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

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Pinus pinaster (Coniferae).—A young hypocotyl, with the tips of the cotyledons still enclosed within the seed-coats, was at first only .35 of an inch in height; but the upper part grew so rapidly that at the end of our observations it was .6 in height,

Fig. 43. Pinus pinaster: circumnutation of hypocotyl, with filament fixed across its summit, traced on horizontal glass, from 10 A.M. March 21st to 9

A.M. 23rd. Seedling kept in darkness. Movement of bead magnified about 35

times.

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and by this time the filament was attached some way down the little stem.

From some unknown cause, the hypocotyl moved far towards the left, but there could be no doubt (Fig. 43) that it circumnutated. Another hypocotyl was similarly observed, and it likewise moved in a strongly zigzag line to the same side. This lateral movement was not caused by the attachment of the glass filaments, nor by the action of light; for no light was allowed to enter when each observation was made, except from vertically above.

The hypocotyl of a seedling was secured to a little stick; it bore nine in appearance distinct cotyledons, arranged in a circle. The movements of two nearly opposite ones were observed. The tip of one was painted white, with a mark placed below, and the figure described (Fig. 44, A) shows that it made an irregular

Fig. 44. Pinus pinaster: circumnutation of two opposite cotyledons, traced on horizontal glass in darkness, from 8.45 A.M. to 8.35 P.M. Nov. 25th.

Movement of tip in A magnified about 22 times, here reduced to one-half of original scale.

circle in the course of about 8 h. during the night it travelled to a considerable distance in the direction indicated by the broken line. A glass filament was attached longitudinally to the other cotyledon, and this nearly completed (Fig, 44, B) an irregular circular figure in about 12

hours. During the night it also moved to a considerable distance, in the direction indicated by the broken line. The cotyledons therefore circumnutate independently of the movement of the hypocotyl. Although they moved much during the night, they did not approach each other so as to stand more vertically than during the day.

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Cycas pectinata (Cycadeae).—The large seeds of this plant in germinating first protrude a single leaf, which breaks through the ground with the petiole bowed into an arch and with the leaflets involuted. A leaf in this condition, which at the close of our observations was 2 � inches in height, had its movements traced in a warm greenhouse by means of a glass filament bearing paper triangles attached across its tip. The tracing (Fig. 45) shows how large, complex, and rapid were the circum-Fig. 45. Cycas pectinata: circumnutation of young leaf whilst emerging from the ground, feebly illuminated from above, traced on vertical glass, from 5

P.M. May 28th to 11 A.M. 31st. Movement magnified 7 times, here reduced to two-thirds of original scale.

nutating movements. The extreme distance from side to side which it passed over amounted to between .6 and .7 of an inch.

Canna Warscewiczii (Cannaceae).—A seedling with the plumule projecting one inch above the ground was observed, but not under fair conditions, as it was brought out of the hot-house and kept in a room not sufficiently warm.

Nevertheless the tracing (Fig. 46) shows that it made two or three incomplete irregular circles or ellipses in the course of 48 hours. The plumule is straight; and this was the first instance observed [page 59]

by us of the part that first breaks through the ground not being arched.

Fig. 46. Canna Warscewiczii: circumnutation of plumule with filament affixed obliquely to outer sheath-like leaf, traced in darkness on horizontal glass from 8.45 A.M. Nov. 9th to 8.10 A.M. 11th. Movement of bead magnified 6 times.

Allium cepa (Liliaceae).—The narrow green leaf, which protrudes from the seed of the common onion as a cotyledon,* breaks through the ground in the form of an arch, in the same manner as the hypocotyl or epicotyl of a dicotyledonous plant. Long after the arch has risen above the surface the apex remains within the seed-coats, evidently absorbing the still abundant contents. The summit or crown of the arch, when it first protrudes from the seed and is still buried beneath the ground, is simply rounded; but before it reaches the surface it is developed into a conical protuberance of a white colour (owing to the absence of chlorophyll), whilst the adjoining parts are green, with the epidermis apparently rather thicker and tougher than elsewhere. We may therefore conclude that this conical protuberance is a special adaptation for breaking through the ground,** and answers the same end as the knife-like white crest on the summit of the straight cotyledon of the Gramineae.

* This is the expression used by Sachs in his ‘Text-book of Botany.’

** Haberlandt has briefly described (‘Die Schutzeinrichtungen…Keimpflanze,’ 1877, p. 77) this curious structure and the purpose which it subserves. He states that good figures of the cotyledon of the onion have been given by Tittmann and by Sachs in his ‘Experimental Physiologie,’ p. 93.

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After a time the apex is drawn out of the empty seed-coats, and rises up, forming a right angle, or more commonly a still larger angle with the lower part, and occasionally the whole becomes nearly straight. The conical protuberance, which originally formed the crown of the arch, is now seated on one side, and appears like a joint or knee, which from acquiring chlorophyll becomes green, and increases in size. In rarely or never becoming perfectly straight, these cotyledons differ remarkably from the ultimate condition of the arched hypocotyls or epicotyls of dicotyledons.

