The Movements and Habits of Climbing Plants, by Charles Darwin

Chapter II. — Leaf-Climbers.

Plants which climb by the aid of spontaneously revolving and sensitive petioles — Clematis — Tropaeolum — Maurandia, flower-peduncles moving spontaneously and sensitive to a touch — Rhodochiton — Lophospermum — internodes sensitive — Solanum, thickening of the clasped petioles — Fumaria — Adlumia — Plants which climb by the aid of their produced midribs — Gloriosa — Flagellaria — Nepenthes — Summary on leaf-climbers.

We now come to our second class of climbing plants, namely, those which ascend by the aid of irritable or sensitive organs. For convenience’ sake the plants in this class have been grouped under two sub-divisions, namely, leaf-climbers, or those which retain their leaves in a functional condition, and tendril-bearers. But these sub-divisions graduate into each other, as we shall see under Corydalis and the Gloriosa lily.

It has long been observed that several plants climb by the aid of their leaves, either by their petioles (foot-stalks) or by their produced midribs; but beyond this simple fact they have not been described. Palm and Mohl class these plants with those which bear tendrils; but as a leaf is generally a defined object, the present classification, though artificial, has at least some advantages. Leaf-climbers are, moreover, intermediate in many respects between twiners and tendril-bearers. Eight species of Clematis and seven of Tropaeolum were observed, in order to see what amount of difference in the manner of climbing existed within the same genus; and the differences are considerable.

CLEMATIS. — C. glandulosa. — The thin upper internodes revolve, moving against the course of the sun, precisely like those of a true twiner, at an average rate, judging from three revolutions, of 3 hrs. 48 m. The leading shoot immediately twined round a stick placed near it; but, after making an open spire of only one turn and a half, it ascended for a short space straight, and then reversed its course and wound two turns in an opposite direction. This was rendered possible by the straight piece between the opposed spires having become rigid. The simple, broad, ovate leaves of this tropical species, with their short thick petioles, seem but ill-fitted for any movement; and whilst twining up a vertical stick, no use is made of them. Nevertheless, if the footstalk of a young leaf be rubbed with a thin twig a few times on any side, it will in the course of a few hours bend to that side; afterwards becoming straight again. The under side seemed to be the most sensitive; but the sensitiveness or irritability is slight compared to that which we shall meet with in some of the following species; thus, a loop of string, weighing 1.64 grain (106.2 mg.) and hanging for some days on a young footstalk, produced a scarcely perceptible effect. A sketch is here given of two young leaves which had naturally caught hold of two thin branches. A forked twig placed so as to press lightly on the under side of a young footstalk caused it, in 12 hrs., to bend greatly, and ultimately to such an extent that the leaf passed to the opposite side of the stem; the forked stick having been removed, the leaf slowly recovered its former position.

The young leaves spontaneously and gradually change their position: when first developed the petioles are upturned and parallel to the stem; they then slowly bend downwards, remaining for a short time at right angles to the stem, and then become so much arched downwards that the blade of the leaf points to the ground with its tip curled inwards, so that the whole petiole and leaf together form a hook. They are thus enabled to catch hold of any twig with which they may be brought into contact by the revolving movement of the internodes. If this does not happen, they retain their hooked shape for a considerable time, and then bending upwards reassume their original upturned position, which is preserved ever afterwards. The petioles which have clasped any object soon become much thickened and strengthened, as may be seen in the drawing.

Clematis montana. — The long, thin petioles of the leaves, whilst young, are sensitive, and when lightly rubbed bend to the rubbed side, subsequently becoming straight. They are far more sensitive than the petioles of C. glandulosa; for a loop of thread weighing a quarter of a grain (16.2 mg.) caused them to bend; a loop weighing only one-eighth of a grain (8.1 mg.) sometimes acted and sometimes did not act. The sensitiveness extends from the blade of the leaf to the stem. I may here state that I ascertained in all cases the weights of the string and thread used by carefully weighing 50 inches in a chemical balance, and then cutting off measured lengths. The main petiole carries three leaflets; but their short, sub-petioles are not sensitive. A young, inclined shoot (the plant being in the greenhouse) made a large circle opposed to the course of the sun in 4 hrs. 20 m., but the next day, being very cold, the time was 5 hrs. 10 m. A stick placed near a revolving stem was soon struck by the petioles which stand out at right angles, and the revolving movement was thus arrested. The petioles then began, being excited by the contact, to slowly wind round the stick. When the stick was thin, a petiole sometimes wound twice round it. The opposite leaf was in no way affected. The attitude assumed by the stem after the petiole had clasped the stick, was that of a man standing by a column, who throws his arm horizontally round it. With respect to the stem’s power of twining, some remarks will be made under C. calycina.

