The Movements and Habits of Climbing Plants, by Charles Darwin

Chapter III. — Tendril-Bearers.

Nature of tendrils — BIGNONIACEAE, various species of, and their different modes of climbing — Tendrils which avoid the light and creep into crevices — Development of adhesive discs — Excellent adaptations for seizing different kinds of supports. — POLEMONIACEAE— Cobaea scandens much branched and hooked tendrils, their manner of action — LEGUMINOSAE— COMPOSITAE— SMILACEAE— Smilax aspera, its inefficient tendrils — FUMARIACEAE— Corydalis claviculata, its state intermediate between that of a leaf-climber and a tendril-bearer.

By tendrils I mean filamentary organs, sensitive to contact and used exclusively for climbing. By this definition, spines, hooks and rootlets, all of which are used for climbing, are excluded. True tendrils are formed by the modification of leaves with their petioles, of flower-peduncles, branches, 24 and perhaps stipules. Mohl, who includes under the name of tendrils various organs having a similar external appearance, classes them according to their homological nature, as being modified leaves, flower-peduncles, &c. This would be an excellent scheme; but I observe that botanists are by no means unanimous on the homological nature of certain tendrils. Consequently I will describe tendril-bearing plants by natural families, following Lindley’s classification; and this will in most cases keep those of the same nature together. The species to be described belong to ten families, and will be given in the following order:— Bignoniaceae, Polemoniaceae, Leguminosae, Compositae, Smilaceae, Fumariaceae, Cucurbitaceae, Vitaceae, Sapindaceae, Passifloraceae. 25

BIGNONIACEAE. — This family contains many tendril-bearers, some twiners, and some root-climbers. The tendrils always consist of modified leaves. Nine species of Bignonia, selected by hazard, are here described, in order to show what diversity of structure and action there may be within the same genus, and to show what remarkable powers some tendrils possess. The species, taken together, afford connecting links between twiners, leaf-climbers, tendril-bearers, and root-climbers.

Bignonia (an unnamed species from Kew, closely allied to B. unguis, but with smaller and rather broader leaves). — A young shoot from a cut-down plant made three revolutions against the sun, at an average rate of 2 hrs. 6m. The stem is thin and flexible; it twined round a slender vertical stick, ascending from left to right, as perfectly and as regularly as any true twining-plant. When thus ascending, it makes no use of its tendrils or petioles; but when it twined round a rather thick stick, and its petioles were brought into contact with it, these curved round the stick, showing that they have some degree of irritability. The petioles also exhibit a slight degree of spontaneous movement; for in one case they certainly described minute, irregular, vertical ellipses. The tendrils apparently curve themselves spontaneously to the same side with the petioles; but from various causes, it was difficult to observe the movement of either the tendrils or petioles, in this and the two following species. The tendrils are so closely similar in all respects to those of B. unguis, that one description will suffice.

Bignonia unguis. — The young shoots revolve, but less regularly and less quickly than those of the last species. The stem twines imperfectly round a vertical stick, sometimes reversing its direction, in the same manner as described in so many leaf-climbers; and this plant though possessing tendrils, climbs to a certain extent like a leaf-climber. Each leaf consists of a petiole bearing a pair of leaflets, and terminates in a tendril, which is formed by the modification of three leaflets, and closely resembles that above figured (fig. 5). But it is a little larger, and in a young plant was about half an inch in length. It is curiously like the leg and foot of a small bird, with the hind toe cut off. The straight leg or tarsus is longer than the three toes, which are of equal length, and diverging, lie in the same plane. The toes terminate in sharp, hard claws, much curved downwards, like those on a bird’s foot. The petiole of the leaf is sensitive to contact; even a small loop of thread suspended for two days caused it to bend upwards; but the sub-petioles of the two lateral leaflets are not sensitive. The whole tendril, namely, the tarsus and the three toes, are likewise sensitive to contact, especially on their under surfaces. When a shoot grows in the midst of thin branches, the tendrils are soon brought by the revolving movement of the internodes into contact with them; and then one toe of the tendril or more, commonly all three, bend, and after several hours seize fast hold of the twigs, like a bird when perched. If the tarsus of the tendril comes into contact with a twig, it goes on slowly bending, until the whole foot is carried quite round, and the toes pass on each side of the tarsus and seize it. In like manner, if the petiole comes into contact with a twig, it bends round, carrying the tendril, which then seizes its own petiole or that of the opposite leaf. The petioles move spontaneously, and thus, when a shoot attempts to twine round an upright stick, those on both sides after a time come into contact with it, and are excited to bend. Ultimately the two petioles clasp the stick in opposite directions, and the foot-like tendrils, seizing on each other or on their own petioles, fasten the stem to the support with surprising security. The tendrils are thus brought into action, if the stem twines round a thin vertical stick; and in this respect the present species differs from the last. Both species use their tendrils in the same manner when passing through a thicket. This plant is one of the most efficient climbers which I have observed; and it probably could ascend a polished stem incessantly tossed by heavy storms. To show how important vigorous health is for the action of all the parts, I may mention that when I first examined a plant which was growing moderately well, though not vigorously, I concluded that the tendrils acted only like the hooks on a bramble, and that it was the most feeble and inefficient of all climbers!

Bignonia Tweedyana. — This species is closely allied to the last, and behaves in the same manner; but perhaps twines rather better round a vertical stick. On the same plant, one branch twined in one direction and another in an opposite direction. The internodes in one case made two circles, each in 2 hrs. 33 m. I was enabled to observe the spontaneous movements of the petioles better in this than in the two preceding species: one petiole described three small vertical ellipses in the course of 11 hrs., whilst another moved in an irregular spire. Some little time after a stem has twined round an upright stick, and is securely fastened to it by the clasping petioles and tendrils, it emits aerial roots from the bases of its leaves; and these roots curve partly round and adhere to the stick. This species of Bignonia, therefore, combines four different methods of climbing generally characteristic of distinct plants, namely, twining, leaf-climbing, tendril-climbing, and root-climbing.

In the three foregoing species, when the foot-like tendril has caught an object, it continues to grow and thicken, and ultimately becomes wonderfully strong, in the same manner as the petioles of leaf-climbers. If the tendril catches nothing, it first slowly bends downwards, and then its power of clasping is lost. Very soon afterwards it disarticulates itself from the petiole, and drops off like a leaf in autumn. I have seen this process of disarticulation in no other tendrils, for these, when they fail to catch an object, merely wither away.

