated through a skylight from above; it then circumnutated on a very small scale round the same spot until about 4 P.M., when the great evening fall commenced. During the latter part of the night or very early on the next morning the leaf rose again. On the second day it fell during the morning till 1 P.M., and this no doubt is its normal habit. From 1 to 4 P.M. it rose in a zigzag line, and soon afterwards the great evening fall commenced. It thus completed a double oscillation during the 24 h.

 

The specific name given to this plant by Ruiz and Pavon, indicates that in its native arid home it is affected in some manner by the dryness or dampness of the atmosphere.* In the Botanic Garden at W�rzburg, there was a plant in a pot out of doors which was daily watered, and another in the open ground which was never watered. After some hot and dry weather there was a great difference in the state of the leaflets on these two plants; those on the unwatered plant in the open ground remaining half, * ‘Systema Veg. Florae Peruvianae et Chilensis,’ tom. i. p. 95, 1798. We cannot understand the account given by the authors of the behaviour of this plant in its native home. There is much about its power of foretelling changes in the weather; and it appears as if the brightness of the sky largely determined the opening and closing of the leaflets.

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or even quite, closed during the day. But twigs cut from this bush, with their ends standing in water, or wholly immersed in it, or kept in damp air under a bell-glass, opened their leaves though exposed to a blazing sun; whilst those on the plant in the ground remained closed. The leaves on this same plant, after some heavy rain, remained open for two days; they then became half closed during two days, and after an additional day were quite closed. This plant was now copiously watered, and on the following morning the leaflets were fully expanded. The other plant growing in a pot, after having been exposed to heavy rain, was placed before a window in the Laboratory, with its leaflets open, and they remained so during the daytime for 48 h.; but after an additional day were half closed. The plant was then watered, and the leaflets on the two following days remained open. On the third day they were again half closed, but on being again watered remained open during the two next days. From these several facts we may conclude that the plant soon feels the want of water; and that as soon as this occurs, it partially or quite closes its leaflets, which in their then imbricated condition expose a small surface to evaporation. It is therefore probable that this sleep-like movement, which occurs only when the ground is dry, is an adaptation against the loss of moisture.

 

A bush about 4 feet in height, a native of Chili, which was thickly covered with leaves, behaved very differently, for during the day it never closed its leaflets. On July 6th the earth in the small pot in which it grew appeared extremely dry, and it was given a very little water. After 21 and 22 days (on the 27th and 28th), during the whole of which time the plant did not receive a drop of water, the leaves began to droop, but they showed no signs of closing during the day. It appeared almost incredible that any plant, except a fleshy one, could have kept alive in soil so dry, which resembled the dust on a road. On the 29th, when the bush was shaken, some leaves fell off, and the remaining ones were unable to sleep at night. It was therefore moderately watered, as well as syringed, late in the evening.

On the next morning (30th) the bush looked as fresh as ever, and at night the leaves went to sleep. It may be added that a small branch while growing on the bush was enclosed, by means of a curtain of bladder, during 13 days in a large bottle half full of quicklime, so that the air within must have been intensely dry; yet the leaves on this branch did not suffer in the [page 338]

least, and did not close at all during the hottest days. Another trial was made with the same bush on August 2nd and 6th (the soil appearing at this latter date extremely dry), for it was exposed out of doors during the whole day to the wind, but the leaflets showed no signs of closing. The Chilian form therefore differs widely from the one at W�rzburg, in not closing its leaflets when suffering from the want of water; and it can live for a surprisingly long time without water.

 

Tropaeolum majus (?) (cultivated var.) (Tropaeoleae).—Several plants in pots stood in the greenhouse, and the blades of the leaves which faced the front-lights were during the day highly inclined and at night vertical; whilst the leaves on the back of the pots, though of course illuminated through the roof, did not become vertical at night. We thought, at first, that this difference in their positions was in some manner due to heliotropism, for the leaves are highly heliotropic. The true explanation, however, is that unless they are well illuminated during at least a part of the day they do not sleep at night; and a little difference in the degree of illumination determines whether or not they shall become vertical at night. We have observed no other so well-marked a case as this, of the influence of previous illumination on nyctitropic movements. The leaves present also another peculiarity in their habit of rising or awaking in the morning, being more strongly fixed or inherited than that of sinking or sleeping at night. The movements are caused by the bending of an upper part of the petiole, between � and 1 inch in length; but the part close to the blade, for about 1/4 of an inch in length, does not bend and always remains at right angles to the blade. The bending portion does not present any external or internal difference in structure from the rest of the petiole.

