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

each side. They all sleep by bending vertically downwards, but do not become at all folded. The petiole is rather long, and, one having been secured to a stick, the movement of the terminal leaflet was traced during 45 h. on a vertical glass. It moved in a very simple manner, sinking rapidly after 5 P.M., and rising rapidly early next morning. During the middle of the day it moved slowly and a little laterally. Consequently the ascending and descending lines did not coincide, and a single great ellipse was formed each day. There was no other evidence of circumnutation, and this fact is of interest, as we shall hereafter see.

 

Oxalis sensitiva.—The leaflets, as in the last species, bend vertically down at night, without becoming folded. The much elongated main petiole rises considerably in the evening, but in [page 328]

some very young plants the rise did not commence until late at night. We have seen that the cotyledons, instead of sinking like the leaflets, rise up vertically at night.

 

Oxalis bupleurifolia.—This species is rendered remarkable by the petioles being foliaceous, like the phyllodes of many Acacias. The leaflets are small, of a paler green and more tender consistence than the foliaceous petioles. The leaflet which was observed was .55 inch in length, and was borne by a petiole 2 inches long and .3 inch broad. It may be suspected that the leaflets are on the road to abortion or obliteration, as has actually occurred with those of another Brazilian species, O. rusciformis.

Nevertheless, in the present species the nyctitropic movements are perfectly performed. The foliaceous petiole was first observed during 48

h., and found to be in continued circumnutation, as shown in the accompanying figure (Fig. 130). It rose during the day and early part of the night, and fell during the remainder of the night and early morning; but the movement was not sufficient to be called sleep. The ascending and descending lines did not coincide, so that an ellipse was formed each day.

There was but little zigzagging; if the filament had been fixed longitudinally, we should probably have seen that there was more lateral movement than appears in the diagram.

 

Fig. 130. Oxalis bupleurifolia: circumnutation of foliaceous petiole, filament fixed obliquely across end of petiole; movements traced on vertical glass from 9 A.M. June 26th to 8.50 A.M. 28th. Apex of leaflet 4 �

inches from the glass, so movement not much magnified. Plant 9 inches high, illuminated from above. Temp. 23 1/2o - 24 1/2o C.

 

A terminal leaflet on another leaf was next observed (the petiole being secured), and its movements are shown in Fig. 131. During the day the leaflets are extended horizontally, and at night depend vertically; and as the petiole rises during the day the leaflets have to bend down in the evening

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more than 90o, so as to assume at night their vertical position. On the first day the leaflet simply moved up and down; on the Fig. 131. Oxalis bupleurifolia: circumnutation and nyctitropic movement of terminal leaflet, with filament affixed along the midrib; traced on a vertical glass from 9 A.M. on June 26th to 8.45 A.M. 28th. Conditions the same as in the last case.

 

second day it plainly circumnutated between 8 A.M. and 4.30 P.M., after which hour the great evening fall commenced.

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Averrhoa bilimbi (Oxalidae).—It has long been known,* firstly, that the leaflets in this genus sleep; secondly, that they move spontaneously during the day; and thirdly, that they are sensitive to a touch; but in none of these respects do they differ essentially from the species of Oxalis. They differ, however, as Mr. R. I. Lynch** has lately shown, in their spontaneous movements being strongly marked. In the case of A. bilimbi, it is a wonderful spectacle to behold on a warm sunny day the leaflets one after the other sinking rapidly downwards, and again ascending slowly.

Their movements rival those of Desmodium gyrans. At night the leaflets hang vertically down; and now

 

Fig. 132. Averrhoa bilimbi: leaf asleep; drawing reduced.

 

they are motionless, but this may be due to the opposite ones being pressed together (Fig. 132). The main petiole is in constant movement during the day, but no careful observations were made on it. The following diagrams are graphic representations of the variations in the angle, which a given leaflet makes with the vertical. The observations were made as follows. The plant growing in a pot was kept in a high temperature, the petiole of the leaf to be observed pointing straight at the observer, being separated from him by a vertical pane of glass. The petiole was secured so that the basal joint, or pulvinus, of one of the lateral leaflets was at the centre of a graduated arc placed close behind the leaflet. A fine glass filament was fixed to the leaf, so as to project like a continuation of the * Dr. Bruce, ‘Philosophical Trans.,’ 1785, p. 356.

 

** ‘Journal Linn. Soc.,’ vol. xvi. 1877, p. 231.

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midrib. This filament acted as an index; and as the leaf rose and fell, rotating about its basal joint, its angular movement Fig. 133. Averrhoa bilimbi: angular movements of a leaflet during its evening descent, when going to sleep. Temp. 78o - 81o F.