Bignonia venusta. — The tendrils differ considerably from those of the previous species. The lower part, or tarsus, is four times as long as the three toes; these are of equal length and diverge equally, but do not lie in the same plane; their tips are bluntly hooked, and the whole tendril makes an excellent grapnel. The tarsus is sensitive on all sides; but the three toes are sensitive only on their outer surfaces. The sensitiveness is not much developed; for a slight rubbing with a twig did not cause the tarsus or the toes to become curved until an hour had elapsed, and then only in a slight degree. Subsequently they straightened themselves. Both the tarsus and toes can seize well hold of sticks. If the stem is secured, the tendrils are seen spontaneously to sweep large ellipses; the two opposite tendrils moving independently of one another. I have no doubt, from the analogy of the two following allied species, that the petioles also move spontaneously; but they are not irritable like those of B. unguis and B. Tweedyana. The young internodes sweep large circles, one being completed in 2 hrs. 15 m., and a second in 2 hrs. 55 m. By these combined movements of the internodes, petioles, and grapnel-like tendrils, the latter are soon brought into contact with surrounding objects. When a shoot stands near an upright stick, it twines regularly and spirally round it. As it ascends, it seizes the stick with one of its tendrils, and, if the stick be thin, the right-and left-hand tendrils are alternately used. This alternation follows from the stem necessarily taking one twist round its own axis for each completed circle.

The tendrils contract spirally a short time after catching any object; those which catch nothing merely bend slowly downwards. But the whole subject of the spiral contraction of tendrils will be discussed after all the tendril-bearing species have been described.

Bignonia littoralis. — The young internodes revolve in large ellipses. An internode bearing immature tendrils made two revolutions, each in 3 hrs. 50 m.; but when grown older with the tendrils mature, it made two ellipses, each at the rate of 2 hrs. 44 m. This species, unlike the preceding, is incapable of twining round a stick: this does not appear to be due to any want of flexibility in the internodes or to the action of the tendrils, and certainly not to any want of the revolving power; nor can I account for the fact. Nevertheless the plant readily ascends a thin upright stick by seizing a point above with its two opposite tendrils, which then contract spirally. If the tendrils seize nothing, they do not become spiral.

The species last described, ascended a vertical stick by twining spirally and by seizing it alternately with its opposite tendrils, like a sailor pulling himself up a rope, hand over hand; the present species pulls itself up, like a sailor seizing with both hands together a rope above his head.

The tendrils are similar in structure to those of the last species. They continue growing for some time, even after they have clasped an object. When fully grown, though borne by a young plant, they are 9 inches in length. The three divergent toes are shorter relatively to the tarsus than in the former species; they are blunt at their tips and but slightly hooked; they are not quite equal in length, the middle one being rather longer than the others. Their outer surfaces are highly sensitive; for when lightly rubbed with a twig, they became perceptibly curved in 4 m. and greatly curved in 7 m. In 7 hrs. they became straight again and were ready to re-act. The tarsus, for the space of one inch close to the toes, is sensitive, but in a rather less degree than the toes; for the latter after a slight rubbing, became curved in about half the time. Even the middle part of the tarsus is sensitive to prolonged contact, as soon as the tendril has arrived at maturity. After it has grown old, the sensitiveness is confined to the toes, and these are only able to curl very slowly round a stick. A tendril is perfectly ready to act, as soon as the three toes have diverged, and at this period their outer surfaces first become irritable. The irritability spreads but little from one part when excited to another: thus, when a stick was caught by the part immediately beneath the three toes, these seldom clasped it, but remained sticking straight out.

The tendrils revolve spontaneously. The movement begins before the tendril is converted into a three-pronged grapnel by the divergence of the toes, and before any part has become sensitive; so that the revolving movement is useless at this early period. The movement is, also, now slow, two ellipses being completed conjointly in 24 hrs. 18 m. A mature tendril made an ellipse in 6 hrs.; so that it moved much more slowly than the internodes. The ellipses which were swept, both in a vertical and horizontal plane, were of large size. The petioles are not in the least sensitive, but revolve like the tendrils. We thus see that the young internodes, the petioles, and the tendrils all continue revolving together, but at different rates. The movements of the tendrils which rise opposite one another are quite independent. Hence, when the whole shoot is allowed freely to revolve, nothing can be more intricate than the course followed by the extremity of each tendril. A wide space is thus irregularly searched for some object to be grasped.

One other curious point remains to be mentioned. In the course of a few days after the toes have closely clasped a stick, their blunt extremities become developed, though not invariably, into irregular disc-like balls which have the power of adhering firmly to the wood. As similar cellular outgrowths will be fully described under B. capreolata, I will here say nothing more about them.

Bignonia aequinoctialis, var. Chamberlaynii. — The internodes, the elongated non-sensitive petioles, and the tendrils all revolve. The stem does not twine, but ascends a vertical stick in the same manner as the last species. The tendrils also resemble those of the last species, but are shorter; the three toes are more unequal in length, the two outer ones being about one-third shorter and rather thinner than the middle toe; but they vary in this respect. They terminate in small hard points; and what is important, cellular adhesive discs are not developed. The reduced size of two of the toes as well as their lessened sensitiveness, seem to indicate a tendency to abortion; and on one of my plants the first-formed tendrils were sometimes simple, that is, were not divided into three toes. We are thus naturally led to the three following species with undivided tendrils

Bignonia speciosa. — The young shoots revolve irregularly, making narrow ellipses, spires or circles, at rates varying from 3 hrs. 30 m. to 4 hrs. 40 m.; but they show no tendency to twine. Whilst the plant is young and does not require a support, tendrils are not developed. Those borne by a moderately young plant were five inches in length. They revolve spontaneously, as do the short and non-sensitive petioles. When rubbed, they slowly bend to the rubbed side and subsequently straighten themselves; but they are not highly sensitive. There is something strange in their behaviour: I repeatedly placed close to them, thick and thin, rough and smooth sticks and posts, as well as string suspended vertically, but none of these objects were well seized. After clasping an upright stick, they repeatedly loosed it again, and often would not seize it at all, or their extremities did not coil closely round. I have observed hundreds of tendrils belonging to various Cucurbitaceous, Passifloraceous, and Leguminous plants, and never saw one behave in this manner. When, however, my plant had grown to a height of eight or nine feet, the tendrils acted much better. They now seized a thin, upright stick horizontally, that is, at a point on their own level, and not some way up the stick as in the case of all the previous species. Nevertheless, the non-twining stem was enabled by this means to ascend the stick.

