Plants climbing by the aid of hooks, or merely scrambling over other plants — Root-climbers, adhesive matter secreted by the rootlets — General conclusions with respect to climbing plants, and the stages of their development.
Hook–Climbers. — In my introductory remarks, I stated that, besides the two first great classes of climbing plants, namely, those which twine round a support, and those endowed with irritability enabling them to seize hold of objects by means of their petioles or tendrils, there are two other classes, hook-climbers and root-climbers. Many plants, moreover, as Fritz Muller has remarked, 40 climb or scramble up thickets in a still more simple fashion, without any special aid, excepting that their leading shoots are generally long and flexible. It may, however, be suspected from what follows, that these shoots in some cases tend to avoid the light. The few hook-climbers which I have observed, namely, Galium aparine, Rubus australis, and some climbing Roses, exhibit no spontaneous revolving movement. If they had possessed this power, and had been capable of twining, they would have been placed in the class of Twiners; for some twiners are furnished with spines or hooks, which aid them in their ascent. For instance, the Hop, which is a twiner, has reflexed hooks as large as those of the Galium; some other twiners have stiff reflexed hairs; and Dipladenia has a circle of blunt spines at the bases of its leaves. I have seen only one tendril-bearing plant, namely, Smilax aspera, which is furnished with reflexed spines; but this is the case with several branch-climbers in South Brazil and Ceylon; and their branches graduate into true tendrils. Some few plants apparently depend solely on their hooks for climbing, and yet do so efficiently, as certain palms in the New and Old Worlds. Even some climbing Roses will ascend the walls of a tall house, if covered with a trellis. How this is effected I know not; for the young shoots of one such Rose, when placed in a pot in a window, bent irregularly towards the light during the day and from the light during the night, like the shoots of any common plant; so that it is not easy to understand how they could have got under a trellis close to the wall. 41
Root-climbers. — A good many plants come under this class, and are excellent climbers. One of the most remarkable is the Marcgravia umbellata, the stem of which in the tropical forests of South America, as I hear from Mr. Spruce, grows in a curiously flattened manner against the trunks of trees; here and there it puts forth claspers (roots), which adhere to the trunk, and, if the latter be slender, completely embrace it. When this plant has climbed to the light, it produces free branches with rounded stems, clad with sharp-pointed leaves, wonderfully different in appearance from those borne by the stem as long as it remains adherent. This surprising difference in the leaves, I have also observed in a plant of Marcgravia dubia in my hothouse. Root-climbers, as far as I have seen, namely, the Ivy (Hedera helix), Ficus repens, and F. barbatus, have no power of movement, not even from the light to the dark. As previously stated, the Hoya carnosa (Asclepiadaceae) is a spiral twiner, and likewise adheres by rootlets even to a flat wall. The tendril-bearing Bignonia Tweedyana emits roots, which curve half round and adhere to thin sticks. The Tecoma radicans (Bignoniaceae), which is closely allied to many spontaneously revolving species, climbs by rootlets; nevertheless, its young shoots apparently move about more than can be accounted for by the varying action of the light.
I have not closely observed many root-climbers, but can give one curious fact. Ficus repens climbs up a wall just like Ivy; and when the young rootlets are made to press lightly on slips of glass, they emit after about a week’s interval, as I observed several times, minute drops of clear fluid, not in the least milky like that exuded from a wound. This fluid is slightly viscid, but cannot be drawn out into threads. It has the remarkable property of not soon drying; a drop, about the size of half a pin’s head, was slightly spread out on glass, and I scattered on it some minute grains of sand. The glass was left exposed in a drawer during hot and dry weather, and if the fluid had been water, it would certainly have dried in a few minutes; but it remained fluid, closely surrounding each grain of sand, during 128 days: how much longer it would have remained I cannot say. Some other rootlets were left in contact with the glass for about ten days or a fortnight, and the drops of secreted fluid were now rather larger, and so viscid that they could be drawn out into threads. Some other rootlets were left in contact during twenty-three days, and these were firmly cemented to the glass. Hence we may conclude that the rootlets first secrete a slightly viscid fluid, subsequently absorb the watery parts, (for we have seen that the fluid will not dry by itself,) and ultimately leave a cement. When the rootlets were torn from the glass, atoms of yellowish matter were left on it, which were partly dissolved by a drop of bisulphide of carbon; and this extremely volatile fluid was rendered very much less volatile by what it had dissolved.
