Summary on the Nature and Action of Tendrils.
With the majority of tendril-bearing plants the young internodes revolve in more or less broad ellipses, like those made by twining plants; but the figures described, when carefully traced, generally form irregular ellipsoidal spires. The rate of revolution varies from one to five hours in different species, and consequently is in some cases more rapid than with any twining plant, and is never so slow as with those many twiners which take more than five hours for each revolution. The direction is variable even in the same individual plant. In Passiflora, the internodes of only one species have the power of revolving. The Vine is the weakest revolver observed by me, apparently exhibiting only a trace of a former power. In the Eccremocarpus the movement is interrupted by many long pauses. Very few tendril-bearing plants can spirally twine up an upright stick. Although the power of twining has generally been lost, either from the stiffness or shortness of the internodes, from the size of the leaves, or from some other unknown cause, the revolving movement of the stem serves to bring the tendrils into contact with surrounding objects.
The tendrils themselves also spontaneously revolve. The movement begins whilst the tendril is young, and is at first slow. The mature tendrils of Bignonia littoralis move much slower than the internodes. Generally, the internodes and tendrils revolve together at the same rate; in Cissus, Cobæa, and most Passifloræ, the tendrils alone revolve; in other cases, as with Lathyrus aphaca, only the internodes move, carrying with them the motionless tendrils; and, lastly (and this is the fourth possible case), neither internodes nor tendrils spontaneously revolve, as with Lathyrus grandiflorus and Ampelopsis. In most Bignonias, Eccremocarpus Mutisia, and the Fumariaceæ, the internodes, petioles and tendrils all move harmoniously together. In every case the conditions of life must be favourable in order that the different parts should act in a perfect manner.
Tendrils revolve by the curvature of their whole length, excepting the sensitive extremity and the base, which parts do not move, or move but little. The movement is of the same nature as that of the revolving internodes, and, from the observations of Sachs and H. de Vries, no doubt is due to the same cause, namely, the rapid growth of a longitudinal band, which travels round the tendril and successively bows each part to the opposite side. Hence, if a line be painted along that surface which happens at the time to be convex, the line becomes first lateral, then concave, then lateral, and ultimately again convex. This experiment can be tried only on the thicker tendrils, which are not affected by a thin crust of dried paint. The extremities are often slightly curved or hooked, and the curvature of this part is never reversed; in this respect they differ from the extremities of twining shoots, which not only reverse their curvature, or at least become periodically straight, but curve themselves in a greater degree than the lower part. In most other respects a tendril acts as if it were one of several revolving internodes, which all move together by successively bending to each point of the compass. There is, however, in many cases this unimportant difference, that the curving tendril is separated from the curving internode by a rigid petiole. With most tendril-bearers the summit of the stem or shoot projects above the point from which the tendril arises; and it is generally bent to one side, so as to be out of the way of the revolutions swept by the tendril. In those plants in which the terminal shoot is not sufficiently out of the way, as we have seen with the Echinocystis, as soon as the tendril comes in its revolving course to this point, it stiffens and straightens itself, and thus rising vertically up passes over the obstacle in an admirable manner.
All tendrils are sensitive, but in various degrees, to contact with an object, and curve towards the touched side. With several plants a single touch, so slight as only just to move the highly flexible tendril, is enough to induce curvature. Passiflora gracilis possesses the most sensitive tendrils which I have observed: a bit of platina wire 0.02 of a grain (1.23 mg.) in weight, gently placed on the concave point, caused a tendril to become hooked, as did a loop of soft, thin cotton thread weighing one thirty-second of a grain (2.02 mg.) With the tendrils of several other plants, loops weighing one sixteenth of a grain (4.05 mg.) sufficed. The point of a tendril of Passiflora gracilis began to move distinctly in 25 seconds after a touch, and in many cases after 30 seconds. Asa Gray also saw movement in the tendrils of the Cucurbitaceous genus, Sicyos, in 30 seconds. The tendrils of some other plants, when lightly rubbed, moved in a few minutes; with Dicentra in half-an-hour; with Smilax in an hour and a quarter or half; and with Ampelopsis still more slowly. The curling movement consequent on a single touch continues to increase for a considerable time, then ceases; after a few hours the tendril uncurls itself, and is again ready to act. When the tendrils of several kinds of plants were caused to bend by extremely light weights suspended on them, they seemed to grow accustomed to so slight a stimulus, and straightened themselves, as if the loops had been removed. It makes no difference what sort of object a tendril touches, with the remarkable exception of other tendrils and drops of water, as was observed with the extremely sensitive-tendrils of Passiflora gracilis and of the Echinocystis. I have, however, seen tendrils of the Bryony which had temporarily caught other tendrils, and often in the case of the vine.