The extremity of the tendril is almost straight and sharp. The whole terminal portion exhibits a singular habit, which in an animal would be called an instinct; for it continually searches for any little crevice or hole into which to insert itself. I had two young plants; and, after having observed this habit, I placed near them posts, which had been bored by beetles, or had become fissured by drying. The tendrils, by their own movement and by that of the internodes, slowly travelled over the surface of the wood, and when the apex came to a hole or fissure it inserted itself; in order to effect this the extremity for a length of half or quarter of an inch, would often bend itself at right angles to the basal part. I have watched this process between twenty and thirty times. The same tendril would frequently withdraw from one hole and insert its point into a second hole. I have also seen a tendril keep its point, in one case for 20 hrs. and in another for 36 hrs., in a minute hole, and then withdraw it. Whilst the point is thus temporarily inserted, the opposite tendril goes on revolving.

The whole length of a tendril often fits itself closely to any surface of wood with which it has come into contact; and I have observed one bent at right angles, from having entered a wide and deep fissure, with its apex abruptly re-bent and inserted into a minute lateral hole. After a tendril has clasped a stick, it contracts spirally; if it remains unattached it hangs straight downwards. If it has merely adapted itself to the inequalities of a thick post, though it has clasped nothing, or if it has inserted its apex into some little fissure, this stimulus suffices to induce spiral contraction; but the contraction always draws the tendril away from the post. So that in every case these movements, which seem so nicely adapted for some purpose, were useless. On one occasion, however, the tip became permanently jammed into a narrow fissure. I fully expected, from the analogy of B. capreolata and B. littoralis, that the tips would have been developed into adhesive discs; but I could never detect even a trace of this process. There is therefore at present something unintelligible about the habits of this plant.

Bignonia picta. — This species closely resembles the last in the structure and movements of its tendrils. I also casually examined a fine growing plant of the allied B. Lindleyi, and this apparently behaved in all respects in the same manner.

Bignonia capreolata. — We now come to a species having tendrils of a different type; but first for the internodes. A young shoot made three large revolutions, following the sun, at an average rate of 2 hrs. 23 m. The stem is thin and flexible, and I have seen one make four regular spiral turns round a thin upright stick, ascending of course from right to left, and therefore in a reversed direction compared with the before described species. Afterwards, from the interference of the tendrils, it ascended either straight up the stick or in an irregular spire. The tendrils are in some respects highly remarkable. In a young plant they were about 2.5 inches in length and much branched, the five chief branches apparently representing two pairs of leaflets and a terminal one. Each branch is, however, bifid or more commonly trifid towards the extremity, with the points blunt yet distinctly hooked. A tendril bends to any side which is lightly rubbed, and subsequently becomes straight again; but a loop of thread weighing 0.25th of a grain produced no effect. On two occasions the terminal branches became slightly curved in 10 m. after they had touched a stick; and in 30 m. the tips were curled quite round it. The basal part is less sensitive. The tendrils revolved in an apparently capricious manner, sometimes very slightly or not at all; at other times they described large regular ellipses. I could detect no spontaneous movement in the petioles of the leaves.

Whilst the tendrils are revolving more or less regularly, another remarkable movement takes place, namely, a slow inclination from the light towards the darkest side of the house. I repeatedly changed the position of my plants, and some little time after the revolving movement had ceased, the successively formed tendrils always ended by pointing to the darkest side. When I placed a thick post near a tendril, between it and the light, the tendril pointed in that direction. In two instances a pair of leaves stood so that one of the two tendrils was directed towards the light and the other to the darkest side of the house; the latter did not move, but the opposite one bent itself first upwards and then right over its fellow, so that the two became parallel, one above the other, both pointing to the dark: I then turned the plant half round; and the tendril which had turned over recovered its original position, and the opposite one which had not before moved, now turned over to the dark side. Lastly, on another plant, three pairs of tendrils were produced at the same time by three shoots, and all happened to be differently directed: I placed the pot in a box open only on one side, and obliquely facing the light; in two days all six tendrils pointed with unerring truth to the darkest corner of the box, though to do this each had to bend in a different manner. Six wind-vanes could not have more truly shown the direction of the wind, than did these branched tendrils the course of the stream of light which entered the box. I left these tendrils undisturbed for above 24 hrs., and then turned the pot half round; but they had now lost their power of movement, and could not any longer avoid the light.

When a tendril has not succeeded in clasping a support, either through its own revolving movement or that of the shoot, or by turning towards any object which intercepts the light, it bends vertically downwards and then towards its own stem, which it seizes together with the supporting stick, if there be one. A little aid is thus given in keeping the stem secure. If the tendril seizes nothing, it does not contract spirally, but soon withers away and drops off. If it seizes an object, all the branches contract spirally.

I have stated that after a tendril has come into contact with a stick, it bends round it in about half an hour; but I repeatedly observed, as in the case of B. speciosa and its allies, that it often again loosed the stick; sometimes seizing and loosing the same stick three or four times. Knowing that the tendrils avoided the light, I gave them a glass tube blackened within, and a well-blackened zinc plate: the branches curled round the tube and abruptly bent themselves round the edges of the zinc plate; but they soon recoiled from these objects with what I can only call disgust, and straightened themselves. I then placed a post with extremely rugged bark close to a pair of tendrils; twice they touched it for an hour or two, and twice they withdrew; at last one of the hooked extremities curled round and firmly seized an excessively minute projecting point of bark, and then the other branches spread themselves out, following with accuracy every inequality of the surface. I afterwards placed near the plant a post without bark but much fissured, and the points of the tendrils crawled into all the crevices in a beautiful manner. To my surprise, I observed that the tips of the immature tendrils, with the branches not yet fully separated, likewise crawled just like roots into the minutest crevices. In two or three days after the tips had thus crawled into the crevices, or after their hooked ends had seized minute points, the final process, now to be described, commenced.

