The revolving movement of a tendril is not stopped by the curving of its extremity after it has been touched. When one of the lateral branches has firmly clasped an object, the middle branch continues to revolve. When a stem is bent down and secured, so that the tendril depends but is left free to move, its previous revolving movement is nearly or quite stopped; but it soon begins to bend upwards, and as soon as it has become horizontal the revolving movement recommences. I tried this four times; the tendril generally rose to a horizontal position in an hour or an hour and a half; but in one case, in which a tendril depended at an angle of 45° beneath the horizon, the uprising took two hours; in half an hour afterwards it rose to 23° above the horizon and then recommenced revolving. This upward movement is independent of the action of light, for it occurred twice in the dark, and on another occasion the light came in on one side alone. The movement no doubt is guided by opposition to the force of gravity, as in the case of the ascent of the plumules of germinating seeds.
A tendril does not long retain its revolving power; and as soon as this is lost, it bends downwards and contracts spirally. After the revolving movement has ceased, the tip still retains for a short time its sensitiveness to contact, but this can be of little or no use to the plant.
Though the tendril is highly flexible, and though the extremity travels, under favourable circumstances, at about the rate of an inch in two minutes and a quarter, yet its sensitiveness to contact is so great that it hardly ever fails to seize a thin stick placed in its path. The following case surprised me much: I placed a thin, smooth, cylindrical stick (and I repeated the experiment seven times) so far from a tendril, that its extremity could only curl half or three-quarters round the stick; but I always found that the tip managed in the course of a few hours to curl twice or even thrice round the stick. I at first thought that this was due to rapid growth on the outside; but by coloured points and measurements I proved that there had been no sensible increase of length within the time. When a stick, flat on one side, was similarly placed, the tip of the tendril could not curl beyond the flat surface, but coiled itself into a helix, which, turning to one side, lay flat on the little flat surface of wood. In one instance a portion of tendril three-quarters of an inch in length was thus dragged on to the flat surface by the coiling in of the helix. But the tendril thus acquires a very insecure hold, and generally after a time slips off. In one case alone the helix subsequently uncoiled itself, and the tip then passed round and clasped the stick. The formation of the helix on the flat side of the stick apparently shows us that the continued striving of the tip to curl itself closely inwards gives the force which drags the tendril round a smooth cylindrical stick. In this latter case, whilst the tendril was slowly and quite insensibly crawling onwards, I observed several times through a lens that the whole surface was not in close contact with the stick; and I can understand the onward progress only by supposing that the movement is slightly undulatory or vermicular, and that the tip alternately straightens itself a little and then again curls inwards. It thus drags itself onwards by an insensibly slow, alternate movement, which may be compared to that of a strong man suspended by the ends of his fingers to a horizontal pole, who works his fingers onwards until he can grasp the pole with the palm of his hand. However this may be, the fact is certain that a tendril which has caught a round stick with its extreme point, can work itself onwards until it has passed twice or even thrice round the stick, and has permanently grasped it.
Hanburya Mexicana.—The young internodes and tendrils of this anomalous member of the family, revolve in the same manner and at about the same rate as those of the Echinocystis. The stem does not twine, but can ascend an upright stick by the aid of its tendrils. The concave tip of the tendril is very sensitive; after it had become rapidly coiled into a ring owing to a single touch, it straightened itself in 50 m. The tendril, when in full action, stands vertically up, with the projecting extremity of the young stem thrown a little on one side, so as to be out of the way; but the tendril bears on the inner side, near its base, a short rigid branch, which projects out at right angles like a spur, with the terminal half bowed a little downwards. Hence, as the main vertical branch revolves, the spur, from its position and rigidity, cannot pass over the extremity of the shoot, in the same curious manner as do the three branches of the tendril of the Echinocystis, namely, by stiffening themselves at the proper point. The spur is therefore pressed laterally against the young stem in one part of the revolving course, and thus the sweep of the lower part of the main branch is much restricted. A nice case of co-adaptation here comes into play: in all the other tendrils observed by me, the several branches become sensitive at the same period: had this been the case with the Hanburya, the inwardly directed, spur-like branch, from being pressed, during the revolving movement, against the projecting end of the shoot, would infallibly have seized it in a useless or injurious manner. But the main branch of the tendril, after revolving for a time in a vertical position, spontaneously bends downwards; and in doing so, raises the spur-like branch, which itself also curves upwards; so that by these combined movements it rises above the projecting end of the shoot, and can now move freely without touching the shoot; and now it first becomes sensitive.