 

could be recorded by reading off at short intervals of time the position of the glass filament on the graduated arc. In order [page 332]

to avoid errors of parallax, all readings were made by looking through a small ring painted on the vertical glass, in a line with the joint of the leaflet and the centre of the graduated arc. In the following diagrams the ordinates represent the angles which the leaflet made with the vertical at successive instants.* It follows that a fall in the curve represents an actual dropping of the leaf, and that the zero line represents a vertically dependent position. Fig. 133 represents the nature of the movements which occur in the evening, as soon as the leaflets begin to assume their nocturnal position. At 4.55 P.M. the leaflet formed an angle of 85o with the vertical, or was only 5o below the horizontal; but in order that the diagram might get into our page, the leaflet is represented falling from 75o instead of 85o. Shortly after 6 P.M. it hung vertically down, and had attained its nocturnal position. Between 6.10 and 6.35 P.M. it performed a number of minute oscillations of about 2o each, occupying periods of 4 or 5

m. The complete state of rest of the leaflet which ultimately followed is not shown in the diagram. It is manifest that each oscillation consists of a gradual rise, followed by a sudden fall. Each time the leaflet fell, it approached nearer to the nocturnal position than it did on the previous fall. The amplitude of the oscillations diminished, while the periods of oscillation became shorter.

 

In bright sunshine the leaflets assume a highly inclined dependent position. A leaflet in diffused light was observed rising for 25 m. A blind was then pulled up so that the plant was brightly illuminated (BR in Fig.

134), and within a minute it began to fall, and ultimately fell 47o, as shown in the diagram. This descent was performed by six descending steps, precisely similar to those by which the nocturnal fall is effected. The plant was then again shaded (SH), and a long slow rise occurred until another series of falls commenced at BR’, when the sun was again admitted.

In this experiment cool air was allowed to enter by the windows being opened at the same time that the blinds were pulled up, so that in spite of the sun shining on the plant the temperature was not raised.

 

The effect of an increase of temperature in diffused light is * In all the diagrams 1 mm. in the horizontal direction represents one minute of time. Each mm. in the vertical direction represents one degree of angular movement. In Figs. 133 and 134 the temperature is represented (along the ordinates) in the scale of 1 mm. to each 0.1 degree C. In Fig.

135 each mm. equals 0.2o F.

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shown in Fig. 135. The temperature began to rise at 11.35 A.M. (in consequence of the fire being lighted), but by 12.42 a marked fall had occurred. It may be seen in the diagram that when the temperature was highest there were rapid oscillations

 

Fig. 134. Averrhoa bilimbi: angular movements of leaflet during a change from bright illumination to shade; temperature (broken line) remaining nearly the same.

 

of small amplitude, the mean position of the leaflet being at the time nearer the vertical. When the temperature began to fall, the oscillations became slower and larger, and the mean position of the leaf again approached the horizontal. The rate of oscillation was sometimes quicker than is represented in the above diagram. Thus, when the temperature was between 31o and

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Fig. 135. Averrhoa bilimbi: angular movement of leaflet during a change of temperature; light remaining the same. The broken line shows the change of temperature.

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32o C., 14 oscillations of a few degrees occurred in 19 m. On the other hand, an oscillation may be much slower; thus a leaflet was observed (temperature 25o C.) to rise during 40 m. before it fell and completed its oscillation.

 

Fig. 136. Porlieria hygrometrica: circumnutation and nyctitropic movements of petiole of leaf, traced from 9.35 A.M. July 7th to about midnight on the 8th. Apex of leaf 7 � inches from the vertical glass. Temp. 19 1/2o - 20

1/2o C.

 

Porlieria hygrometrica (Zygophylleae).—The leaves of this plant (Chilian form) are from 1 to 1 � inch in length, and bear as many as 16 or 17 small leaflets on each side, which do not stand opposite one another. They are articulated to the petiole, and the petiole to the branch by a pulvinus. We must premise that apparently two forms are confounded under the same name: the leaves on a bush from Chili, which was sent to us from Kew, bore many leaflets, whilst those on plants in the Botanic Garden at W�rzburg bore only 8 or 9 pairs; and the whole character of the bushes appeared somewhat different. We shall also see that they differ in a remarkable physiological peculiarity. On the Chilian plant the petioles of the younger leaves on upright branches, stood horizontally during the day, and at night sank down vertically so as to depend parallel and close to the branch beneath. The petioles of rather older leaves did not become at night vertically depressed, but only highly inclined. In one instance we found a branch which had grown perpendicularly downwards, and the petioles on it moved in the same direction relatively to the branch as just stated, and therefore moved upwards. On horizontal branches the younger petioles likewise move at night in the same direction as before, that is, towards the branch, and are consequently then extended horizontally; but it is remarkable that the older petioles on the

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same branch, though moving a little in the same direction, also bend downwards; they thus occupy a somewhat different position, relatively to the centre of the earth and to the branch, from that of the petioles on the upright branches. With respect to the leaflets, they move at night towards the apex of the petiole until their midribs stand nearly parallel to it; and they then lie neatly imbricated one over the other. Thus half of the upper surface of each leaflet is in close contact with half of the lower surface of the one next in advance; and all the leaflets, excepting the basal ones, have the whole of their upper surfaces and half of their lower surfaces well protected.