Clematis Sieboldi. — A shoot made three revolutions against the sun at an average rate of 3 hrs. 11 m. The power of twining is like that of the last species. Its leaves are nearly similar in structure and in function, excepting that the sub-petioles of the lateral and terminal leaflets are sensitive. A loop of thread, weighing one-eighth of a grain, acted on the main petiole, but not until two or three days had elapsed. The leaves have the remarkable habit of spontaneously revolving, generally in vertical ellipses, in the same manner, but in a less degree, as will be described under C. microphylla.

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 0.125th 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 4.625 inches, and that by the apex of the penultimate internode only 1.125 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. 10m. 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 1.375, and in another case 1.75 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.

The following observations are more precise. After trying heavier pieces of string and thread, I placed a loop of fine string, weighing 1.04 gr. (67.4 mg.) on a terminal sub-petiole: in 6 hrs. 40 m. a curvature could be seen; in 24 hrs. the petiole formed an open ring round the string; in 48 hrs. the ring had almost closed on the string, and in 72 hrs. seized it so firmly, that some force was necessary for its withdrawal. A loop weighing 0.52 of a grain (33.7 mg.) caused in 14 hrs. a lateral sub-petiole just perceptibly to curve, and in 24 hrs. it moved through ninety degrees. These observations were made during the summer: the following were made in the spring, when the petioles apparently are more sensitive:— A loop of thread, weighing one-eighth of a grain (8.1 mg.), produced no effect on the lateral sub-petioles, but placed on a terminal one, caused it, after 24 hrs., to curve moderately; the curvature, though the loop remained suspended, was after 48 hrs. diminished, but never disappeared; showing that the petiole had become partially accustomed to the insufficient stimulus. This experiment was twice repeated with nearly the same result. Lastly, a loop of thread, weighing only one-sixteenth of a grain (4.05 mg.) was twice gently placed by a forceps on a terminal sub-petiole (the plant being, of course, in a still and closed room), and this weight certainly caused a flexure, which very slowly increased until the petiole moved through nearly ninety degrees: beyond this it did not move; nor did the petiole, the loop remaining suspended, ever become perfectly straight again.

When we consider, on the one hand, the thickness and stiffness of the petioles, and, on the other hand, the thinness and softness of fine cotton thread, and what an extremely small weight one-sixteenth of a grain (4.05 mg.) is, these facts are remarkable. But I have reason to believe that even a less weight excites curvature when pressing over a broader surface than that acted on by a thread. Having noticed that the end of a suspended string which accidentally touched a petiole, caused it to bend, I took two pieces of thin twine, 10 inches in length (weighing 1.64 gr.), and, tying them to a stick, let them hang as nearly perpendicularly downwards as their thinness and flexuous form, after being stretched, would permit; I then quietly placed their ends so as just to rest on two petioles, and these certainly became curved in 36 hrs. One of the ends touched the angle between a terminal and lateral sub-petiole, and it was in 48 hours caught between them as by a forceps. In these cases the pressure, though spread over a wider surface than that touched by the cotton thread, must have been excessively slight.

Clematis vitalba. — The plants were in pots and not healthy, so that I dare not trust my observations, which indicate much similarity in habits with C. flammula. I mention this species only because I have seen many proofs that the petioles in a state of nature are excited to movement by very slight pressure. For instance, I have found them embracing thin withered blades of grass, the soft young leaves of a maple, and the flower-peduncles of the quaking-grass or Briza. The latter are about as thick as the hair of a man’s beard, but they were completely surrounded and clasped. The petioles of a leaf, so young that none of the leaflets were expanded, had partially seized a twig. Those of almost all the old leaves, even when unattached to any object, are much convoluted; but this is owing to their having come, whilst young, into contact during several hours with some object subsequently removed. With none of the above-described species, cultivated in pots and carefully observed, was there any permanent bending of the petioles without the stimulus of contact. In winter, the blades of the leaves of C. vitalba drop off; but the petioles (as was observed by Mohl) remain attached to the branches, sometimes during two seasons; and, being convoluted, they curiously resemble true tendrils, such as those possessed by the allied genus Naravelia. The petioles which have clasped some object become much more stiff, hard, and polished than those which have failed in this their proper function.

TROPAEOLUM. — I observed T. tricolorum, T. azureum, T. pentaphyllum, T. peregrinum, T. elegans, T. tuberosum, and a dwarf variety of, as I believe, T. minus.

