The shoot pointed away from the observer; and a glass filament pointing towards the vertical glass on which the tracing was made, was fixed to the convex surface of the curved portion. Therefore the descending lines in the figure represent the straightening of the curved portion as it grew older. The tracing (Fig. 123, p. 274) was begun at 9 A.M. on July 10th; the filament at first moved but little in a zigzag line, but at 2 P.M. it began rising and continued to do so till 9 P.M.; and this proves that the terminal portion was being more bent downwards. After 9 P.M. on the 10th an opposite movement commenced, and the curved portion began to straighten itself, and this continued till 11.10 A.M. on the 12th, but was interrupted by some small oscillations and zigzags, showing movement in different directions. After 11.10 A.M. on the 12th this part of the stem, still considerably curved, circumnutated in a conspicuous manner until nearly 3 P.M. on the 13th; but during all this time a downward movement of the filament prevailed, caused by the continued straightening of the stem. By the afternoon of the 13th, the summit, which had originally been deflected more than a right angle from the perpendicular, had grown so nearly straight that the tracing could no longer be continued on the vertical glass. There can therefore be no doubt that the straightening of the abruptly curved portion of the growing stem of this plant, which appears to be wholly due to hyponasty, is the result of modified circumnutation. We will only add that a filament was fixed in a different manner across the curved summit of another plant, and the same general kind of movement was observed.

Trifolium repens.--In many, but not in all the species of Trifolium, as the separate little flowers wither, the sub-peduncles bend downwards, so as to depend parallel to the upper part of the main peduncle. In Tr. subterraneum the main peduncle curves downwards for the sake of burying its capsules, and in this species the sub-peduncles of the separate flowers bend

* 'Ueber Orthotrope und Plagiotrope Pflanzentheile;' 'Arbeiten des Bot. Inst., in Würzburg,' Heft ii. 1879, p. 226. [page 277]

Fig. 124. Trifolium repens: circumnutating and epinastic movements of the sub-peduncle of a single flower, traced on a vertical glass under a skylight, in A from 11.30 A.M. Aug. 27th to 7 A.M. 30th; in B from 7 A.M. Aug. 30th to a little after 6 P.M. Sept. 8th. [page 278]

upwards, so as to occupy the same position relatively to the upper part of the main peduncle as in Tr. repens. This fact alone would render it probable that the movements of the sub-peduncles in Tr. repens were independent of geotropism. Nevertheless, to make sure, some flower-heads were tied to little sticks upside down and others in a horizontal position; their sub-peduncles, however, all quickly curved upwards through the action of heliotropism. We therefore protected some flower-heads, similarly secured to sticks, from the light, and although some of them rotted, many of their sub-peduncles turned very slowly from their reversed or from their horizontal positions, so as to stand in the normal manner parallel to the upper part of the main peduncle. These facts show that the movement is independent of geotropism or apheliotropism; it must there[fore] be attributed to epinasty, which however is checked, at least as long as the flowers are young, by heliotropism. Most of the above flowers were never fertilised owing to the exclusion of bees; they consequently withered very slowly, and the movements of the sub-peduncles were in like manner much retarded.

To ascertain the nature of the movement of the sub-peduncle, whilst bending downwards, a filament was fixed across the summit of the calyx of a not fully expanded and almost upright flower, nearly in the centre of the head. The main peduncle was secured to a stick close beneath the head. In order to see the marks on the glass filament, a few flowers had to be cut away on the lower side of the head. The flower under observation at first diverged a little from its upright position, so as to occupy the open space caused by the removal of the adjoining flowers.

Charles Darwin

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