The Movements and Habits of Climbing Plants · Charles Darwin
CHAPTER IV. TENDRIL-BEARERS—(_continued_).
_Echinocystis lobata_.—Numerous observations were made on this plant (raised from seed sent me by Prof. Asa Gray), for the spontaneous revolving movements of the internodes and tendrils were first observed by me in this case, and greatly perplexed me. My observations may now be much condensed. I observed thirty-five revolutions of the internodes and tendrils; the slowest rate was 2 hrs. and the average rate, with no great fluctuations, 1 hr. 40 m. Sometimes I tied the internodes, so that the tendrils alone moved; at other times I cut off the tendrils whilst very young, so that the internodes revolved by themselves; but the rate was not thus affected. The course generally pursued was with the sun, but often in an opposite direction. Sometimes the movement during a short time would either stop or be reversed; and this apparently was due to interference from the light, as, for instance, when I placed a plant close to a window. In one instance, an old tendril, which had nearly ceased revolving, moved in one direction, whilst a young tendril above moved in an opposite course. The two uppermost internodes alone revolve; and as soon as the lower one grows old, only its upper part continues to move. The ellipses or circles swept by the summits of the internodes are about three inches in diameter; whilst those swept by the tips of the tendrils, are from 15 to 16 inches in diameter. During the revolving movement, the internodes become successively curved to all points of the compass; in one part of their course they are often inclined, together with the tendrils, at about 45° to the horizon, and in another part stand vertically up. There was something in the appearance of the revolving internodes which continually gave the false impression that their movement was due to the weight of the long and spontaneously revolving tendril; but, on cutting off the latter with sharp scissors, the top of the shoot rose only a little, and went on revolving. This false appearance is apparently due to the internodes and tendrils all curving and moving harmoniously together.
A revolving tendril, though inclined during the greater part of its course at an angle of about 45° (in one case of only 37°) above the horizon, stiffened and straightened itself from tip to base in a certain part of its course, thus becoming nearly or quite vertical. I witnessed this repeatedly; and it occurred both when the supporting internodes were free and when they were tied up; but was perhaps most conspicuous in the latter case, or when the whole shoot happened to be much inclined. The tendril forms a very acute angle with the projecting extremity of the stem or shoot; and the stiffening always occurred as the tendril approached, and had to pass over the shoot in its circular course. If it had not possessed and exercised this curious power, it would infallibly have struck against the extremity of the shoot and been arrested. As soon as the tendril with its three branches begins to stiffen itself in this manner and to rise from an inclined into a vertical position, the revolving motion becomes more rapid; and as soon as the tendril has succeeded in passing over the extremity of the shoot or point of difficulty, its motion, coinciding with that from its weight, often causes it to fall into its previously inclined position so quickly, that the apex could be seen travelling like the minute hand of a gigantic clock.
The tendrils are thin, from 7 to 9 inches in length, with a pair of short lateral branches rising not far from the base. The tip is slightly and permanently curved, so as to act to a limited extent as a hook. The concave side of the tip is highly sensitive to a touch; but not so the convex side, as was likewise observed to be the case with other species of the family by Mohl (p. 65). I repeatedly proved this difference by lightly rubbing four or five times the convex side of one tendril, and only once or twice the concave side of another tendril, and the latter alone curled inwards. In a few hours afterwards, when the tendrils which had been rubbed on the concave side had straightened themselves, I reversed the process of rubbing, and always with the same result. After touching the concave side, the tip becomes sensibly curved in one or two minutes; and subsequently, if the touch has been at all rough, it coils itself into a helix. But the helix will, after a time, straighten itself, and be again ready to act. A loop of thin thread only one-sixteenth of a grain in weight caused a temporary flexure. The lower part was repeatedly rubbed rather roughly, but no curvature ensued; yet this part is sensitive to prolonged pressure, for when it came into contact with a stick, it would slowly wind round it.