Tropaeolum tricolorum, var. grandiflorum. — The flexible shoots, which first rise from the tubers, are as thin as fine twine. One such shoot revolved in a course opposed to the sun, at an average rate, judging from three revolutions, of 1 hr. 23 m.; but no doubt the direction of the revolving movement is variable. When the plants have grown tall and are branched, all the many lateral shoots revolve. The stem, whilst young, twines regularly round a thin vertical stick, and in one case I counted eight spiral turns in the same direction; but when grown older, the stem often runs straight up for a space, and, being arrested by the clasping petioles, makes one or two spires in a reversed direction. Until the plant grows to a height of two or three feet, requiring about a month from the time when the first shoot appears above ground, no true leaves are produced, but, in their place, filaments coloured like the stem. The extremities of these filaments are pointed, a little flattened, and furrowed on the upper surface. They never become developed into leaves. As the plant grows in height new filaments are produced with slightly enlarged tips; then others, bearing on each side of the enlarged medial tip a rudimentary segment of a leaf; soon other segments appear, and at last a perfect leaf is formed, with seven deep segments. So that on the same plant we may see every step, from tendril-like clasping filaments to perfect leaves with clasping petioles. After the plant has grown to a considerable height, and is secured to its support by the petioles of the true leaves, the clasping filaments on the lower part of the stem wither and drop off; so that they perform only a temporary service.

These filaments or rudimentary leaves, as well as the petioles of the perfect leaves, whilst young, are highly sensitive on all sides to a touch. The slightest rub caused them to curve towards the rubbed side in about three minutes, and one bent itself into a ring in six minutes; they subsequently became straight. When, however, they have once completely clasped a stick, if this is removed, they do not straighten themselves. The most remarkable fact, and one which I have observed in no other species of the genus, is that the filaments and the petioles of the young leaves, if they catch no object, after standing for some days in their original position, spontaneously and slowly oscillate a little from side to side, and then move towards the stem and clasp it. They likewise often become, after a time, in some degree spirally contracted. They therefore fully deserve to be called tendrils, as they are used for climbing, are sensitive to a touch, move spontaneously, and ultimately contract into a spire, though an imperfect one. The present species would have been classed amongst the tendril-bearers, had not these characters been confined to early youth. During maturity it is a true leaf-climber.

Tropaeolum azureum. — An upper internode made four revolutions, following the sun, at an average rate of 1 hr. 47 m. The stem twined spirally round a support in the same irregular manner as that of the last species. Rudimentary leaves or filaments do not exist. The petioles of the young leaves are very sensitive: a single light rub with a twig caused one to move perceptibly in 5 m., and another in 6 m. The former became bent at right angles in 15 min., and became straight again in between 5 hrs. and 6 hrs. A loop of thread weighing 0.125th of a grain caused another petiole to curve.

Tropaeolum pentaphyllum. — This species has not the power of spirally twining, which seems due, not so much to a want of flexibility in the stem, as to continual interference from the clasping petioles. An upper internode made three revolutions, following the sun, at an average rate of 1 hr. 46 m. The main purpose of the revolving movement in all the species of Tropaeolum manifestly is to bring the petioles into contact with some supporting object. The petiole of a young leaf, after a slight rub, became curved in 6 m.; another, on a cold day, in 20 m., and others in from 8 m. to 10 m. Their curvature usually increased greatly in from 15 m. to 20 m., and they became straight again in between 5 hrs. and 6 hrs., but on one occasion in 3 hrs. When a petiole has fairly clasped a stick, it is not able, on the removal of the stick, to straighten itself. The free upper part of one, the base of which had already clasped a stick, still retained the power of movement. A loop of thread weighing 0.125th of a grain caused a petiole to curve; but the stimulus was not sufficient, the loop remaining suspended, to cause a permanent flexure. If a much heavier loop be placed in the angle between the petiole and the stem, it produces no effect; whereas we have seen with Clematis montana that the angle between the stem and petiole is sensitive.

Tropaeolum peregrinum. — The first-formed internodes of a young plant did not revolve, resembling in this respect those of a twining plant. In an older plant the four upper internodes made three irregular revolutions, in a course opposed to the sun, at an average rate of 1 hr. 48 min. It is remarkable that the average rate of revolution (taken, however, but from few observations) is very nearly the same in this and the two last species, namely, 1 hr. 47 m., 1 hr. 46 m., and 1 hr. 48 m. The present species cannot twine spirally, which seems mainly due to the rigidity of the stem. In a very young plant, which did not revolve, the petioles were not sensitive. In older plants the petioles of quite young leaves, and of leaves as much as an inch and a quarter in diameter, are sensitive. A moderate rub caused one to curve in 10 m., and others in 20 m. They became straight again in between 5 hrs. 45m. and 8 hrs. Petioles which have naturally come into contact with a stick, sometimes take two turns round it. After they have clasped a support, they become rigid and hard. They are less sensitive to a weight than in the previous species; for loops of string weighing 0.82 of a grain (53.14 mg.), did not cause any curvature, but a loop of double this weight (1.64 gr.) acted.