As the bisulphide of carbon has a strong power of softening indurated caoutchouc, I soaked in it during a short time several rootlets of a plant which had grown up a plaistered wall; and I then found many extremely thin threads of transparent, not viscid, excessively elastic matter, precisely like caoutchouc, attached to two sets of rootlets on the same branch. These threads proceeded from the bark of the rootlet at one end, and at the other end were firmly attached to particles of silex or mortar from the wall. There could be no mistake in this observation, as I played with the threads for a long time under the microscope, drawing them out with my dissecting-needles and letting them spring back again. Yet I looked repeatedly at other rootlets similarly treated, and could never again discover these elastic threads. I therefore infer that the branch in question must have been slightly moved from the wall at some critical period, whilst the secretion was in the act of drying, through the absorption of its watery parts. The genus Ficus abounds with caoutchouc, and we may conclude from the facts just given that this substance, at first in solution and ultimately modified into an unelastic cement, 42 is used by the Ficus repens to cement its rootlets to any surface which it ascends. Whether other plants, which climb by their rootlets, emit any cement I do not know; but the rootlets of the Ivy, placed against glass, barely adhered to it, yet secreted a little yellowish matter. I may add, that the rootlets of the Marcgravia dubia can adhere firmly to smooth painted wood.
Vanilla aromatica emits aerial roots a foot in length, which point straight down to the ground. According to Mohl (p. 49), these crawl into crevices, and when they meet with a thin support, wind round it, as do tendrils. A plant which I kept was young, and did not form long roots; but on placing thin sticks in contact with them, they certainly bent a little to that side, in the course of about a day, and adhered by their rootlets to the wood; but they did not bend quite round the sticks, and afterwards they re-pursued their downward course. It is probable that these slight movements of the roots are due to the quicker growth of the side exposed to the light, in comparison with the other side, and not because the roots are sensitive to contact in the same manner as true tendrils. According to Mohl, the rootlets of certain species of Lycopodium act as tendrils. 43
Concluding Remarks on Climbing Plants.
Plants become climbers, in order, as it may be presumed, to reach the light, and to expose a large surface of their leaves to its action and to that of the free air. This is effected by climbers with wonderfully little expenditure of organized matter, in comparison with trees, which have to support a load of heavy branches by a massive trunk. Hence, no doubt, it arises that there are so many climbing plants in all quarters of the world, belonging to so many different orders. These plants have been arranged under four classes, disregarding those which merely scramble over bushes without any special aid. Hook-climbers are the least efficient of all, at least in our temperate countries, and can climb only in the midst of an entangled mass of vegetation. Root-climbers are excellently adapted to ascend naked faces of rock or trunks of trees; when, however, they climb trunks they are compelled to keep much in the shade; they cannot pass from branch to branch and thus cover the whole summit of a tree, for their rootlets require long-continued and close contact with a steady surface in order to adhere. The two great classes of twiners and of plants with sensitive organs, namely, leaf-climbers and tendril-bearers taken together, far exceed in number and in the perfection of their mechanism the climbers of the two first classes. Those which have the power of spontaneously revolving and of grasping objects with which they come in contact, easily pass from branch to branch, and securely ramble over a wide, sun-lit surface.
The divisions containing twining plants, leaf-climbers, and tendril-bearers graduate to a certain extent into one another, and nearly all have the same remarkable power of spontaneously revolving. Does this gradation, it may be asked, indicate that plants belonging to one subdivision have actually passed during the lapse of ages, or can pass, from one state to the other? Has, for instance, any tendril-bearing plant assumed its present structure without having previously existed as a leaf-climber or a twiner? If we consider leaf-climbers alone, the idea that they were primordially twiners is forcibly suggested. The internodes of all, without exception, revolve in exactly the same manner as twiners; some few can still twine well, and many others in an imperfect manner. Several leaf-climbing genera are closely allied to other genera which are simple twiners. It should also be observed, that the possession of leaves with sensitive petioles, and with the consequent power of clasping an object, would be of comparatively little use to a plant, unless associated with revolving internodes, by which the leaves are brought into contact with a support; although no doubt a scrambling plant would be apt, as Professor Jaeger has remarked, to rest on other plants by its leaves. On the other hand, revolving internodes, without any other aid, suffice to give the power of climbing; so that it seems probable that leaf-climbers were in most cases at first twiners, and subsequently became capable of grasping a support; and this, as we shall presently see, is a great additional advantage.