One of my plants bore two shoots near together, and the tendrils were repeatedly drawn across one another, but it is a singular fact that they did not once catch each other. It would appear as if they had become habituated to contact of this kind, for the pressure thus caused must have been much greater than that caused by a loop of soft thread weighing only the one-sixteenth of a grain. I have, however, seen several tendrils of _Bryonia dioica_ interlocked, but they subsequently released one another. The tendrils of the Echinocystis are also habituated to drops of water or to rain; for artificial rain made by violently flirting a wet brush over them produced not the least effect. 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.
_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.
Various authors (Palm, p. 55; Mohl, p. 45; Lindley, &c.) believe that the tendrils of the vine are modified flower-peduncles. I here give a drawing (fig. 10) of the ordinary state of a young flower-stalk: it consists of the “common peduncle” (A); of the “flower-tendril” (B), which is represented as having caught a twig; and of the “sub-peduncle” (C) bearing the flower-buds. The whole moves spontaneously, like a true tendril, but in a less degree; the movement, however, is greater when the sub-peduncle (C) does not bear many flower-buds. The common peduncle (A) has not the power of clasping a support, nor has the corresponding part of a true tendril. The flower-tendril (B) is always longer than the sub-peduncle (C) and has a scale at its base; it sometimes bifurcates, and therefore corresponds in every detail with the longer scale-bearing branch (B, fig. 9) of the true tendril. It is, however, inclined backwards from the sub-peduncle (C), or stands at right angles with it, and is thus adapted to aid in carrying the future bunch of grapes. When rubbed, it curves and subsequently straightens itself; and it can, as is shown in the drawing, securely clasp a support. I have seen an object as soft as a young vine-leaf caught by one.
The lower and naked part of the sub-peduncle (C) is likewise slightly sensitive to a rub, and I have seen it bent round a stick and even partly round a leaf with which it had come into contact. That the sub-peduncle has the same nature as the corresponding branch of an ordinary tendril, is well shown when it bears only a few flowers; for in this case it becomes less branched, increases in length, and gains both in sensitiveness and in the power of spontaneous movement. I have twice seen sub-peduncles which bore from thirty to forty flower-buds, and which had become considerably elongated and were completely wound round sticks, exactly like true tendrils. The whole length of another sub-peduncle, bearing only eleven flower-buds, quickly became curved when slightly rubbed; but even this scanty number of flowers rendered the stalk less sensitive than the other branch, that is, the flower-tendril; for the latter after a lighter rub became curved more quickly and in a greater degree. I have seen a sub-peduncle thickly covered with flower-buds, with one of its higher lateral branchlets bearing from some cause only two buds; and this one branchlet had become much elongated and had spontaneously caught hold of an adjoining twig; in fact, it formed a little sub-tendril. The increasing length of the sub-peduncle (C) with the decreasing number of the flower-buds is a good instance of the law of compensation. In accordance with this same principle, the true tendril as a whole is always longer than the flower-stalk; for instance, on the same plant, the longest flower-stalk (measured from the base of the common peduncle to the tip of the flower-tendril) was 8½ inches in length, whilst the longest tendril was nearly double this length, namely 16 inches.
The gradations from the ordinary state of a flower-stalk, as represented in the drawing (fig. 10), to that of a true tendril (fig. 9) are complete. We have seen that the sub-peduncle (C), whilst still bearing from thirty to forty flower-buds, sometimes becomes a little elongated and partially assumes all the characters of the corresponding branch of a true tendril. From this state we can trace every stage till we come to a full-sized perfect tendril, bearing on the branch which corresponds with the sub-peduncle one single flower-bud! Hence there can be no doubt that the tendril is a modified flower-peduncle. Another kind of gradation well deserves notice. Flower-tendrils (B, fig. 10) sometimes produce a few flower-buds. For instance, on a vine growing against my house, there were thirteen and twenty-two flower-buds respectively on two flower-tendrils, which still retained their characteristic qualities of sensitiveness and spontaneous movement, but in a somewhat lessened degree. On vines in hothouses, so many flowers are occasionally produced on the flower-tendrils that a double bunch of grapes is the result; and this is technically called by gardeners a “cluster.” In this state the whole bunch of flowers presents scarcely any resemblance to a tendril; and, judging from the facts already given, it would probably possess little power of clasping a support, or of spontaneous movement. Such flower-stalks closely resemble in structure those borne by _Cissus_. This genus, belonging to the same family of the Vitaceæ, produces well-developed tendrils and ordinary bunches of flowers; but there are no gradations between the two states. If the genus _Vitis_ had been unknown, the boldest believer in the modification of species would never have surmised that the same individual plant, at the same period of growth, would have yielded every possible gradation between ordinary flower-stalks for the support of the flowers and fruit, and tendrils used exclusively for climbing. But the vine clearly gives us such a case; and it seems to me as striking and curious an instance of transition as can well be conceived.
