The Movements and Habits of Climbing Plants · Charles Darwin
CHAPTER III. TENDRIL-BEARERS.
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; {102} 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_ (_Bignoniaceæ_).—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° with the stem, in an hour afterwards making an angle of 106°. 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.
POLEMONIACEÆ.—_Cobæa 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 _Cobæa_ 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 co-adaptation, 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 _Cobæa_ 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.
LEGUMINOSÆ.—_Pisum sativum_.—The common pea was the subject of a valuable memoir by Dutrochet, {111} 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 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°, 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° 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.
COMPOSITÆ.—_Mutisia clematis_.—The immense family of the Compositæ 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. Müller 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.
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.
Text from Project Gutenberg, public domain in the United States.