This process I discovered by having accidentally left a piece of wool near a tendril; and this led me to bind a quantity of flax, moss, and wool loosely round sticks, and to place them near tendrils. The wool must not be dyed, for these tendrils are excessively sensitive to some poisons. The hooked points soon caught hold of the fibres, even loosely floating fibres, and now there was no recoiling; on the contrary, the excitement caused the hooks to penetrate the fibrous mass and to curl inwards, so that each hook caught firmly one or two fibres, or a small bundle of them. The tips and the inner surfaces of the hooks now began to swell, and in two or three days were visibly enlarged. After a few more days the hooks were converted into whitish, irregular balls, rather above the 0.05th of an inch (1.27 mm.) in diameter, formed of coarse cellular tissue, which sometimes wholly enveloped and concealed the hooks themselves. The surfaces of these balls secrete some viscid resinous matter, to which the fibres of the flax, &c., adhere. When a fibre has become fastened to the surface, the cellular tissue does not grow directly beneath it, but continues to grow closely on each side; so that when several adjoining fibres, though excessively thin, were caught, so many crests of cellular matter, each not as thick as a human hair, grew up between them, and these, arching over on both sides, adhered firmly together. As the whole surface of the ball continues to grow, fresh fibres adhere and are afterwards enveloped; so that I have seen a little ball with between fifty and sixty fibres of flax crossing it at various angles and all embedded more or less deeply. Every gradation in the process could be followed — some fibres merely sticking to the surface, others lying in more or less deep furrows, or deeply embedded, or passing through the very centre of the cellular ball. The embedded fibres are so closely clasped that they cannot be withdrawn. The outgrowing tissue has so strong a tendency to unite, that two balls produced by distinct tendrils sometimes unite and grow into a single one.

On one occasion, when a tendril had curled round a stick, half an inch in diameter, an adhesive disc was formed; but this does not generally occur in the case of smooth sticks or posts. If, however, the tip catches a minute projecting point, the other branches form discs, especially if they find crevices to crawl into. The tendrils failed to attach themselves to a brick wall.

I infer from the adherence of the fibres to the discs or balls, that these secrete some resinous adhesive matter; and more especially from such fibres becoming loose if immersed in sulphuric ether. This fluid likewise removes small, brown, glistening points which can generally be seen on the surfaces of the older discs. If the hooked extremities of the tendrils do not touch anything, discs, as far as I have seen, are never formed; 26 but temporary contact during a moderate time suffices to cause their development. I have seen eight discs formed on the same tendril. After their development the tendrils contract spirally, and become woody and very strong. A tendril in this state supported nearly seven ounces, and would apparently have supported a considerably greater weight, had not the fibres of flax to which the discs were attached yielded.

From the facts now given, we may infer that though the tendrils of this Bignonia can occasionally adhere to smooth cylindrical sticks and often to rugged bark, yet that they are specially adapted to climb trees clothed with lichens, mosses, or other such productions; and I hear from Professor Asa Gray that the Polypodium incanum abounds on the forest-trees in the districts of North America where this species of Bignonia grows. Finally, I may remark how singular a fact it is that a leaf should be metamorphosed into a branched organ which turns from the light, and which can by its extremities either crawl like roots into crevices, or seize hold of minute projecting points, these extremities afterwards forming cellular outgrowths which secrete an adhesive cement, and then envelop by their continued growth the finest fibres.

Eccremocarpus scaber (Bignoniaceae). — Plants, though growing pretty well in my green-house, showed no spontaneous movements in their shoots or tendrils; but when removed to the hot-house, the young internodes revolved at rates varying from 3 hrs. 15 m. to 1 hr. 13 m. One large circle was swept at this latter unusually quick rate; but generally the circles or ellipses were small, and sometimes the course pursued was quite irregular. An internode, after making several revolutions, sometimes stood still for 12 hrs. or 18 hrs., and then recommenced revolving. Such strongly marked interruptions in the movements of the internodes I have observed in hardly any other plant.

The leaves bear four leaflets, themselves subdivided, and terminate in much-branched tendrils. The main petiole of the leaf, whilst young, moves spontaneously, and follows nearly the same irregular course and at about the same rate as the internodes. The movement to and from the stem is the most conspicuous, and I have seen the chord of a curved petiole which formed an angle of 59 degrees with the stem, in an hour afterwards making an angle of 106 degrees. The two opposite petioles do not move together, and one is sometimes so much raised as to stand close to the stem, whilst the other is not far from horizontal. The basal part of the petiole moves less than the distal part. The tendrils, besides being carried by the moving petioles and internodes, themselves move spontaneously; and the opposite tendrils occasionally move in opposite directions. By these combined movements of the young internodes, petioles, and tendrils, a considerable space is swept in search of a support.

In young plants the tendrils are about three inches in length: they bear two lateral and two terminal branches; and each branch bifurcates twice, with the tips terminating in blunt double hooks, having both points directed to the same side. All the branches are sensitive on all sides; and after being lightly rubbed, or after coming into contact with a stick, bend in about 10 m. One which had become curved in 10 m. after a light rub, continued bending for between 3 hrs. and 4 hrs., and became straight again in 8 hrs. or 9 hrs. Tendrils, which have caught nothing, ultimately contract into an irregular spire, as they likewise do, only much more quickly, after clasping a support. In both cases the main petiole bearing the leaflets, which is at first straight and inclined a little upwards, moves downwards, with the middle part bent abruptly into a right angle; but this is seen in E. miniatus more plainly than in E. scaber. The tendrils in this genus act in some respects like those of Bignonia capreolata; but the whole does not move from the light, nor do the hooked tips become enlarged into cellular discs. After the tendrils have come into contact with a moderately thick cylindrical stick or with rugged bark, the several branches may be seen slowly to lift themselves up, change their positions, and again come into contact with the supporting surface. The object of these movements is to bring the double-hooks at the extremities of the branches, which naturally face in all directions, into contact with the wood. I have watched a tendril, half of which had bent itself at right angles round the sharp corner of a square post, neatly bring every single hook into contact with both rectangular surfaces. The appearance suggested the belief, that though the whole tendril is not sensitive to light, yet that the tips are so, and that they turn and twist themselves towards any dark surface. Ultimately the branches arrange themselves very neatly to all the irregularities of the most rugged bark, so that they resemble in their irregular course a river with its branches, as engraved on a map. But when a tendril has wound round a rather thick stick, the subsequent spiral contraction generally draws it away and spoils the neat arrangement. So it is, but not in quite so marked a manner, when a tendril has spread itself over a large, nearly flat surface of rugged bark. We may therefore conclude that these tendrils are not perfectly adapted to seize moderately thick sticks or rugged bark. If a thin stick or twig is placed near a tendril, the terminal branches wind quite round it, and then seize their own lower branches or the main stem. The stick is thus firmly, but not neatly, grasped. What the tendrils are really adapted for, appears to be such objects as the thin culms of certain grasses, or the long flexible bristles of a brush, or thin rigid leaves such as those of the Asparagus, all of which they seize in an admirable manner. This is due to the extremities of the branches close to the little hooks being extremely sensitive to a touch from the thinnest object, which they consequently curl round and clasp. When a small brush, for instance, was placed near a tendril, the tips of each sub-branch seized one, two, or three of the bristles; and then the spiral contraction of the several branches brought all these little parcels close together, so that thirty or forty bristles were drawn into a single bundle, which afforded an excellent support.

