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

Lupinus (Tribe 2).—The palmate or digitate leaves of the species in this large genus sleep in three different manners. One of the simplest, is that all the leaflets become steeply inclined downwards at night, having been during the day extended horizontally. This is shown in the accompanying figures (Fig. 137), of a leaf of L. pilosus, as seen during the day from vertically above, and of another leaf asleep with the leaflets inclined downwards. As in this position they are crowded together, and as they do not become folded like those in the genus Oxalis, they cannot occupy a vertically dependent position; but they are often inclined at an angle of 50o beneath the horizon. In this species, whilst the leaflets are sinking, the petioles rise up, in two instances when the angles were measured to the extent of 23o. The leaflets of L. sub-carnosus and arboreus, which were horizontal during the day, sank down at night in nearly the same manner; the former to an angle of 38o and the latter of 36o, beneath the horizon; but their petioles

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did not move in any plainly perceptible degree. It is, however, quite possible, as we shall presently see, that if a large number of plants of the three foregoing and of the following species Fig. 137. Lupinus pilosus: A, leaf seen from vertically above in daytime; B, leaf asleep, seen laterally at night.

were to be observed at all seasons, some of the leaves would be found to sleep in a different manner.

In the two following species the leaflets, instead of moving downwards, rise at night. With L. Hartwegii some stood at noon at a mean angle of 36o above the horizon, and at night at 51o, thus forming together a hollow cone with moderately steep sides. The petiole of one leaf rose 14o and of a second 11o at night. With L. luteus a leaflet rose from 47o at noon to 65o above the horizon at night, and another on a distinct leaf rose from 45o to 69o. The petioles, however, sink at night to a small extent, viz., in three instances by 2o, 6o, and 9o 30 seconds. Owing to this movement of the petioles, the outer and longer leaflets have to bend up a little more than the shorter and inner ones, in order that all should stand symmetrically at night. We shall presently see that some leaves on the same individual plants of L. luteus sleep in a very different manner.

We now come to a remarkable position of the leaves when asleep, which is common to several species of Lupines. On the same leaf the shorter leaflets, which generally face the centre of the plant, sink at night, whilst the longer ones on the opposite side rise; the intermediate and lateral ones merely twisting on their own axes. But there is some variability with respect to which leaflets rise or fall. As might have been expected from such diverse and complicated movements, the [page 342]

base of each leaflet is developed (at least in the case of L. luteus) into a pulvinus. The result is that all the leaflets on the same leaf stand at night more or less highly inclined, or even quite vertically, forming in this latter case a vertical star. This occurs with the leaves of a species purchased under the name of

Fig. 138. Lupinus pubescens: A, leaf viewed laterally during the day; B, same leaf at night; C, another leaf with the leaflet forming a vertical star at night. Figures reduced.

L. pubescens; and in the accompanying figures we see at A (Fig. 138) the leaves in their diurnal position; and at B the same plant at night with the two upper leaves having their leaflets almost vertical. At C another leaf, viewed laterally, is shown with the leaflets quite vertical. It is chiefly or exclusively the youngest leaves which form at night vertical stars. But there

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is much variability in the position of the leaves at night on the same plant; some remaining with their leaflets almost horizontal, others forming more or less highly inclined or vertical stars, and some with all their leaflets sloping downwards, as in our first class of cases. It is also a remarkable fact, that although all the plants produced from the same lot of seeds were identical in appearance, yet some individuals at night had the leaflets of all their leaves arranged so as to form more or less highly inclined stars; others had them all sloping downwards and never forming a star; and others, again, retained them either in a horizontal position or raised them a little.

We have as yet referred only to the different positions of the leaflets of L. pubescens at night; but the petioles likewise differ in their movements.

That of a young leaf which formed a highly inclined star at night, stood at noon at 42o above the horizon, and during the night at 72o, so had risen 30o. The petiole of another leaf, the leaflets of which occupied a similar position at night, rose only 6o. On the other hand, the petiole of a leaf with all its leaflets sloping down at night, fell at this time 4o. The petioles of two rather older leaves were subsequently observed; both of which stood during the day at exactly the same angle, viz., 50o above the horizon, and one of these rose 7o - 8o, and the other fell 3o - 4o at night.