From analogous reasons, it is probable that all tendril-bearers were primordially twiners, that is, are the descendants of plants having this power and habit. For the internodes of the majority revolve; and, in a few species, the flexible stem still retains the capacity of spirally twining round an upright stick. Tendril-bearers have undergone much more modification than leaf-climbers; hence it is not surprising that their supposed primordial habits of revolving and twining have been more frequently lost or modified than in the case of leaf-climbers. The three great tendril-bearing families in which this loss has occurred in the most marked manner, are the Cucurbitaceae, Passifloraceae, and Vitaceae. In the first, the internodes revolve; but I have heard of no twining form, with the exception (according to Palm, p. 29. 52) of Momordica balsamina, and this is only an imperfect twiner. In the two other families I can hear of no twiners; and the internodes rarely have the power of revolving, this power being confined to the tendrils. The internodes, however, of Passiflora gracilis have the power in a perfect manner, and those of the common Vine in an imperfect degree: so that at least a trace of the supposed primordial habit has been retained by some members of all the larger tendril-bearing groups.
On the view here given, it may be asked, Why have the species which were aboriginally twiners been converted in so many groups into leaf-climbers or tendril-bearers? Of what advantage has this been to them? Why did they not remain simple twiners? We can see several reasons. It might be an advantage to a plant to acquire a thicker stem, with short internodes bearing many or large leaves; and such stems are ill fitted for twining. Any one who will look during windy weather at twining plants will see that they are easily blown from their support; not so with tendril-bearers or leaf-climbers, for they quickly and firmly grasp their support by a much more efficient kind of movement. In those plants which still twine, but at the same time possess tendrils or sensitive petioles, as some species of Bignonia, Clematis, and Tropaeolum, it can readily be observed how incomparably better they grasp an upright stick than do simple twiners. Tendrils, from possessing this power of grasping an object, can be made long and thin; so that little organic matter is expended in their development, and yet they sweep a wide circle in search of a support. Tendril-bearers can, from their first growth, ascend along the outer branches of any neighbouring bush, and they are thus always fully exposed to the light; twiners, on the contrary, are best fitted to ascend bare stems, and generally have to start in the shade. Within tall and dense tropical forests, twining plants would probably succeed better than most kinds of tendril-bearers; but the majority of twiners, at least in our temperate regions, from the nature of their revolving movement, cannot ascend thick trunks, whereas this can be affected by tendril-bearers if the trunks are branched or bear twigs, and by some species if the bark is rugged.
The advantage gained by climbing is to reach the light and free air with as little expenditure of organic matter as possible; now, with twining plants, the stem is much longer than is absolutely necessary; for instance, I measured the stem of a kidney-bean, which had ascended exactly two feet in height, and it was three feet in length: the stem of a pea, on the other hand, which had ascended to the same height by the aid of its tendrils, was but little longer than the height reached. That this saving of the stem is really an advantage to climbing plants, I infer from the species that still twine but are aided by clasping petioles or tendrils, generally making more open spires than those made by simple twiners. Moreover, the plants thus aided, after taking one or two turns in one direction, generally ascend for a space straight, and then reverse the direction of their spire. By this means they ascend to a considerably greater height, with the same length of stem, than would otherwise have been possible; and they do this with safety, as they secure themselves at intervals by their clasping petioles or tendrils.
We have seen that tendrils consist of various organs in a modified state, namely, leaves, flower-peduncles, branches, and perhaps stipules. With respect to leaves, the evidence of their modification is ample. In young plants of Bignonia the lower leaves often remain quite unchanged, whilst the upper ones have their terminal leaflets converted into perfect tendrils; in Eccremocarpus I have seen a single lateral branch of a tendril replaced by a perfect leaflet; in Vicia sativa, on the other hand, leaflets are sometimes replaced by tendril-branches; and many other such cases could be given. But he who believes in the slow modification of species will not be content simply to ascertain the homological nature of different kinds of tendrils; he will wish to learn, as far as is possible, by what actual steps leaves, flower-peduncles, &c., have had their functions wholly changed, and have come to serve merely as prehensile organs.
In the whole group of leaf-climbers abundant evidence has been given that an organ, still subserving the functions of a leaf, may become sensitive to a touch, and thus grasp an adjoining object. With several leaf-climbers the true leaves spontaneously revolve; and their petioles, after clasping a support grow thicker and stronger. We thus see that leaves may acquire all the leading and characteristic qualities of tendrils, namely, sensitiveness, spontaneous movement, and subsequently increased strength. If their blades or laminae were to abort, they would form true tendrils. And of this process of abortion we can follow every step, until no trace of the original nature of the tendril is left. In Mutisia clematis, the tendril, in shape and colour, closely resembles the petiole of one of the ordinary leaves, together with the midribs of the leaflets, but vestiges of the laminae are still occasionally retained. In four genera of the Fumariaceae we can follow the whole process of transformation. The terminal leaflets of the leaf-climbing Fumaria officinalis are not smaller than the other leaflets; those of the leaf-cli............