It is, also, a singular circumstance that the attenuated extremity of the upper bent portion invariably withers and dies.

A filament, 1.7 inch in length, was affixed nearly upright beneath the knee to the basal and vertical portion of a cotyledon; and its movements were traced during 14 h. in the usual manner. The tracing here given (Fig. 47) indicates circumnutation. The movement of the upper part above the knee of the same cotyledon, which projected at about an angle of 45o above the horizon, was observed at the same time. A filament was not affixed to it, but a mark was placed beneath the apex, which was almost white from beginning to wither, and its movements were thus traced. The figure described resembled pretty closely that above given; and this shows that the chief seat of movement is in the lower or basal part of the cotyledon.

Fig. 47. Allium cepa: circumnutation of basal half of arched cotyledon, traced in darkness on horizontal glass, from 8.15 A.M. to 10 P.M. Oct.

31st. Movement of bead magnified about 17 times.

Asparagus officinalis (Asparageae).—The tip of a straight plumule or cotyledon (for we do not know which it should be called) was found at a depth of .1 inch beneath the surface, and the earth was then removed all round to the dept of .3 inch. a glass filament was affixed obliquely to it, and the movement of the bead, magnified 17 times, was traced in darkness.

During the first 1 h. 15 m. the plumule moved to the right, and during the next two hours it returned in a roughly parallel but strongly zigzag course. From some unknown cause it had grown up through the soil in an inclined direction, and now through apogeotropism it moved during nearly 24

h. in

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the same general direction, but in a slightly zigzag manner, until it became upright. On the following morning it changed its course completely.

There can therefore hardly be a doubt that the plumule circumnutates, whilst buried beneath the ground, as much as the pressure of the surrounding earth will permit. The surface of the soil in the pot was now covered with a thin layer of very fine argillaceous sand, which was kept damp; and after the tapering seedlings had grown a few tenths of an inch in height, each was found surrounded by a little open space or circular crack; and this could be accounted for only by their having circumnutated and thus pushed away the sand on all sides; for there was no vestige of a crack in any other part.

In order to prove that there was circumnutation, the move-Fig. 48. Asparagus officinalis: circumnutation of plumules with tips whitened and marks placed beneath, traced on a horizontal glass. A, young plumule; movement traced from 8.30 A.M. Nov. 30th to 7.15 A.M. next morning; magnified about 35 times. B, older plumule; movement traced from 10.15 A.M. to 8.10 P.M. Nov. 29th; magnified 9 times, but here reduced to one-half of original scale.

ments of five seedlings, varying in height from .3 inch to 2 inches, were traced. They were placed within a box and illuminated from above; but in all five cases the longer axes of the figures described were directed to nearly the same point; so that more light seemed to have come through the glass roof of the greenhouse on one side than on any other. All five tracings resembled each other to a certain extent, and it will suffice to give two of them. In A (Fig. 48) the seedling was only .45 of an [page 62]

inch in height, and consisted of a single internode bearing a bud on its summit. The apex described between 8.30 A.M. and 10.20 P.M. (i.e. during nearly 14 hours) a figure which would probably have consisted of 3 �

ellipses, had not the stem been drawn to one side until 1 P.M., after which hour it moved backwards. On the following morning it was not far distant from the point whence it had first started. The actual amount of movement of the apex from side to side was very small, viz. about 1/18th of an inch.

The seedling of which the movements are shown in Fig. 48, B, was 1 3/4 inch in height, and consisted of three internodes besides the bud on the summit.

The figure, which was described during 10 h., apparently represents two irregular and unequal ellipses or circles. The actual amount of movement of the apex, in the line not influenced by the light, was .11 of an inch, and in that thus influenced .37 of an inch. With a seedling 2 inches in height it was obvious, even without the aid of any tracing, that the uppermost part of the stem bent successively to all points of the compass, like the stem of a twining plant. A little increase in the power of circumnutating and in the flexibility of the stem, would convert the common asparagus into a twining plant, as has occurred with one species in this genus, namely, A.

scandens.

Phalaris Canariensis (Gramineae).—With the Gramineae the part which first rises above the ground has been called by some authors the pileole; and various views have been expressed on its homological nature. It is considered by some great authorities to be a cotyledon, which term we will use without venturing to express any opinion on the subject.* It consists in the present case of a slightly flattened reddish sheath, terminating upwards in a sharp white edge; it encloses a true green leaf, which protrudes from the sheath through a slit-like orifice, close beneath and at right angles to the sharp edge on the summit. The sheath is not arched when it breaks through the ground.

The movements of three rather old seedlings, about 1 � inch in height, shortly before the protrusion of the leaves, were first traced. They were illuminated exclusively from above; for, as will hereafter be shown, they are excessively sensitive to the

* We are indebted to the Rev. G. Henslow for an abstract