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

the petiole. The second pair of leaflets likewise moves a little backwards, but less than the terminal pair; and the third pair moves vertically downwards, or even a little forwards. Thus all the leaflets, in those species which bear only 3 or 4 pairs, tend to form a single packet, with their upper surfaces in contact, and their lower surfaces turned outwards. Lastly, the main petiole rises at night, but with leaves of different ages to very different degrees, namely some rose through an angle of only 12o, and others as much as 41o.

 

Cassia calliantha.—The leaves bear a large number of leaflets, which move at night in nearly the same manner as just described; but the petioles apparently do not rise, and one which was carefully observed certainly fell 3o.

Cassia pubescens.—The chief difference in the nyctitropic Fig. 155. Cassia pubescens: A, upper part of plant during the day; B, same plant at night. Figures reduced from photographs.

 

movements of this species, compared with those of the former species, consists in the leaflets not rotating nearly so much; [page 372]

therefore their lower surfaces face but little outwards at night. The petioles, which during the day are inclined only a little above the horizon, rise at night in a remarkable manner, and stand nearly or quite vertically. This, together with the dependent position of the leaflets, makes the whole plant wonderfully compact at night. In the two foregoing figures, copied from photographs, the same plant is represented awake and asleep (Fig. 155), and we see how different is its appearance.

 

Cassia mimosoides.—At night the numerous leaflets on each leaf rotate on their axes, and their tips move towards the apex of the leaf; they thus become imbricated with their lower surfaces directed upwards, and with their midribs almost parallel to the petiole. Consequently, this species differs from all the others seen by us, with the exception of the following one, in the leaflets not sinking down at night. A petiole, the movement of which was measured, rose 8o at night.

 

Cassia Barclayana.—The leaflets of this Australian species are numerous, very narrow, and almost linear. At night they rise up a little, and also move towards the apex of the leaf. For instance, two opposite leaflets which diverged from one another during the day at an angle of 104o, diverted at night only 72o; so that each had risen 16o above its diurnal position. The petiole of a young leaf rose at night 34o, and that of an older leaf 19o. Owing to the slight movement of the leaflets and the considerable movement of the petiole, the bush presents a different appearance at night to what it does by day; yet the leaves can hardly be said to sleep.

 

The circumnutating movements of the leaves of C. floribunda, calliantha, and pubescens were observed, each during three or four days; they were essentially alike, those of the last-named species being the simplest. The petiole of C. floribunda was secured to a stick at the base of the two terminal leaflets, and a filament was fixed along the midrib of one of them. Its movements were traced from 1 P.M. on August 13th to 8.30 A.M.

17th; but those during the last 2 h. are alone given in Fig. 156. From 8

A.M. on each day (by which hour the leaf had assumed its diurnal position) to 2 or 3 P.M., it either zigzagged or circumnutated over nearly the same small space; at between 2 and 3 P.M. the great evening fall commenced. The lines representing this fall and the early morning rise are oblique, owing to the peculiar manner in which the leaflets sleep, as already described.

After the leaflet was asleep at 6 P.M., and whilst the glass filament hung [page 373]

perpendicularly down, the movement of its apex was traced until 10.30 P.M.; and during this whole time it swayed from side to side, completing more than one ellipse.

 

Fig 156. Cassia floribunda: circumnutation and nyctitropic movement of a terminal leaflet (1 5/6 inch in length) traced from 8.30 A.M. to same hour on following morning. Apex of leaflet 5 � inches from the vertical glass.

Main petiole 3 3/4 inches long. Temp. 16o - 17 1/2o C. Figure reduced to one-half of the original scale.

 

Bauhinia (Tribe 15).—The nyctitropic movements of four species were alike, and were highly peculiar. A plant raised from seed sent us from South Brazil by Fritz M�ller, was more especially observed. The leaves are large and deeply notched at their ends. At night the two halves rise up and close completely together, like the opposite leaflets of many Leguminosae. With very young plants the petioles rise considerably at the same time; one, which was inclined at noon 45o above the horizon, at night stood at 75o; it thus rose 30o; another rose 34o. Whilst the two halves of the leaf are closing, the midrib at first sinks vertically downwards and afterwards bends backwards, so as to pass close along one side of its own upwardly inclined petiole; the midrib being thus directed towards the stem or axis of the plant. The angle which the midrib formed with the horizon was measured in one case at different hours: at noon it stood horizontally; late in the evening it depended vertically; then rose to the opposite side, and at 10.15 P.M. stood at only 27o beneath the horizon, being directed towards the stem. It had thus travelled through 153o.

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Owing to this movement—to the leaves being folded—and to the petioles rising, the whole plant is as much more compact at night than during the day, as a fastigiate Lombardy poplar is compared with any other species of poplar. It is remarkable that when our plants had grown a little older, viz., to a height of 2 or 3 feet, the petioles did not rise at night, and the midribs of the folded leaves were no longer bent back along one side of the petiole. We have noticed in some other genera that the petioles of very young plants rise much more at night than do those of older plants.