Tropaeolum elegans. — I did not make many observations on this species. The short and stiff internodes revolve irregularly, describing small oval figures. One oval was completed in 3 hrs. A young petiole, when rubbed, became slightly curved in 17 m.; and afterwards much more so. It was nearly straight again in 8 hrs.

Tropaeolum tuberosum. — On a plant nine inches in height, the internodes did not move at all; but on an older plant they moved irregularly and made small imperfect ovals. These movements could be detected only by being traced on a bell-glass placed over the plant. Sometimes the shoots stood still for hours; during some days they moved only in one direction in a crooked line; on other days they made small irregular spires or circles, one being completed in about 4 hrs. The extreme points reached by the apex of the shoot were only about one or one and a half inches asunder; yet this slight movement brought the petioles into contact with some closely surrounding twigs, which were then clasped. With the lessened power of spontaneously revolving, compared with that of the previous species, the sensitiveness of the petioles is also diminished. These, when rubbed a few times, did not become curved until half an hour had elapsed; the curvature increased during the next two hours, and then very slowly decreased; so that they sometimes required 24 hrs. to become straight again. Extremely young leaves have active petioles; one with the lamina only 0.15 of an inch in diameter, that is, about a twentieth of the full size, firmly clasped a thin twig. But leaves grown to a quarter of their full size can likewise act.

Tropaeolum minus (?). — The internodes of a variety named “dwarf crimson Nasturtium” did not revolve, but moved in a rather irregular course during the day to the light, and from the light at night. The petioles, when well rubbed, showed no power of curving; nor could I see that they ever clasped any neighbouring object. We have seen in this genus a gradation from species such as T. tricolorum, which have extremely sensitive petioles, and internodes which rapidly revolve and spirally twine up a support, to other species such as T. elegans and T. tuberosum, the petioles of which are much less sensitive, and the internodes of which have very feeble revolving powers and cannot spirally twine round a support, to this last species, which has entirely lost or never acquired these faculties. From the general character of the genus, the loss of power seems the more probable alternative.

In the present species, in T. elegans, and probably in others, the flower-peduncle, as soon as the seed-capsule begins to swell, spontaneously bends abruptly downwards and becomes somewhat convoluted. If a stick stands in the way, it is to a certain extent clasped; but, as far as I have been able to observe, this clasping movement is independent of the stimulus from contact.

ANTIRRHINEAE. — In this tribe (Lindley) of the Scrophulariaceae, at least four of the seven included genera have leaf-climbing species.

Maurandia Barclayana. — A thin, slightly bowed shoot made two revolutions, following the sun, each in 3 hrs. 17 min.; on the previous day this same shoot revolved in an opposite direction. The shoots do not twine spirally, but climb excellently by the aid of their young and sensitive petioles. These petioles, when lightly rubbed, move after a considerable interval of time, and subsequently become straight again. A loop of thread weighing 0.125th of a grain caused them to bend.