We will now give the experiments on which the above conclusions are founded.

 

A large pot with several plants was brought on the morning of Sept. 3rd out of the greenhouse and placed before a north-east window, in the same position as before with respect to the light, as far as that was possible.

On the front of the plants, 24 leaves were marked with thread, some of which had their blades horizontal, but the greater number were inclined at about 45o, beneath the horizon; at night all these, without exception, became vertical. Early on the following morning (4th) they reassumed their former positions, and at night again became vertical. On the 5th the shutters were opened at 6.15 A.M., and

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by 8.18 A.M., after the leaves had been illuminated for 2 h. 3 m. and had acquired their diurnal position, they were placed in a dark cupboard. They were looked at twice during the day and thrice in the evening, the last time at 10.30 P.M., and not one had become vertical. At 8 A.M. on the following morning (6th) they still retained the same diurnal position, and were now replaced before the north-east window. At night all the leaves which had faced the light had their petioles curved and their blades vertical; whereas none of the leaves on the back of the plants, although they had been moderately illuminated by the diffused light of the room, were vertical. They were now at night placed in the same dark cupboard; at 9 A.M. on the next morning (7th) all those which had been asleep had reassumed their diurnal position. The pot was then placed for 3 h. in the sunshine, so as to stimulate the plants; at noon they were placed before the same north-east window, and at night the leaves slept in the usual manner and awoke on the following morning. At noon on this day (8th) the plants, after having been left before the north-east window for 5 h. 45 m.

and thus illuminated (though not brightly, as the sky was cloudy during the whole time), were replaced in the dark cupboard, and at 3 P.M. the position of the leaves was very little, if at all, altered, so that they are not quickly affected by darkness; but by 10.15 P.M. all the leaves which had faced the north-east sky during the 5 h. 45 m. of illumination stood vertical, whereas those on the back of the plant retained their diurnal position. On the following morning (9th) the leaves awoke as on the two former occasions in the dark, and they were kept in the dark during the whole day; at night a very few of them became vertical, and this was the one instance in which we observed any inherited tendency or habit in this plant to sleep at the proper time. That it was real sleep was shown by these same leaves reassuming their diurnal position on the following morning (10th) whilst still kept in the dark.

 

The pot was then (9.45 A.M. 10th) replaced, after having been kept for 36

h. in darkness, before the north-east window; and at night the blades of all the leaves (excepting a few on the back of the plants) became conspicuously vertical.

At 6.45 A.M. (11th) after the plants had been illuminated on the same side as before during only 25 m., the pot was turned round, so that the leaves which had faced the light now faced the interior of the room, and not one of these went to sleep at night;

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whilst some, but not many, of those which had formerly stood facing the back of the room and which had never before been well illuminated or gone to sleep, now assumed a vertical position at night. On the next day (12th) the plant was turned round into its original position, so that the same leaves faced the light as formerly, and these now went to sleep in the usual manner. We will only add that with some young seedlings kept in the greenhouse, the blades of the first pair of true leaves (the cotyledons being hypogean) stood during the day almost horizontally and at night almost vertically.

 

A few observations were subsequently made on the circumnutation of three leaves, whilst facing a north-east window; but the tracings are not given, as the leaves moved somewhat towards the light. It was, however, manifest that they rose and fell more than once during the daytime, the ascending and descending lines being in parts extremely zigzag. The nocturnal fall commenced about 7 P.M., and the leaves had risen considerably by 6.45 A.M.

on the following morning.

 

Leguminosae.—This Family includes many more genera with sleeping species than all the other families put together. The number of the tribes to which each genus belongs, according to Bentham and Hooker’s arrangement, has been added.

 

Crotolaria (sp.?) (Tribe 2).—This plant is monophyllous, and we are informed by Mr. T. Thiselton Dyer