 

Tamarindus Indica (Tribe 16).—The leaflets approach or meet each other at night, and are all directed towards the apex of the leaf. They thus become imbricated with their midribs parallel to the petiole. The movement is closely similar to that of Haematoxylon (see Fig. 153), but more striking from the greater number of the leaflets.

 

Adenanthera, Prosopis, and Neptunia (Tribe 20).—With Adenanthera pavonia the leaflets turn edgeways and sink at night. In Prosopis they turn upwards. With Neptunia oleracea the leaflets on the opposite sides of the same pinna come into contact at night and are directed forwards. The pinnae themselves move downwards, and at the same time backwards or towards the stem of the plant. The main petiole rises.

 

Mimosa pudica (Tribe 20).—This plant has been the subject of innumerable observations; but there are some points in relation to our subject which have not been sufficiently attended to. At night, as is well known, the opposite leaflets come into contact and point towards the apex of the leaf; they thus become neatly imbricated with their upper surfaces protected. The four pinnae also approach each other closely, and the whole leaf is thus rendered very compact. The main petiole sinks downwards during the day till late in the evening, and rises until very early in the morning. The stem is continually circumnutating at a rapid rate, though not to a wide extent.

Some very young plants, kept in darkness, were observed during two days, and although subjected to a rather low temperature of 57o - 59o F., the stem of one described four small ellipses in the course of 12 h. We shall immediately see that the main petiole is likewise continually circumnutating, as is each separate pinna and each separate leaflet.

Therefore, if the movement of the apex of any one leaflet were to be traced, the course described would be compounded of the movements of four separate parts.

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A filament had been fixed on the previous evening, longitudinally to the main petiole of a nearly full-grown, highly-sensitive leaf (four inches in length), the stem having been secured to a stick at its base; and a tracing was made on a vertical glass in the hot-house under a high temperature. In the figure given (Fig. 157), the first dot was made at 8.30 A.M. August 2nd, and the last at 7 P.M. on the 3rd. During 12 h. on the first day the petiole moved thrice downwards and twice upwards. Within the same length of time on the second day, it moved five times downwards and four times upwards. As the ascending and descending lines do not coincide, the petiole manifestly circumnutates; the great evening fall and nocturnal rise being an exaggeration of one of the circumnutations. It should, however, be observed that the petiole fell much lower down in the evenings than could be seen on the vertical glass or is represented in the diagram. After 7

P.M. on the 3rd (when the last dot in Fig. 157 was made) the pot was carried into a bed-room, and the petiole was found at 12.50 A.M. (i.e.

after midnight) standing almost upright, and much more highly inclined than it was at 10.40 P.M. When observed again at 4 A.M. it had begun to fall, and continued falling till 6.15 A.M., after which hour it zigzagged and again circumnutated. Similar observations were made on another petiole, with nearly the same result.

 

Fig. 157 Mimosa pudica: circumnutation and nyctitropic movement of main petiole, traced during 34 h. 30 m.

 

On two other occasions the movement of the main petiole [page 376]

was observed every two or three minutes, the plants being kept at a rather high temperature, viz., on the first occasion at 77o - 81o F., and the filament then described 2 � ellipses in 69 m. On the second occasion, when the temperature was 81o - 86o F., it made rather more than 3 ellipses in 67

m. therefore, Fig. 157, though now sufficiently complex, would have been incomparably more so, if dots had been made on the glass every 2 or 3

minutes, instead of every hour or half-hour. Although the main petiole is continually and rapidly describing small ellipses during the day, yet after the great nocturnal rising movement has commenced, if dots are made every 2

or 3 minutes, as was done for an hour between 9.30 and 10.30 P.M. (temp.

84o F.), and the dots are then joined, an almost absolutely straight line is the result.

 

To show that the movement of the petiole is in all probability due to the varying turgescence of the pulvinus, and not to growth (in accordance with the conclusions of Pfeffer), a very old leaf, with some of its leaflets yellowish and hardly at all sensitive, was selected for observation, and the plant was kept at the highly favourable temp. of 80o F. The petiole fell from 8 A.M. till 10.15 A.M., it then rose a little in a somewhat zigzag line, often remaining stationary, till 5 P.M., when the great evening fall commenced, which was continued till at least 10 P.M. By 7 A.M.

on the following morning it had risen to the same level as on the previous morning, and then descended in a zigzag line. But from 10.30 A.M. till 4.15

P.M. it remained almost motionless, all power of movement being now lost.

The petiole, therefore, of this very old leaf, which must have long ceased growing, moved periodically; but instead of circumnutating several times during the day, it moved only twice down and twice up in the course of 24

h., with the ascending and descending lines not coincident.

 

It has already been stated that the pinnae move independently of the main petiole.