We meet with cases like that of L. pubescens with some other species. On a single plant of L. mutabilis some leaves, which stood horizontally during the day, formed highly inclined stars at night, and the petiole of one rose 7o. Other leaves which likewise stood horizontally during the day, had at night all their leaflets sloping downwards at 46o beneath the horizon, but their petioles had hardly moved. Again, L. luteus offered a still more remarkable case, for on two leaves, the leaflets which stood at noon at about 45o above the horizon, rose at night to 65o and 69o, so that they formed a hollow cone with steep sides. Four leaves on the same plant, which had their leaflets horizontal at noon, formed vertical stars at night; and three other leaves equally horizontal at noon, had all their leaflets sloping downwards at night. So that the leaves on this one plant assumed at night three different positions. Though we cannot account for this fact, we can see that such a stock might readily give birth to species having widely different nyctitropic habits.

Little more need be said about the sleep of the species of Lupinus; several, namely, L. polyphyllus, nanus, Menziesii, speciosus, [page 344]

and albifrons, though observed out of doors and in the greenhouse, did not change the position of their leaves sufficiently at night to be said to sleep. From observations made on two sleeping species, it appears that, as with Tropaeolum majus, the leaves must be well illuminated during the day in order to sleep at night. For several plants, kept all day in a sitting-room with north-east windows, did not sleep at night; but when the pots were placed on the following day out of doors, and were brought in at night, they slept in the usual manner. the trial was repeated on the following day and night with the same result.

Some observations were made on the circumnutation of the leaves of L.

luteus and arboreus. It will suffice to say that the leaflets of the latter exhibited a double oscillation in the course of 24 h.; for they fell from the early morning until 10.15 A.M., then rose and zigzagged greatly till 4

P.M., after which hour the great nocturnal fall commenced. By 8 A.M. on the following morning the leaflets had risen to their proper height. We have seen in the fourth chapter, that the leaves of Lupinus speciosus, which do not sleep, circumnutate to an extraordinary extent, making many ellipses in the course of the day.

Cytisus (Tribe 2), Trigonella and Medicago (Tribe 3).—Only Fig. 139. Medicago marina: A, leaves during the day; B, leaves asleep at night.

a few observations were made on these three genera. The petioles on a young plant, about a foot in height, of Cytisus fragrans rose at night, on one occasion 23o and on another 33o. The three leaflets also bend upwards, and at the same time

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approach each other, so that the base of the central leaflet overlaps the bases of the two lateral leaflets. They bend up so much that they press against the stem; and on looking down on one of these young plants from vertically above, the lower surfaces of the leaflets are visible; and thus their upper surfaces, in accordance with the general rule, are best protected from radiation. Whilst the leaves on these young plants were thus behaving, those on an old bush in full flower did not sleep at night.

Trigonella Cretica resembles a Melilotus in its sleep, which will be immediately described. According to M. Royer,* the leaves of Medicago maculata rise up at night, and “se renversent un peu de mani�re � presenter obliquement au ciel leur face inf�rieure.” A drawing is here given (Fig.

139) of the leaves of M. marina awake and asleep; and this would almost serve for Cytisus fragrans in the same two states.

Melilotus (Tribe 3).—The species in this genus sleep in a remarkable manner. The three leaflets of each leaf twist through an angle of 90o, so that their blades stand vertically at night with one lateral edge presented to the zenith (Fig. 140). We shall best understand the other and more complicated movements, if we imagine ourselves always to hold the leaf with the tip of the terminal leaflet pointed to the north. The leaflets in becoming vertical at night could of course twist so that their upper surfaces should face to either side; but the two lateral leaflets always twist so that this surface tends to face the north, but as they move at the same time towards the terminal leaflet, the upper surface of the one faces about N.N.W., and that of the other N.N.E. The terminal leaflet behaves differently, for it twists to either side, the upper surface facing sometimes east and sometimes west, but rather more commonly west than east.

The terminal leaflet also moves in another and more remarkable manner, for whilst its blade is twisting and becoming vertical, the whole leaflet bends to one side, and invariably to the side towards which the upper surface is directed; so that if this surface faces the west the whole leaflet bends to the west, until it comes into contact with the upper and vertical surface of the western lateral leaflet. Thus the upper surface of the terminal and of one of the two lateral leaflets is well protected.

The fact of the terminal leaflet twisting indifferently to either * ‘Annales des Sc. Nat. Bot.’ (5th series), ix. 1868, p. 368.

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side and afterwards bending to the same side, seemed to us so remarkable, that we endeavoured to discover the cause. We imagined that at the commencement of the movement it might be determined by one of the two halves of the leaflet being a little heavier than the other. Therefore bits of wood were gummed on one side of several leaflets, but this produced no effect; and they continued to twist in the same direction as Fig. 140. Melilotus officinalis: A, leaf during the daytime. B, another leaf asleep. C, a leaf asleep as viewed from vertically above; but in this case the terminal leaflet did not happen to be in such close contact with the lateral one, as is usual.