I have described the above case in some detail, because, as far as I have seen, it is rare to find any special adaptations with twining plants, in which respect they differ much from the more highly organized tendril-bearers. The Solanum dulcamara, as we shall presently see, can twine only round stems which are both thin and flexible. Most twining plants are adapted to ascend supports of moderate though of different thicknesses. Our English twiners, as far as I have seen, never twine round trees, excepting the honeysuckle (Lonicera periclymenum), which I have observed twining up a young beech-tree nearly 4.5 inches in diameter. Mohl (p. 134) found that the Phaseolus multiflorus and Ipomoea purpurea could not, when placed in a room with the light entering on one side, twine round sticks between 3 and 4 inches in diameter; for this interfered, in a manner presently to be explained, with the revolving movement. In the open air, however, the Phaseolus twined round a support of the above thickness, but failed in twining round one 9 inches in diameter. Nevertheless, some twiners of the warmer temperate regions can manage this latter degree of thickness; for I hear from Dr. Hooker that at Kew the Ruscus androgynus has ascended a column 9 inches in diameter; and although a Wistaria grown by me in a small pot tried in vain for weeks to get round a post between 5 and 6 inches in thickness, yet at Kew a plant ascended a trunk above 6 inches in diameter. The tropical twiners, on the other hand, can ascend thicker trees; I hear from Drs. Thomson and Hooker that this is the case with the Butea parviflora, one of the Menispermaceae, and with some Dalbergias and other Leguminosae. {19} This power would be necessary for any species which had to ascend by twining the large trees of a tropical forest; otherwise they would hardly ever be able to reach the light. In our temperate countries it would be injurious to the twining plants which die down every year if they were enabled to twine round trunks of trees, for they could not grow tall enough in a single season to reach the summit and gain the light.

By what means certain twining plants are adapted to ascend only thin stems, whilst others can twine round thicker ones, I do not know. It appeared to me probable that twining plants with very long revolving shoots would be able to ascend thick supports; accordingly I placed Ceropegia Gardnerii near a post 6 inches in diameter, but the shoots entirely failed to wind round it; their great length and power of movement merely aid them in finding a distant stem round which to twine. The Sphaerostemma marmoratum is a vigorous tropical twiner; and as it is a very slow revolver, I thought that this latter circumstance might help it in ascending a thick support; but though it was able to wind round a 6-inch post, it could do this only on the same level or plane, and did not form a spire and thus ascend.

As ferns differ so much in structure from phanerogamic plants, it may be worth while here to show that twining ferns do not differ in their habits from other twining plants. In Lygodium articulatum the two internodes of the stem (properly the rachis) which are first formed above the root-stock do not move; the third from the ground revolves, but at first very slowly. This species is a slow revolver: but L. scandens made five revolutions, each at the average rate of 5 hrs. 45 m.; and this represents fairly well the usual rate, taking quick and slow movers, amongst phanerogamic plants. The rate was accelerated by increased temperature. At each stage of growth only the two upper internodes revolved. A line painted along the convex surface of a revolving internode becomes first lateral, then concave, then lateral and ultimately again convex. Neither the internodes nor the petioles are irritable when rubbed. The movement is in the usual direction, namely, in opposition to the course of the sun; and when the stem twines round a thin stick, it becomes twisted on its own axis in the same direction. After the young internodes have twined round a stick, their continued growth causes them to slip a little upwards. If the stick be soon removed, they straighten themselves, and recommence revolving. The extremities of the depending shoots turn upwards, and twine on themselves. In all these respects we have complete identity with twining phanerogamic plants; and the above enumeration may serve as a summary of the leading characteristics of all twining plants.

Charles Darwin

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