POLEMONIACEAE. — Cobaea scandens. — This is an excellently constructed climber. The tendrils on a fine plant were eleven inches long, with the petiole bearing two pairs of leaflets, only two and a half inches in length. They revolve more rapidly and vigorously than those of any other tendril-bearer observed by me, with the exception of one kind of Passiflora. Three large, nearly circular sweeps, directed against the sun were completed, each in 1 hr. 15 m.; and two other circles in 1 hr. 20 m. and 1 hr. 23 m. Sometimes a tendril travels in a much inclined position, and sometimes nearly upright. The lower part moves but little and the petiole not at all; nor do the internodes revolve; so that here we have the tendril alone moving. On the other hand, with most of the species of Bignonia and the Eccremocarpus, the internodes, tendrils, and petioles all revolved. The long, straight, tapering main stem of the tendril of the Cobaea bears alternate branches; and each branch is several times divided, with the finer branches as thin as very thin bristles and extremely flexible, so that they are blown about by a breath of air; yet they are strong and highly elastic. The extremity of each branch is a little flattened, and terminates in a minute double (though sometimes single) hook, formed of a hard, translucent, woody substance, and as sharp as the finest needle. On a tendril which was eleven inches long I counted ninety-four of these beautifully constructed little hooks. They readily catch soft wood, or gloves, or the skin of the naked hand. With the exception of these hardened hooks, and of the basal part of the central stem, every part of every branchlet is highly sensitive on all sides to a slight touch, and bends in a few minutes towards the touched side. By lightly rubbing several sub-branches on opposite sides, the whole tendril rapidly assumed an extraordinarily crooked shape. These movements from contact do not interfere with the ordinary revolving movement. The branches, after becoming greatly curved from being touched, straighten themselves at a quicker rate than in almost any other tendril seen by me, namely, in between half an hour and an hour. After the tendril has caught any object, spiral contraction likewise begins after an unusually short interval of time, namely, in about twelve hours.

Before the tendril is mature, the terminal branchlets cohere, and the hooks are curled closely inwards. At this period no part is sensitive to a touch; but as soon as the branches diverge and the hooks stand out, full sensitiveness is acquired. It is a singular circumstance that immature tendrils revolve at their full velocity before they become sensitive, but in a useless manner, as in this state they can catch nothing. This want of perfect coadaptation, though only for a short time, between the structure and the functions of a climbing-plant is a rare event. A tendril, as soon as it is ready to act, stands, together with the supporting petiole, vertically upwards. The leaflets borne by the petiole are at this time quite small, and the extremity of the growing stem is bent to one side so as to be out of the way of the revolving tendril, which sweeps large circles directly over head. The tendrils thus revolve in a position well adapted for catching objects standing above; and by this means the ascent of the plant is favoured. If no object is caught, the leaf with its tendril bends downwards and ultimately assumes a horizontal position. An open space is thus left for the next succeeding and younger tendril to stand vertically upwards and to revolve freely. As soon as an old tendril bends downwards, it loses all power of movement, and contracts spirally into an entangled mass. Although the tendrils revolve with unusual rapidity, the movement lasts for only a short time. In a plant placed in the hot-house and growing vigorously, a tendril revolved for not longer than 36 hours, counting from the period when it first became sensitive; but during this period it probably made at least 27 revolutions.

When a revolving tendril strikes against a stick, the branches quickly bend round and clasp it. The little hooks here play an important part, as they prevent the branches from being dragged away by the rapid revolving movement, before they have had time to clasp the stick securely. This is especially the case when only the extremity of a branch has caught hold of a support. As soon as a tendril has bent a smooth stick or a thick rugged post, or has come into contact with planed wood (for it can adhere temporarily even to so smooth a surface as this), the same peculiar movements may be observed as those described under Bignonia capreolata and Eccremocarpus. The branches repeatedly lift themselves up and down; those which have their hooks already directed downwards remaining in this position and securing the tendril, whilst the others twist about until they succeed in arranging themselves in conformity with every irregularity of the surface, and in bringing their hooks into contact with the wood. The use of the hooks was well shown by giving the tendrils tubes and slips of glass to catch; for these, though temporarily seized, were invariably lost, either during the re-arrangement of the branches or ultimately when spiral contraction ensued.

The perfect manner in which the branches arranged themselves, creeping like rootlets over every inequality of the surface and into any deep crevice, is a pretty sight; for it is perhaps more effectually performed by this than by any other species. The action is certainly more conspicuous, as the upper surfaces of the main stem, as well as of every branch to the extreme hooks, are angular and green, whilst the lower surfaces are rounded and purple. I was led to infer, as in former cases, that a less amount of light guided these movements of the branches of the tendrils. I made many trials with black and white cards and glass tubes to prove it, but failed from various causes; yet these trials countenanced the belief. As a tendril consists of a leaf split into numerous segments, there is nothing surprising in all the segments turning their upper surfaces towards the light, as soon as the tendril is caught and the revolving movement is arrested. But this will not account for the whole movement, for the segments actually bend or curve to the dark side besides turning round on their axes so that their upper surfaces may face the light.