The tips of both branches, when they come into contact with a stick, grasp it like any ordinary tendril. But in the course of a few days, the lower surface swells and becomes developed into a cellular layer, which adapts itself closely to the wood, and firmly adheres to it. This layer is analogous to the adhesive discs formed by the extremities of the tendrils of some species of Bignonia and of Ampelopsis; but in the Hanburya the layer is developed along the terminal inner surface, sometimes for a length of 1¾ inches, and not at the extreme tip. The layer is white, whilst the tendril is green, and near the tip it is sometimes thicker than the tendril itself; it generally spreads a little beyond the sides of the tendril, and is fringed with free elongated cells, which have enlarged globular or retort-shaped heads. This cellular layer apparently secretes some resinous cement; for its adhesion to the wood was not lessened by an immersion of 24 hrs. in alcohol or water, but was quite loosened by a similar immersion in ether or turpentine. After a tendril has once firmly coiled itself round a stick, it is difficult to imagine of what use the adhesive cellular layer can be. Owing to the spiral contraction which soon ensues, the tendrils were never able to remain, excepting in one instance, in contact with a thick post or a nearly flat surface; if they had quickly become attached by means of the adhesive layer, this would evidently have been of service to the plant.
The tendrils of Bryonia dioica, Cucurbita ovifera, and Cucumis sativa are sensitive and revolve. Whether the internodes likewise revolve I did not observe. In Anguria Warscewiczii, the internodes, though thick and stiff, revolve: in this plant the lower surface of the tendril, some time after clasping a stick, produces a coarsely cellular layer or cushion, which adapts itself closely to the wood, like that formed by the tendril of the Hanburya; but it is not in the least adhesive. In Zanonia Indica, which belongs to a different tribe of the family, the forked tendrils and the internodes revolve in periods between 2 hrs. 8 m. and 3 hrs. 35 m., moving against the sun.
Vitaceæ.—In this family and in the two following, namely, the Sapindacæ and Passifloraceæ, the tendrils are modified flower-peduncles; and are therefore axial in their nature. In this respect they differ from all those previously described, with the exception, perhaps, of the Cucurbitaceæ. The homological nature, however, of a tendril seems to make no difference in its action.
Vitis vinifera.—The tendril is thick and of great length; one from a vine growing out of doors and not vigorously, was 16 inches long. It consists of a peduncle (A), bearing two branches which diverge equally from it. One of the branches (B) has a scale at its base; it is always, as far as I have seen, longer than the other and often bifurcates. The branches when rubbed become curved, and subsequently straighten themselves. After a tendril has clasped any object with its extremity, it contracts spirally; but this does not occur (Palm, p. 56) when no object has been seized. The tendrils move spontaneously from side to side; and on a very hot day, one made two elliptical revolutions, at an average rate of 2 hrs. 15 m. During these movements a coloured line, painted along the convex surface, appeared after a time on one side, then on the concave side, then on the opposite side, and lastly again on the convex side. The two branches of the same tendril have independent movements. After a tendril has spontaneously revolved for a time, it bends from the light towards the dark: I do not state this on my own authority, but on that of Mohl and Dutrochet. Mohl (p. 77) says that in a vine planted against a wall the tendrils point towards it, and in a vineyard generally more or less to the north.
The young internodes revolve spontaneously; but the movement is unusually slight. A shoot faced a window, and I traced its course on the glass during two perfectly calm and hot days. On one of these days it described, in the course of ten hours, a spire, representing two and a half ellipses. I also placed a bell-glass over a young Muscat grape in the hot-house, and it made each day three or four very small oval revolutions; the shoot moving less than half an inch from side to side. Had it not made at least three revolutions whilst the sky was uniformly overcast, I should have attributed this slight degree of movement to the varying action of the light. The extremity of the stem is more or less bent downwards, but it never reverses its curvature, as so generally occurs with twining plants.