Maurandia semperflorens. — This freely growing species climbs exactly like the last, by the aid of its sensitive petioles. A young internode made two circles, each in 1 hr. 46 mm.; so that it moved almost twice as rapidly as the last species. The internodes are not in the least sensitive to a touch or pressure. I mention this because they are sensitive in a closely allied genus, namely, Lophospermum. The present species is unique in one respect. Mohl asserts (p. 45) that “the flower-peduncles, as well as the petioles, wind like tendrils;” but he classes as tendrils such objects as the spiral flower-stalks of the Vallisneria. This remark, and the fact of the flower-peduncles being decidedly flexuous, led me carefully to examine them. They never act as true tendrils; I repeatedly placed thin sticks in contact with young and old peduncles, and I allowed nine vigorous plants to grow through an entangled mass of branches; but in no one instance did they bend round any object. It is indeed in the highest degree improbable that this should occur, for they are generally developed on branches which have already securely clasped a support by the petioles of their leaves; and when borne on a free depending branch, they are not produced by the terminal portion of the internode which alone has the power of revolving; so that they could be brought only by accident into contact with any neighbouring object. Nevertheless (and this is the remarkable fact) the flower-peduncles, whilst young, exhibit feeble revolving powers, and are slightly sensitive to a touch. Having selected some stems which had firmly clasped a stick by their petioles, and having placed a bell-glass over them, I traced the movements of the young flower — peduncles. The tracing generally formed a short and extremely irregular line, with little loops in its course. A young peduncle 1.5 inch in length was carefully observed during a whole day, and it made four and a half narrow, vertical, irregular, and short ellipses — each at an average rate of about 2 hrs. 25 m. An adjoining peduncle described during the same time similar, though fewer, ellipses. As the plant had occupied for some time exactly the same position, these movements could not be attributed to any change in the action of the light. Peduncles, old enough for the coloured petals to be just visible, do not move. With respect to irritability, 21 I rubbed two young peduncles (1.5 inch in length) a few times very lightly with a thin twig; one was rubbed on the upper, and the other on the lower side, and they became in between 4 hrs. and 5 hrs. distinctly bowed towards these sides; in 24 hrs. subsequently, they straightened themselves. Next day they were rubbed on the opposite sides, and they became perceptibly curved towards these sides. Two other and younger peduncles (three-fourths of an inch in length) were lightly rubbed on their adjoining sides, and they became so much curved towards one another, that the arcs of the bows stood at nearly right angles to their previous direction; and this was the greatest movement seen by me. Subsequently they straightened themselves. Other peduncles, so young as to be only three-tenths of an inch in length, became curved when rubbed. On the other hand, peduncles above 1.5 inch in length required to be rubbed two or three times, and then became only just perceptibly bowed. Loops of thread suspended on the peduncles produced no effect; loops of string, however, weighing 0.82 and 1.64 of a grain sometimes caused a slight curvature; but they were never closely clasped, as were the far lighter loops of thread by the petioles.

In the nine vigorous plants observed by me, it is certain that neither the slight spontaneous movements nor the slight sensitiveness of the flower-peduncles aided the plants in climbing. If any member of the Scrophulariaceae had possessed tendrils produced by the modification of flower-peduncles, I should have thought that this species of Maurandia had perhaps retained a useless or rudimentary vestige of a former habit; but this view cannot be maintained. We may suspect that, owing to the principle of correlation, the power of movement has been transferred to the flower-peduncles from the young internodes, and sensitiveness from the young petioles. But to whatever cause these capacities are due, the case is interesting; for, by a little increase in power through natural selection, they might easily have been rendered as useful to the plant in climbing, as are the flower-peduncles (hereafter to be described) of Vitis or Cardiospermum.

Rhodochiton volubile. — A long flexible shoot swept a large circle, following the sun, in 5 hrs. 30 m.; and, as the day became warmer, a second circle was completed in 4 hrs. 10 m. The shoots sometimes make a whole or a half spire round a vertical stick, they then run straight up for a space, and afterwards turn spirally in an opposite direction. The petioles of very young leaves about one-tenth of their full size, are highly sensitive, and bend towards the side which is touched; but they do not move quickly. One was perceptibly curved in 1 hr. 10 m., after being lightly rubbed, and became considerably curved in 5 hrs. 40 m.; some others were scarcely curved in 5 hrs. 30 m., but distinctly so in 6 hrs. 30 m. A curvature was perceptible in one petiole in between 4 hrs. 30 m. and 5 hrs., after the suspension of a little loop of string. A loop of fine cotton thread, weighing one sixteenth of a grain (4.05 mg.), not only caused a petiole slowly to bend, but was ultimately so firmly clasped that it could be withdrawn only by some little force. The petioles, when coming into contact with a stick, take either a complete or half a turn round it, and ultimately increase much in thickness. They do not possess the power of spontaneously revolving.

Lophospermum scandens, var. purpureum. — Some long, moderately thin internodes made four revolutions at an average rate of 3 hrs. 15 m. The course pursued was very irregular, namely, an extremely narrow ellipse, a large circle, an irregular spire or a zigzag line, and sometimes the apex stood still. The young petioles, when brought by the revolving movement into contact with sticks, clasped them, and soon increased considerably in thickness. But they are not quite so sensitive to a weight as those of the Rhodochiton, for loops of thread weighing one-eighth of a grain did not always cause them to bend.

This plant presents a case not observed by me in any other leaf-climber or twiner, 22 namely, that the young internodes of the stem are sensitive to a touch. When a petiole of this species clasps a stick, it draws the base of the internode against it; and then the internode itself bends towards the stick, which is caught between the stem and the petiole as by a pair of pincers. The internode afterwards straightens itself, excepting the part in actual contact with the stick. Young internodes alone are sensitive, and these are sensitive on all sides along their whole length. I made fifteen trials by twice or thrice lightly rubbing with a thin twig several internodes; and in about 2 hrs., but in one case in 3 hrs., all were bent: they became straight again in about 4 hrs. afterwards. An internode, which was rubbed as often as six or seven times, became just perceptibly curved in 1 hr. 15 m., and in 3 hrs. the curvature increased much; it became straight again in the course of the succeeding night. I rubbed some internodes one day on one side, and the next day either on the opposite side or at right angles to the first side; and the curvature was always towards the rubbed side.