Tendrils of which the extremities are permanently and slightly curved, are sensitive only on the concave surface; other tendrils, such as those of the Cobæa (though furnished with horny hooks directed to one side) and those of Cissus discolor, are sensitive on all sides. Hence the tendrils of this latter plant, when stimulated by a touch of equal force on opposite sides, did not bend. The inferior and lateral surfaces of the tendrils of Mutisia are sensitive, but not the upper surface. With branched tendrils, the several branches act alike; but in the Hanburya the lateral spur-like branch does not acquire (for excellent reasons which have been explained) its sensitiveness nearly so soon as the main branch. With most tendrils the lower or basal part is either not at all sensitive, or sensitive only to prolonged contact. We thus see that the sensitiveness of tendrils is a special and localized capacity. It is quite independent of the power of spontaneously revolving; for the curling of the terminal portion from touch does not in the least interrupt the former movement. In Bignonia unguis and its close allies, the petioles of the leaves, as well as the tendrils, are sensitive to a touch.
Twining plants when they come into contact with a stick, curl round it invariably in the direction of their revolving movement; but tendrils curl indifferently to either side, in accordance with the position of the stick and the side which is first touched. The clasping movement of the extremity is apparently not steady, but undulatory or vermicular in its nature, as may be inferred from the curious manner in which the tendrils of the Echinocystis slowly crawled round a smooth stick.
As with a few exceptions tendrils spontaneously revolve, it may be asked,—why have they been endowed with sensitiveness?—why, when they come into contact with a stick, do they not, like twining plants, spirally wind round it? One reason may be that they are in most cases so flexible and thin, that when brought into contact with any object, they would almost certainly yield and be dragged onwards by the revolving movement. Moreover, the sensitive extremities have no revolving power as far as I have observed, and could not by this means curl round a support. With twining plants, on the other hand, the extremity spontaneously bends more than any other part; and this is of high importance for the ascent of the plant, as may be seen on a windy day. It is, however, possible that the slow movement of the basal and stiffer parts of certain tendrils, which wind round sticks placed in their path, may be analogous to that of twining plants. But I hardly attended sufficiently to this point, and it would have been difficult to distinguish between a movement due to extremely dull irritability, from the arrestment of the lower part, whilst the upper part continued to move onwards.
Tendrils which are only three-fourths grown, and perhaps even at an earlier age, but not whilst extremely young, have the power of revolving and of grasping any object which they touch. These two capacities are generally acquired at about the same period, and both fail when the tendril is full grown. But in Cobæa and Passiflora punctata the tendrils begin to revolve in a useless manner, before they have become sensitive. In the Echinocystis they retain their sensitiveness for some time after they have ceased to revolve and after they have sunk downwards; in this position, even if they were able to seize an object, such power would be of no service in supporting the stem. It is a rare circumstance thus to detect any superfluity or imperfection in the action of tendrils—organs which are so excellently adapted for the functions which they have to perform; but we see that they are not always perfect, and it would be rash to assume that any existing tendril has reached the utmost limit of perfection.
Some tendrils have their revolving motion accelerated or retarded, in moving to or from the light; others, as with the Pea, seem indifferent to its action; others move steadily from the light to the dark, and this aids them in an important manner in finding a support. For instance, the tendrils of Bignonia capreolata bend from the light to the dark as truly as a wind-vane from the wind. In the Eccremocarpus the extremities alone twist and turn about so as to bring their finer branches and hooks into close contact with any dark surface, or into crevices and holes.