When the Cobaea grows in the open air, the wind must aid the extremely flexible tendrils in seizing a support, for I found that a mere breath sufficed to cause the extreme branches to catch hold by their hooks of twigs, which they could not have reached by the revolving movement. It might have been thought that a tendril, thus hooked by the extremity of a single branch, could not have fairly grasped its support. But several times I watched cases like the following: tendril caught a thin stick by the hooks of one of its two extreme branches; though thus held by the tip, it still tried to revolve, bowing itself to all sides, and by this movement the other extreme branch soon caught the stick. The first branch then loosed itself, and, arranging its hooks, again caught hold. After a time, from the continued movement of the tendril, the hooks of a third branch caught hold. No other branches, as the tendril then stood, could possibly have touched the stick. But before long the upper part of the main stem began to contract into an open spire. It thus dragged the shoot which bore the tendril towards the stick; and as the tendril continually tried to revolve, a fourth branch was brought into contact. And lastly, from the spiral contraction travelling down both the main stem and the branches, all of them, one after another, were ultimately brought into contact with the stick. They then wound themselves round it and round one another, until the whole tendril was tied together in an inextricable knot. The tendrils, though at first quite flexible, after having clasped a support for a time, become more rigid and stronger than they were at first. Thus the plant is secured to its support in a perfect manner.

LEGUMINOSAE. — Pisum sativum. — The common pea was the subject of a valuable memoir by Dutrochet, 27 who discovered that the internodes and tendrils revolve in ellipses. The ellipses are generally very narrow, but sometimes approach to circles. I several times observed that the longer axis slowly changed its direction, which is of importance, as the tendril thus sweeps a wider space. Owing to this change of direction, and likewise to the movement of the stem towards the light, the successive irregular ellipses generally form an irregular spire. I have thought it worth while to annex a tracing of the course pursued by the upper internode (the movement of the tendril being neglected) of a young plant from 8.40 A.M. to 9.15 P.M. The course was traced on a hemispherical glass placed over the plant, and the dots with figures give the hours of observation; each dot being joined by a straight line. No doubt all the lines would have been curvilinear if the course had been observed at much shorter intervals. The extremity of the petiole, from which the young tendril arose, was two inches from the glass, so that if a pencil two inches in length could have been affixed to the petiole, it would have traced the annexed figure on the under side of the glass; but it must be remembered that the figure is reduced by one-half. Neglecting the first great sweep towards the light from the figure 1 to 2, the end of the petiole swept a space 4 inches across in one direction, and 3 inches in another. As a full-grown tendril is considerably above two inches in length, and as the tendril itself bends and revolves in harmony with the internode, a considerably wider space is swept than is here represented on a reduced scale. Dutrochet observed the completion of an ellipse in 1 hr. 20 m.; and I saw one completed in 1 hr. 30 m. The direction followed is variable, either with or against the sun.

Dutrochet asserts that the petioles of the leaves spontaneously revolve, as well as the young internodes and tendrils; but he does not say that he secured the internodes; when this was done, I could never detect any movement in the petiole, except to and from the light.

The tendrils, on the other hand, when the internodes and petioles are secured, describe irregular spires or regular ellipses, exactly like those made by the internodes. A young tendril, only 1.125 of an inch in length, revolved. Dutrochet has shown that when a plant is placed in a room, so that the light enters laterally, the internodes travel much quicker to the light than from it: on the other hand, he asserts that the tendril itself moves from the light towards the dark side of the room. With due deference to this great observer, I think he was mistaken, owing to his not having secured the internodes. I took a young plant with highly sensitive tendrils, and tied the petiole so that the tendril alone could move; it completed a perfect ellipse in 1 hr. 30 m.; I then turned the plant partly round, but this made no change in the direction of the succeeding ellipse. The next day I watched a plant similarly secured until the tendril (which was highly sensitive) made an ellipse in a line exactly to and from the light; the movement was so great that the tendril at the two ends of its elliptical course bent itself a little beneath the horizon, thus travelling more than 180 degrees; but the curvature was fully as great towards the light as towards the dark side of the room. I believe Dutrochet was misled by not having secured the internodes, and by having observed a plant of which the internodes and tendrils no longer curved in harmony together, owing to inequality of age.

Dutrochet made no observations on the sensitiveness of the tendrils. These, whilst young and about an inch in length with the leaflets on the petiole only partially expanded, are highly sensitive; a single light touch with a twig on the inferior or concave surface near the tip caused them to bend quickly, as did occasionally a loop of thread weighing one-seventh of a grain (9.25 mg.). The upper or convex surface is barely or not at all sensitive. Tendrils, after bending from a touch, straighten themselves in about two hours, and are then ready to act again. As soon as they begin to grow old, the extremities of their two or three pairs of branches become hooked, and they then appear to form an excellent grappling instrument; but this is not the case. For at this period they have generally quite lost their sensitiveness; and when hooked on to twigs, some were not at all affected, and others required from 18 hrs. to 24 hrs. before clasping such twigs; nevertheless, they were able to utilise the last vestige of irritability owing to their extremities being hooked. Ultimately the lateral branches contract spirally, but not the middle or main stem.

Lathyrus aphaca. — This plant is destitute of leaves, except during a very early age, these being replaced by tendrils, and the leaves themselves by large stipules. It might therefore have been expected that the tendrils would have been highly organized, but this is not so. They are moderately long, thin, and unbranched, with their tips slightly curved. Whilst young they are sensitive on all sides, but chiefly on the concave side of the extremity. They have no spontaneous revolving power, but are at first inclined upwards at an angle of about 45 degrees, then move into a horizontal position, and ultimately bend downwards. The young internodes, on the other hand, revolve in ellipses, and carry with them the tendrils. Two ellipses were completed, each in nearly 5 hrs.; their longer axes were directed at about an angle of 45 degrees to the axis of the previously made ellipse.

Lathyrus grandiflorus. — The plants observed were young and not growing vigorously, yet sufficiently so, I think, for my observations to be trusted. If so, we have the rare case of neither internodes nor tendrils revolving. The tendrils of vigorous plants are above 4 inches in length, and are often twice divided into three branches; the tips are curved and are sensitive on their concave sides; the lower part of the central stem is hardly at all sensitive. Hence this plant appears to climb simply by its tendrils being brought, through the growth of the stem, or more efficiently by the wind, into contact with surrounding objects, which they then clasp. I may add that the tendrils, or the internodes, or both, of Vicia sativa revolve.