According to Palm (p. 63), the petioles of Linaria cirrhosa and, to a limited degree, those of L. elatine have the power of clasping a support.

SOLANACEAE. — Solanum jasminoides. — Some of the species in this large genus are twiners; but the present species is a true leaf-climber. A long, nearly upright shoot made four revolutions, moving against the sun, very regularly at an average rate of 3 hrs. 26 m. The shoots, however, sometimes stood still. It is considered a greenhouse plant; but when kept there, the petioles took several days to clasp a stick: in the hothouse a stick was clasped in 7 hrs. In the greenhouse a petiole was not affected by a loop of string, suspended during several days and weighing 2.5 grains (163 mg.); but in the hothouse one was made to curve by a loop weighing 1.64 gr. (106.27 mg.); and, on the removal of the string, it became straight again. Another petiole was not at all acted on by a loop weighing only 0.82 of a grain (53.14 mg.) We have seen that the petioles of some other leaf-climbing plants are affected by one-thirteenth of this latter weight. In this species, and in no other leaf-climber seen by me, a full-grown leaf is capable of clasping a stick; but in the greenhouse the movement was so extraordinarily slow that the act required several weeks; on each succeeding week it was clear that the petiole had become more and more curved, until at last it firmly clasped the stick.

The flexible petiole of a half or a quarter grown leaf which has clasped an object for three or four days increases much in thickness, and after several weeks becomes so wonderfully hard and rigid that it can hardly be removed from its support. On comparing a thin transverse slice of such a petiole with one from an older leaf growing close beneath, which had not clasped anything, its diameter was found to be fully doubled, and its structure greatly changed. In two other petioles similarly compared, and here represented, the increase in diameter was not quite so great. In the section of the petiole in its ordinary state (A), we see a semilunar band of cellular tissue (not well shown in the woodcut) differing slightly in appearance from that outside it, and including three closely approximate groups of dark vessels. Near the upper surface of the petiole, beneath two exterior ridges, there are two other small circular groups of vessels. In the section of the petiole (B) which had clasped during several weeks a stick, the two exterior ridges have become much less prominent, and the two groups of woody vessels beneath them much increased in diameter. The semilunar band has been converted into a complete ring of very hard, white, woody tissue, with lines radiating from the centre. The three groups of vessels, which, though near together, were before distinct, are now completely blended. The upper part of this ring of woody vessels, formed by the prolongation of the horns of the original semilunar band, is narrower than the lower part, and slightly less compact. This petiole after clasping the stick had actually become thicker than the stem from which it arose; and this was chiefly due to the increased thickness of the ring of wood. This ring presented, both in a transverse and longitudinal section, a closely similar structure to that of the stem. It is a singular morphological fact that the petiole should thus acquire a structure almost identically the same with that of the axis; and it is a still more singular physiological fact that so great a change should have been induced by the mere act of clasping a support. 23

FUMARIACEAE. — Fumaria officinalis. — It could not have been anticipated that so lowly a plant as this Fumaria should have been a climber. It climbs by the aid of the main and lateral petioles of its compound leaves; and even the much-flattened terminal portion of the petiole can seize a support. I have seen a substance as soft as a withered blade of grass caught. Petioles which have clasped any object ultimately become rather thicker and more cylindrical. On lightly rubbing several petioles with a twig, they became perceptibly curved in 1 hr. 15 m., and subsequently straightened themselves. A stick gently placed in the angle between two sub-petioles excited them to move, and was almost clasped in 9 hrs. A loop of thread, weighing one-eighth of a grain, caused, after 12 hrs. and before 20 hrs, had elapsed, a considerable curvature; but it was never fairly clasped by the petiole. The young internodes are in continual movement, which is considerable in extent, but very irregular; a zigzag line, or a spire crossing itself; or a figure of 8 being formed. The course during 12 hrs., when traced on a bell-glass, apparently represented about four ellipses. The leaves themselves likewise move spontaneously, the main petioles curving themselves in accordance with the movements of the internodes; so that when the latter moved to one side, the petioles moved to the same side, then, becoming straight, reversed their curvature. The petioles, however, do not move over a wide space, as could be seen when a shoot was securely tied to a stick. The leaf in this case followed an irregular course, like that made by the internodes.