_Cissus discolor_.—The young shoots show no more movement than can be accounted for by daily variations in the action of the light. The tendrils, however, revolve with much regularity, following the sun; and, in the plants observed by me, swept circles of about 5 inches in diameter. Five circles were completed in the following times:—4 hrs. 45 m., 4 hrs. 50 m., 4 hrs. 45 m., 4 hrs. 30 m., and 5 hrs. The same tendril continues to revolve during three or four days. The tendrils are from 3½ to 5 inches in length. They are formed of a long foot-stalk, bearing two short branches, which in old plants again bifurcate. The two branches are not of quite equal length; and as with the vine, the longer one has a scale at its base. The tendril stands vertically upwards; the extremity of the shoot being bent abruptly downwards, and this position is probably of service to the plant by allowing the tendril to revolve freely and vertically.
Both branches of the tendril, whilst young, are highly sensitive. A touch with a pencil, so gentle as only just to move a tendril borne at the end of a long flexible shoot, sufficed to cause it to become perceptibly curved in four or five minutes. It became straight again in rather above one hour. A loop of soft thread weighing one-seventh of a grain (9.25 mg.) was thrice tried, and each time caused the tendril to become curved in 30 or 40 m. Half this weight produced no effect. The long foot-stalk is much less sensitive, for a slight rubbing produced no effect, although prolonged contact with a stick caused it to bend. The two branches are sensitive on all sides, so that they converge if touched on their inner sides, and diverge if touched on their outer sides. If a branch be touched at the same time with equal force on opposite sides, both sides are equally stimulated and there is no movement. Before examining this plant, I had observed only tendrils which are sensitive on one side alone, and these when lightly pressed between the finger and thumb become curved; but on thus pinching many times the tendrils of the _Cissus_ no curvature ensued, and I falsely inferred at first that they were not at all sensitive.
_Cissus antarcticus_.—The tendrils on a young plant were thick and straight, with the tips a little curved. When their concave surfaces were rubbed, and it was necessary to do this with some force, they very slowly became curved, and subsequently straight again. They are therefore much less sensitive than those of the last species; but they made two revolutions, following the sun, rather more rapidly, viz., in 3 hrs. 30 m. and 4 hrs. The internodes do not revolve. _Ampelopsis hederacea_ (_Virginian Creeper_).—The internodes apparently do not move more than can be accounted for by the varying action of the light. The tendrils are from 4 to 5 inches in length, with the main stem sending off several lateral branches, which have their tips curved, as may be seen in the upper figure (fig. 11). They exhibit no true spontaneous revolving movement, but turn, as was long ago observed by Andrew Knight, {145} from the light to the dark. I have seen several tendrils move in less than 24 hours, through an angle of 180° to the dark side of a case in which a plant was placed, but the movement is sometimes much slower. The several lateral branches often move independently of one another, and sometimes irregularly, without any apparent cause. These tendrils are less sensitive to a touch than any others observed by me. By gentle but repeated rubbing with a twig, the lateral branches, but not the main stem, became in the course of three or four hours slightly curved; but they seemed to have hardly any power of again straightening themselves. The tendrils of a plant which had crawled over a large box-tree clasped several of the branches; but I have repeatedly seen that they will withdraw themselves after seizing a stick. When they meet with a flat surface of wood or a wall (and this is evidently what they are adapted for), they turn all their branches towards it, and, spreading them widely apart, bring their hooked tips laterally into contact with it. In effecting this, the several branches, after touching the surface, often rise up, place themselves in a new position, and again come down into contact with it.
Text from Project Gutenberg, public domain in the United States.