The Movements and Habits of Climbing Plants

by Charles Darwin

Clematis calycina.—The young shoots are thin and flexible: one revolved, describing a broad oval, in 5 hrs. 30 m., and another in 6 hrs. 12 m. They followed the course of the sun; but the course, if observed long enough, would probably be found to vary in this species, as well as in all the others of the genus. It is a rather better twiner than the two last species: the stem sometimes made two spiral turns round a thin stick, if free from twigs; it then ran straight up for a space, and reversing its course took one or two turns in an opposite direction. This reversal of the spire occurred in all the foregoing species. The leaves are so small compared with those of most of the other species, that the petioles at first seem ill-adapted for clasping. Nevertheless, the main service of the revolving movement is to bring them into contact with surrounding objects, which are slowly but securely seized. The young petioles, which alone are sensitive, have their ends bowed a little downwards, so as to be in a slight degree hooked; ultimately the whole leaf, if it catches nothing, becomes level. I gently rubbed with a thin twig the lower surfaces of two young petioles; and in 2 hrs. 30 m. they were slightly curved downwards; in 5 hrs., after being rubbed, the end of one was bent completely back, parallel to the basal portion; in 4 hrs. subsequently it became nearly straight again. To show how sensitive the young petioles are, I may mention that I just touched the under sides of two with a little water-colour, which when dry formed an excessively thin and minute crust; but this sufficed in 24 hrs. to cause both to bend downwards. Whilst the plant is young, each leaf consists of three divided leaflets, which barely have distinct petioles, and these are not sensitive; but when the plant is well grown, the petioles of the two lateral and terminal leaflets are of considerable length, and become sensitive so as to be capable of clasping an object in any direction.

When a petiole has clasped a twig, it undergoes some remarkable changes, which may be observed with the other species, but in a less strongly marked manner, and will here be described once for all. The clasped petiole in the course of two or three days swells greatly, and ultimately becomes nearly twice as thick as the opposite one which has clasped nothing. When thin transverse slices of the two are placed under the microscope their difference is conspicuous: the side of the petiole which has been in contact with the support, is formed of a layer of colourless cells with their longer axes directed from the centre, and these are very much larger than the corresponding cells in the opposite or unchanged petiole; the central cells, also, are in some degree enlarged, and the whole is much indurated. The exterior surface generally becomes bright red. But a far greater change takes place in the nature of the tissues than that which is visible: the petiole of the unclasped leaf is flexible and can be snapped easily, whereas the clasped one acquires an extraordinary degree of toughness and rigidity, so that considerable force is required to pull it into pieces. With this change, great durability is probably acquired; at least this is the case with the clasped petioles of Clematis vitalba. The meaning of these changes is obvious, namely, that the petioles may firmly and durably support the stem.

Clematis microphylla, var. leptophylla.—The long and thin internodes of this Australian species revolve sometimes in one direction and sometimes in an opposite one, describing long, narrow, irregular ellipses or large circles. Four revolutions were completed within five minutes of the same average rate of 1 hr. 51 m.; so that this species moves more quickly than the others of the genus. The shoots, when placed near a vertical stick, either twine round it, or clasp it with the basal portions of their petioles. The leaves whilst young are nearly of the same shape as those of C. viticella, and act in the same manner like a hook, as will be described under that species. But the leaflets are more divided, and each segment whilst young terminates in a hardish point, which is much curved downwards and inwards; so that the whole leaf readily catches hold of any neighbouring object. The petioles of the young terminal leaflets are acted on by loops of thread weighing ⅛th and even 0.0625th of a grain. The basal portion of the main petiole is much less sensitive, but will clasp a stick against which it presses.

The leaves, whilst young, are continually and spontaneously moving slowly. A bell-glass was placed over a shoot secured to a stick, and the movements of the leaves were traced on it during several days. A very irregular line was generally formed; but one day, in the course of eight hours and three quarters, the figure clearly represented three and a half irregular ellipses, the most perfect one of which was completed in 2 hrs. 35 m. The two opposite leaves moved independently of each other. This movement of the leaves would aid that of the internodes in bringing the petioles into contact with surrounding objects. I discovered this movement too late to be enabled to observe it in the other species; but from analogy I can hardly doubt that the leaves of at least C. viticella, C. flammula, and C. vitalba move spontaneously; and, judging from C. Sieboldi, this probably is the case with C. montana and C. calycina. I ascertained that the simple leaves of C. glandulosa exhibited no spontaneous revolving movement.

Clematis viticella, var. venosa.—In this and the two following species the power of spirally twining is completely lost, and this seems due to the lessened flexibility of the internodes and to the interference caused by the large size of the leaves. But the revolving movement, though restricted, is not lost. In our present species a young internode, placed in front of a window, made three narrow ellipses, transversely to the direction of the light, at an average rate of 2 hrs. 40 m. When placed so that the movements were to and from the light, the rate was greatly accelerated in one half of the course, and retarded in the other, as with twining plants. The ellipses were small; the longer diameter, described by the apex of a shoot bearing a pair of not expanded leaves, was only 4⅝ inches, and that by the apex of the penultimate internode only 1⅛ inch. At the most favourable period of growth each leaf would hardly be carried to and fro by the movement of the internodes more than two or three inches, but, as above stated, it is probable that the leaves themselves move spontaneously. The movement of the whole shoot by the wind and by its rapid growth, would probably be almost equally efficient as these spontaneous movements, in bringing the petioles into contact with surrounding objects.