COMPOSITAE. — Mutisia clematis. — The immense family of the Compositae is well known to include very few climbing plants. We have seen in the Table in the first chapter that Mikania scandens is a regular twiner, and F. Muller informs me that in S. Brazil there is another species which is a leaf-climber. Mutisia is the only genus in the family, as far as I can learn, which bears tendrils: it is therefore interesting to find that these, though rather less metamorphosed from their primordial foliar condition than are most other tendrils, yet display all the ordinary characteristic movements, both those that are spontaneous and those which are excited by contact.

The long leaf bears seven or eight alternate leaflets, and terminates in a tendril which, in a plant of considerable size, was 5 inches in length. It consists generally of three branches; and these, although much elongated, evidently represent the petioles and midribs of three leaflets; for they closely resemble the same parts in an ordinary leaf, in being rectangular on the upper surface, furrowed, and edged with green. Moreover, the green edging of the tendrils of young plants sometimes expands into a narrow lamina or blade. Each branch is curved a little downwards, and is slightly hooked at the extremity.

A young upper internode revolved, judging from three revolutions, at an average rate of 1 hr. 38 m.; it swept ellipses with the longer axes directed at right angles to one another; but the plant, apparently, cannot twine. The petioles and the tendrils are both in constant movement. But their movement is slower and much less regularly elliptical than that of the internodes. They appear to be much affected by the light, for the whole leaf usually sinks down during the night and rises during the day, moving, also, during the day in a crooked course to the west. The tip of the tendril is highly sensitive on the lower surface; and one which was just touched with a twig became perceptibly curved in 3 m., and another in 5 m.; the upper surface is not at all sensitive; the sides are moderately sensitive, so that two branches which were rubbed on their inner sides converged and crossed each other. The petiole of the leaf and the lower parts of the tendril, halfway between the upper leaflet and the lowest branch, are not sensitive. A tendril after curling from a touch became straight again in about 6 hrs., and was ready to re-act; but one that had been so roughly rubbed as to have coiled into a helix did not become perfectly straight until after 13 hrs. The tendrils retain their sensibility to an unusually late age; for one borne by a leaf with five or six fully developed leaves above, was still active. If a tendril catches nothing, after a considerable interval of time the tips of the branches curl a little inwards; but if it clasps some object, the whole contracts spirally.

SMILACEAE. — Smilax aspera, var. maculata. — Aug. St.-Hilaire 28 considers that the tendrils, which rise in pairs from the petiole, are modified lateral leaflets; but Mohl (p. 41) ranks them as modified stipules. These tendrils are from 1.5 to 1.75 inches in length, are thin, and have slightly curved, pointed extremities. They diverge a little from each other, and stand at first nearly upright. When lightly rubbed on either side, they slowly bend to that side, and subsequently become straight again. The back or convex side when placed in contact with a stick became just perceptibly curved in 1 hr. 20 m., but did not completely surround it until 48 hrs. had elapsed; the concave side of another became considerably curved in 2 hrs. and clasped a stick in 5 hrs. As the pairs of tendrils grow old, one tendril diverges more and more from the other, and both slowly bend backwards and downwards, so that after a time they project on the opposite side of the stem to that from which they arise. They then still retain their sensitiveness, and can clasp a support placed BEHIND the stem. Owing to this power, the plant is able to ascend a thin upright stick. Ultimately the two tendrils belonging to the same petiole, if they do not come into contact with any object, loosely cross each other behind the stem, as at B, in fig. 7. This movement of the tendrils towards and round the stem is, to a certain extent, guided by their avoidance of the light; for when a plant stood so that one of the two tendrils was compelled in thus slowly moving to travel towards the light, and the other from the light, the latter always moved, as I repeatedly observed, more quickly than its fellow. The tendrils do not contract spirally in any case. Their chance of finding a support depends on the growth of the plant, on the wind, and on their own slow backward and downward movement, which, as we have just seen, is guided, to a certain extent, by the avoidance of the light; for neither the internodes nor the tendrils have any proper revolving movement. From this latter circumstance, from the slow movements of the tendrils after contact (though their sensitiveness is retained for an unusual length of time), from their simple structure and shortness, this plant is a less perfect climber than any other tendril-bearing species observed by me. The plant whilst young and only a few inches in height, does not produce any tendrils; and considering that it grows to only about 8 feet in height, that the stem is zigzag and is furnished, as well as the petioles, with spines, it is surprising that it should be provided with tendrils, comparatively inefficient though these are. The plant might have been left, one would have thought, to climb by the aid of its spines alone, like our brambles. As, however, it belongs to a genus, some of the species of which are furnished with much longer tendrils, we may suspect that it possesses these organs solely from being descended from progenitors more highly organized in this respect.

FUMARIACEAE. — Corydalis claviculata. — According to Mohl (p. 43), the extremities of the branched stem, as well as the leaves, are converted into tendrils. In the specimens examined by me all the tendrils were certainly foliar, and it is hardly credible that the same plant should produce tendrils of a widely different homological nature. Nevertheless, from this statement by Mohl, I have ranked this species amongst the tendril-bearers; if classed exclusively by its foliar tendrils, it would be doubtful whether it ought not to have been placed amongst the leaf-climbers, with its allies, Fumaria and Adlumia. A large majority of its so-called tendrils still bear leaflets, though excessively reduced in size; but some few of them may properly be designated as tendrils, for they are completely destitute of laminae or blades. Consequently, we here behold a plant in an actual state of transition from a leaf-climber to a tendril-bearer. Whilst the plant is rather young, only the outer leaves, but when full-grown all the leaves, have their extremities converted into more or less perfect tendrils. I have examined specimens from one locality alone, viz. Hampshire; and it is not improbable that plants growing under different conditions might have their leaves a little more or less changed into true tendrils.

Whilst the plant is quite young, the first-formed leaves are not modified in any way, but those next formed have their terminal leaflets reduced in size, and soon all the leaves assume the structure represented in the following drawing. This leaf bore nine leaflets; the lower ones being much subdivided. The terminal portion of the petiole, about 1.5 inch in length (above the leaflet f), is thinner and more elongated than the lower part, and may be considered as the tendril. The leaflets borne by this part are greatly reduced in size, being, on an average, about the tenth of an inch in length and very narrow; one small leaflet measured one-twelfth of an inch in length and one-seventy-fifth in breadth (2.116 mm. and 0.339 mm.), so that it was almost microscopically minute. All the reduced leaflets have branching nerves, and terminate in little spines, like those of the fully developed leaflets. Every gradation could be traced, until we come to branchlets (as a and d in the figure) which show no vestige of a lamina or blade. Occasionally all the terminal branchlets of the petiole are in this condition, and we then have a true tendril.