Adlumia cirrhosa. — I raised some plants late in the summer; they formed very fine leaves, but threw up no central stem. The first-formed leaves were not sensitive; some of the later ones were so, but only towards their extremities, which were thus enabled to clasp sticks. This could be of no service to the plant, as these leaves rose from the ground; but it showed what the future character of the plant would have been, had it grown tall enough to climb. The tip of one of these basal leaves, whilst young, described in 1 hr. 36 m. a narrow ellipse, open at one end, and exactly three inches in length; a second ellipse was broader, more irregular, and shorter, viz., only 2.5 inches in length, and was completed in 2 hrs. 2 m. From the analogy of Fumaria and Corydalis, I have no doubt that the internodes of Adlumia have the power of revolving.

Corydalis claviculata. — This plant is interesting from being in a condition so exactly intermediate between a leaf-climber and a tendril-bearer, that it might have been described under either head; but, for reasons hereafter assigned, it has been classed amongst tendril-bearers.

Besides the plants already described, Bignonia unguis and its close allies, though aided by tendrils, have clasping petioles. According to Mohl (p. 40), Cocculus Japonicus (one of the Menispermaceae) and a fern, the Ophioglossum Japonicum (p. 39), climb by their leaf-stalks.

We now come to a small section of plants which climb by means of the produced midribs or tips of their leaves.

LILIACEAE. — Gloriosa Plantii. — The stem of a half-grown plant continually moved, generally describing an irregular spire, but sometimes oval figures with the longer axes directed in different lines. It either followed the sun, or moved in an opposite course, and sometimes stood still before reversing its direction. One oval was completed in 3 hrs. 40 m.; of two horseshoe-shaped figures, one was completed in 4 hrs. 35 m. and the other in 3 hrs. The shoots, in their movements, reached points between four and five inches asunder. The young leaves, when first developed, stand up nearly vertically; but by the growth of the axis, and by the spontaneous bending down of the terminal half of the leaf, they soon become much inclined, and ultimately horizontal. The end of the leaf forms a narrow, ribbon-like, thickened projection, which at first is nearly straight, but by the time the leaf gets into an inclined position, the end bends downwards into a well-formed hook. This hook is now strong and rigid enough to catch any object, and, when caught, to anchor the plant and stop the revolving movement. Its inner surface is sensitive, but not in nearly so high a degree as that of the many before-described petioles; for a loop of string, weighing 1.64 grain, produced no effect. When the hook has caught a thin twig or even a rigid fibre, the point may be perceived in from 1 hr. to 3 hrs. to have curled a little inwards; and, under favourable circumstances, it curls round and permanently seizes an object in from 8 hrs. to 10 hrs. The hook when first formed, before the leaf has bent downwards, is but little sensitive. If it catches hold of nothing, it remains open and sensitive for a long time; ultimately the extremity spontaneously and slowly curls inwards, and makes a button-like, flat, spiral coil at the end of the leaf. One leaf was watched, and the hook remained open for thirty-three days; but during the last week the tip had curled so much inwards that only a very thin twig could have been inserted within it. As soon as the tip has curled so much inwards that the hook is converted into a ring, its sensibility is lost; but as long as it remains open some sensibility is retained.

Whilst the plant was only about six inches in height, the leaves, four or five in number, were broader than those subsequently produced; their soft and but little-attenuated tips were not sensitive, and did not form hooks; nor did the stem then revolve. At this early period of growth, the plant can support itself; its climbing powers are not required, and consequently are not developed. So again, the leaves on the summit of a full-grown flowering plant, which would not require to climb any higher, were not sensitive and could not clasp a stick. We thus see how perfect is the economy of nature.

COMMELYNACEAE. — Flagellaria Indica. — From dried specimens it is manifest that this plant climbs exactly like the Gloriosa. A young plant 12 inches in height, and bearing fifteen leaves, had not a single leaf as yet produced into a hook or tendril-like filament; nor did the stem revolve. Hence this plant acquires its climbing powers later in life than does the Gloriosa lily. According to Mohl (p. 41), Uvularia (Melanthaceae) also climbs like Gloriosa.