The leaves are of large size. Each bears three pairs of lateral leaflets and a terminal one, all supported on rather long sub-petioles. The main petiole bends a little angularly downwards at each point where a pair of leaflets arises (see fig. 2), and the petiole of the terminal leaflet is bent downwards at right angles; hence the whole petiole, with its rectangularly bent extremity, acts as a hook. This hook, the lateral petioles being directed a little upwards; forms an excellent grappling apparatus, by which the leaves readily become entangled with surrounding objects. If they catch nothing, the whole petiole ultimately grows straight. The main petiole, the sub-petioles, and the three branches into which each basi-lateral sub-petiole is generally subdivided, are all sensitive. The basal portion of the main petiole, between the stem and the first pair of leaflets, is less sensitive than the remainder; it will, however, clasp a stick with which it is left in contact. The inferior surface of the rectangularly bent terminal portion (carrying the terminal leaflet), which forms the inner side of the end of the hook, is the most sensitive part; and this portion is manifestly best adapted to catch a distant support. To show the difference in sensibility, I gently placed loops of string of the same weight (in one instance weighing only 0.82 of a grain or 53.14 mg.) on the several lateral sub-petioles and on the terminal one; in a few hours the latter was bent, but after 24 hrs. no effect was produced on the other sub-petioles. Again, a terminal sub-petiole placed in contact with a thin stick became sensibly curved in 45 m., and in 1 hr. 10 m. moved through ninety degrees; whilst a lateral sub-petiole did not become sensibly curved until 3 hrs. 30 m. had elapsed. In all cases, if the sticks are taken away, the petioles continue to move during many hours afterwards; so they do after a slight rubbing; but they become straight again, after about a day’s interval, that is if the flexure has not been very great or long continued.

The graduated difference in the extension of the sensitiveness in the petioles of the above-described species deserves notice. In C. montana it is confined to the main petiole, and has not spread to the sub-petioles of the three leaflets; so it is with young plants of C. calycina, but in older plants it spreads to the three sub-petioles. In C. viticella the sensitiveness has spread to the petioles of the seven leaflets, and to the subdivisions of the basi-lateral sub-petioles. But in this latter species it has diminished in the basal part of the main petiole, in which alone it resided in C. montana; whilst it has increased in the abruptly bent terminal portion.

Clematis flammula.—The rather thick, straight, and stiff shoots, whilst growing vigorously in the spring, make small oval revolutions, following the sun in their course. Four were made at an average rate of 3 hrs. 45 m. The longer axis of the oval, described by the extreme tip, was directed at right angles to the line joining the opposite leaves; its length was in one case only 1⅜, and in another case 1¾ inch; so that the young leaves were moved a very short distance. The shoots of the same plant observed in midsummer, when growing not so quickly, did not revolve at all. I cut down another plant in the early summer, so that by August 1st it had formed new and moderately vigorous shoots; these, when observed under a bell-glass, were on some days quite stationary, and on other days moved to and fro only about the eighth of an inch. Consequently the revolving power is much enfeebled in this species, and under unfavourable circumstances is completely lost. The shoot must depend for coming into contact with surrounding objects on the probable, though not ascertained spontaneous movement of the leaves, on rapid growth, and on movement from the wind. Hence, perhaps, it is that the petioles have acquired a high degree of sensitiveness as a compensation for the little power of movement in the shoots.

The petioles are bowed downwards, and have the same general hook-like form as in C. viticella. The medial petiole and the lateral sub-petioles are sensitive, especially the much bent terminal portion. As the sensitiveness is here greater than in any other species of the genus observed by me, and is in itself remarkable, I will give fuller details. The petioles, when so young that they have not separated from one another, are not sensitive; when the lamina of a leaflet has grown to a quarter of an inch in length (that is, about one-sixth of its full size), the sensitiveness is highest; but at this period the petioles are relatively much more fully developed than are the blades of the leaves. Full-grown petioles are not in the least sensitive. A thin stick placed so as to press lightly against a petiole, having a leaflet a quarter of an inch in length, caused the petiole to bend in 3 hrs. 15 m. In another case a petiole curled completely round a stick in 12 hrs. These petioles were left curled for 24 hrs., and the sticks were then removed; but they never straightened themselves. I took a twig, thinner than the petiole itself, and with it lightly rubbed several petioles four times up and down; these in 1 hr. 45 m. became slightly curled; the curvature increased during some hours and then began to decrease, but after 25 hrs. from the time of rubbing a vestige of the curvature remained. Some other petioles similarly rubbed twice, that is, once up and once down, became perceptibly curved in about 2 hrs. 30 m., the terminal sub-petiole moving more than the lateral sub-petioles; they all became straight again in between 12 hrs. and 14 hrs. Lastly, a length of about one-eighth of an inch of a sub-petiole, was lightly rubbed with the same twig only once; it became slightly curved in 3 hrs., remaining so during 11 hrs., but by the next morning was quite straight.