The several terminal branches of the petiole bearing the much reduced leaflets (a, b, c, d) are highly sensitive, for a loop of thread weighing only the one-sixteenth of a grain (4.05 mg.) caused them to become greatly curved in under 4 hrs. When the loop was removed, the petioles straightened themselves in about the same time. The petiole (e) was rather less sensitive; and in another specimen, in which the corresponding petiole bore rather larger leaflets, a loop of thread weighing one-eighth of a grain did not cause curvature until 18 hrs. had elapsed. Loops of thread weighing one-fourth of a grain, left suspended on the lower petioles (f to l) during several days, produced no effect. Yet the three petioles f, g, and h were not quite insensible, for when left in contact with a stick for a day or two they slowly curled round it. Thus the sensibility of the petiole gradually diminishes from the tendril-like extremity to the base. The internodes of the stem are not at all sensitive, which makes Mohl’s statement that they are sometimes converted into tendrils the more surprising, not to say improbable.

The whole leaf, whilst young and sensitive, stands almost vertically upwards, as we have seen to be the case with many tendrils. It is in continual movement, and one that I observed swept at an average rate of about 2 hrs. for each revolution, large, though irregular, ellipses, which were sometimes narrow, sometimes broad, with their longer axes directed to different points of the compass. The young internodes, likewise revolved irregularly in ellipses or spires; so that by these combined movements a considerable space was swept for a support. If the terminal and attenuated portion of a petiole fails to seize any object, it ultimately bends downwards and inwards, and soon loses all irritability and power of movement. This bending down differs much in nature from that which occurs with the extremities of the young leaves in many species of Clematis; for these, when thus bent downwards or hooked, first acquire their full degree of sensitiveness.

Dicentra thalictrifolia. — In this allied plant the metamorphosis of the terminal leaflets is complete, and they are converted into perfect tendrils. Whilst the plant is young, the tendrils appear like modified branches, and a distinguished botanist thought that they were of this nature; but in a full-grown plant there can be no doubt, as I am assured by Dr. Hooker, that they are modified leaves. When of full size, they are above 5 inches in length; they bifurcate twice, thrice, or even four times; their extremities are hooked and blunt. All the branches of the tendrils are sensitive on all sides, but the basal portion of the main stem is only slightly so. The terminal branches when lightly rubbed with a twig became curved in the course of from 30 m. to 42 m., and straightened themselves in between 10 hrs. and 20 hrs. A loop of thread weighing one-eighth of a grain plainly caused the thinner branches to bend, as did occasionally a loop weighing one-sixteenth of a grain; but this latter weight, though left suspended, was not sufficient to cause a permanent flexure. The whole leaf with its tendril, as well as the young upper internodes, revolves vigorously and quickly, though irregularly, and thus sweeps a wide space. The figure traced on a bell-glass was either an irregular spire or a zigzag line. The nearest approach to an ellipse was an elongated figure of 8, with one end a little open, and this was completed in 1 hr. 53 m. During a period of 6 hrs. 17 m. another shoot made a complex figure, apparently representing three and a half ellipses. When the lower part of the petiole bearing the leaflets was securely fastened, the tendril itself described similar but much smaller figures.

This species climbs well. The tendrils after clasping a stick become thicker and more rigid; but the blunt hooks do not turn and adapt themselves to the supporting surface, as is done in so perfect a manner by some Bignoniaceae and Cobaea. The tendrils of young plants, two or three feet in height, are only half the length of those borne by the same plant when grown taller, and they do not contract spirally after clasping a support, but only become slightly flexuous. Full-sized tendrils, on the other hand, contract spirally, with the exception of the thick basal portion. Tendrils which have caught nothing simply bend downwards and inwards, like the extremities of the leaves of the Corydalis claviculata. But in all cases the petiole after a time is angularly and abruptly bent downwards like that of Eccremocarpus.

24 Never having had the opportunity of examining tendrils produced by the modification of branches, I spoke doubtfully about them in this essay when originally published. But since then Fritz Muller has described (Journal of Linn. Soc. vol. ix. p. 344) many striking cases in South Brazil. In speaking of plants which climb by the aid of their branches, more or less modified, he states that the following stages of development can be traced: (1.) Plants supporting themselves simply by their branches stretched out at right angles — for example, Chiococca. (2.) Plants clasping a support with their unmodified branches, as with Securidaca. (3.) Plants climbing by the extremities of their branches which appear like tendrils, as is the case according to Endlicher with Helinus. (4.) Plants with their branches much modified and temporarily converted into tendrils, but which may be again transformed into branches, as with certain Papilionaceous plants. (5.) Plants with their branches forming true tendrils, and used exclusively for climbing — as with Strychnos and Caulotretus. Even the unmodified branches become much thickened when they wind round a support. I may add that Mr. Thwaites sent me from Ceylon a specimen of an Acacia which had climbed up the trunk of a rather large tree, by the aid of tendril-like, curved or convoluted branchlets, arrested in their growth and furnished with sharp recurved hooks.

25 As far as I can make out, the history of our knowledge of tendrils is as follows:— We have seen that Palm and von Mohl observed about the same time the singular phenomenon of the spontaneous revolving movement of twining-plants. Palm (p. 58), I presume, observed likewise the revolving movement of tendrils; but I do not feel sure of this, for he says very little on the subject. Dutrochet fully described this movement of the tendril in the common pea. Mohl first discovered that tendrils are sensitive to contact; but from some cause, probably from observing too old tendrils, he was not aware how sensitive they were, and thought that prolonged pressure was necessary to excite their movement. Professor Asa Gray, in a paper already quoted, first noticed the extreme sensitiveness and rapidity of the movements of the tendrils of certain Cucurbitaceous plants.

26 Fritz Muller states (ibid. p. 348) that in South Brazil the trifid tendrils of Haplolophium, (one of the Bignoniaceae) without having come into contact with any object, terminate in smooth shining discs. These, however, after adhering to any object, sometimes become considerably enlarged.

27 Comptes Rendus, tom. xvii. 1843, p. 989.

28 ‘Lecons de Botanique,’ &c., 1841, p. 170.

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