These three last-named genera are Monocotyledons; but there is one Dicotyledon, namely Nepenthes, which is ranked by Mohl (p. 41) amongst tendril-bearers; and I hear from Dr. Hooker that most of the species climb well at Kew. This is effected by the stalk or midrib between the leaf and the pitcher coiling round any support. The twisted part becomes thicker; but I observed in Mr. Veitch’s hothouse that the stalk often takes a turn when not in contact with any object, and that this twisted part is likewise thickened. Two vigorous young plants of N. laevis and N. distillatoria, in my hothouse, whilst less than a foot in height, showed no sensitiveness in their leaves, and had no power of climbing. But when N. laevis had grown to a height of 16 inches, there were signs of these powers. The young leaves when first formed stand upright, but soon become inclined; at this period they terminate in a stalk or filament, with the pitcher at the extremity hardly at all developed. The leaves now exhibited slight spontaneous movements; and when the terminal filaments came into contact with a stick, they slowly bent round and firmly seized it. But owing to the subsequent growth of the leaf, this filament became after a time quite slack, though still remaining firmly coiled round the stick. Hence it would appear that the chief use of the coiling, at least whilst the plant is young, is to support the pitcher with its load of secreted fluid.

Summary on Leaf-climbers. — Plants belonging to eight families are known to have clasping petioles, and plants belonging to four families climb by the tips of their leaves. In all the species observed by me, with one exception, the young internodes revolve more or less regularly, in some cases as regularly as those of a twining plant. They revolve at various rates, in most cases rather rapidly. Some few can ascend by spirally twining round a support. Differently from most twiners, there is a strong tendency in the same shoot to revolve first in one and then in an opposite direction. The object gained by the revolving movement is to bring the petioles or the tips of the leaves into contact with surrounding objects; and without this aid the plant would be much less successful in climbing. With rare exceptions, the petioles are sensitive only whilst young. They are sensitive on all sides, but in different degrees in different plants; and in some species of Clematis the several parts of the same petiole differ much in sensitiveness. The hooked tips of the leaves of the Gloriosa are sensitive only on their inner or inferior surfaces. The petioles are sensitive to a touch and to excessively slight continued pressure, even from a loop of soft thread weighing only the one-sixteenth of a grain (4.05 mg.); and there is reason to believe that the rather thick and stiff petioles of Clematis flammula are sensitive to even much less weight if spread over a wide surface. The petioles always bend towards the side which is pressed or touched, at different rates in different species, sometimes within a few minutes, but generally after a much longer period. After temporary contact with any object, the petiole continues to bend for a considerable time; afterwards it slowly becomes straight again, and can then re-act. A petiole excited by an extremely slight weight sometimes bends a little, and then becomes accustomed to the stimulus, and either bends no more or becomes straight again, the weight still remaining suspended. Petioles which have clasped an object for some little time cannot recover their original position. After remaining clasped for two or three days, they generally increase much in thickness either throughout their whole diameter or on one side alone; they subsequently become stronger and more woody, sometimes to a wonderful degree; and in some cases they acquire an internal structure like that of the stem or axis.

The young internodes of the Lophospermum as well as the petioles are sensitive to a touch, and by their combined movement seize an object. The flower-peduncles of the Maurandia semperflorens revolve spontaneously and are sensitive to a touch, yet are not used for climbing. The leaves of at least two, and probably of most, of the species of Clematis, of Fumaria and Adlumia, spontaneously curve from side to side, like the internodes, and are thus better adapted to seize distant objects. The petioles of the perfect leaves of Tropaeolum tricolorum, as well as the tendril-like filaments of the plants whilst young, ultimately move towards the stem or the supporting stick, which they then clasp. These petioles and filaments also show some tendency to contract spirally. The tips of the uncaught leaves of the Gloriosa, as they grow old, contract into a flat spire or helix. These several facts are interesting in relation to true tendrils.

With leaf climbers, as with twining plants, the first internodes which rise from the ground do not, at least in the cases observed by me, spontaneously revolve; nor are the petioles or tips of the first-formed leaves sensitive. In certain species of Clematis, the large size of the leaves, together with their habit of revolving, and the extreme sensitiveness of their petioles, appear to render the revolving movement of the internodes superfluous; and this latter power has consequently become much enfeebled. In certain species of Tropaeolum, both the spontaneous movements of the internodes and the sensitiveness of the petioles have become much enfeebled, and in one species have been completely lost.

21 It appears from A. Kerner’s interesting observations, that the flower-peduncles of a large number of plants are irritable, and bend when they are rubbed or shaken: Die Schutzmittel des Pollens, 1873, p. 34.

22 I have already referred to the case of the twining stem of Cuscuta, which, according to H. de Vries (ibid. p. 322) is sensitive to a touch like a tendril.

23 Dr. Maxwell Masters informs me that in almost all petioles which are cylindrical, such as those bearing peltate leaves, the woody vessels form a closed ring; semilunar bands of vessels being confined to petioles which are channelled along their upper surfaces. In accordance with this statement, it may be observed that the enlarged and clasped petiole of the Solanum, with its closed ring of woody vessels, has become more cylindrical than it was in its original unclasped condition.

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