This Essay first appeared in the ninth volume of the ‘Journal of the Linnean Society,’ published in 1865. It is here reproduced in a corrected and, I hope, clearer form, with some additional facts. The illustrations were drawn by my son, George Darwin. Fritz Müller, after the publication of my paper, sent to the Linnean Society (Journal, vol. ix., p. 344) some interesting observations on the climbing plants of South Brazil, to which I shall frequently refer. Recently two important memoirs, chiefly on the difference in growth between the upper and lower sides of tendrils, and on the mechanism of the movements of twining-plants, by Dr. Hugo de Vries, have appeared in the ‘Arbeiten des Botanischen Instituts in Würzburg,’ Heft. iii., 1873. These memoirs ought to be carefully studied by every one interested in the subject, as I can here give only references to the more important points. This excellent observer, as well as Professor Sachs, [iv] attributes all the movements of tendrils to rapid growth along one side; but, from reasons assigned towards the close of my fourth chapter, I cannot persuade myself that this holds good with respect to those due to a touch. In order that the reader may know what points have interested me most, I may call his attention to certain tendril-bearing plants; for instance, Bignonia capreolata, Cobæa, Echinocystis, and Hanburya, which display as beautiful adaptations as can be found in any part of the kingdom of nature. It is, also, an interesting fact that intermediate states between organs fitted for widely different functions, may be observed on the same individual plant of Corydalis claviculata and the common vine; and these cases illustrate in a striking manner the principle of the gradual evolution of species.
Since the publication of this Edition two papers by eminent botanists have appeared; Schwendener, ‘Das Winden der Pflanzen’ (Monatsberichte der Berliner Akademie, Dec. 1881), and J. Sachs, ‘Notiz über Schlingpflanzen’ (Arbeiten des botanischen Instituts in Würzburg, Bd. ii. p. 719, 1882). The view “that the capacity of revolving, on which most climbers depend, is inherent, though undeveloped, in almost every plant in the vegetable kingdom” (‘Climbing Plants,’ p. 205), has been confirmed by the observations on circumnutation since given in ‘The Power of Movement in Plants.’
On pp. 28, 32, 40, 53, statements are made with reference to the supposed acceleration of the revolving movement towards the light. It appears from the observations given in ‘The Power of Movement in Plants,’ p. 451, that these conclusions were drawn from insufficient observations, and are erroneous.
Introductory remarks—Description of the twining of the Hop—Torsion of the stems—Nature of the revolving movement, and manner of ascent—Stems not irritable—Rate of revolution in various plants—Thickness of the support round which plants can twine—Species which revolve in an anomalous manner.
I was led to this subject by an interesting, but short paper by Professor Asa Gray on the movements of the tendrils of some Cucurbitaceous plants. [1a] My observations were more than half completed before I learnt that the surprising phenomenon of the spontaneous revolutions of the stems and tendrils of climbing plants had been long ago observed by Palm and by Hugo von Mohl, [1b] and had subsequently been the subject of two memoirs by Dutrochet. [1c] Nevertheless, I believe that my observations, founded on the examination of above a hundred widely distinct living species, contain sufficient novelty to justify me in publishing them.
Climbing plants may be divided into four classes. First, those which twine spirally round a support, and are not aided by any other movement. Secondly, those endowed with irritable organs, which when they touch any object clasp it; such organs consisting of modified leaves, branches, or flower-peduncles. But these two classes sometimes graduate to a certain extent into one another. Plants of the third class ascend merely by the aid of hooks; and those of the fourth by rootlets; but as in neither class do the plants exhibit any special movements, they present little interest, and generally when I speak of climbing plants I refer to the two first great classes.
This is the largest subdivision, and is apparently the primordial and simplest condition of the class. My observations will be best given by taking a few special cases. When the shoot of a Hop (Humulus lupulus) rises from the ground, the two or three first-formed joints or internodes are straight and remain stationary; but the next-formed, whilst very young, may be seen to bend to one side and to travel slowly round towards all points of the compass, moving, like the hands of a watch, with the sun. The movement very soon acquires its full ordinary velocity. From seven observations made during August on shoots proceeding from a plant which had been cut down, and on another plant during April, the average rate during hot weather and during the day is 2 hrs. 8 m. for each revolution; and none of the revolutions varied much from this rate. The revolving movement continues as long as the plant continues to grow; but each separate internode, as it becomes old, ceases to move.
To ascertain more precisely what amount of movement each internode underwent, I kept a potted plant, during the night and day, in a well-warmed room to which I was confined by illness. A long shoot projected beyond the upper end of the supporting stick, and was steadily revolving. I then took a longer stick and tied up the shoot, so that only a very young internode, 1¾ of an inch in length, was left free. This was so nearly upright that its revolution could not be easily observed; but it certainly moved, and the side of the internode which was at one time convex became concave, which, as we shall hereafter see, is a sure sign of the revolving movement. I will assume that it made at least one revolution during the first twenty-four hours. Early the next morning its position was marked, and it made a second revolution in 9 hrs.; during the latter part of this revolution it moved much quicker, and the third circle was performed in the evening in a little over 3 hrs. As on the succeeding morning I found that the shoot revolved in 2 hrs. 45 m., it must have made during the night four revolutions, each at the average rate of a little over 3 hrs. I should add that the temperature of the room varied only a little. The shoot had now grown 3½ inches in length, and carried at its extremity a young internode 1 inch in length, which showed slight changes in its curvature. The next or ninth revolution was effected in 2 hrs. 30 m. From this time forward, the revolutions were easily observed. The thirty-sixth revolution was performed at the usual rate; so was the last or thirty-seventh, but it was not completed; for the internode suddenly became upright, and after moving to the centre, remained motionless. I tied a weight to its upper end, so as to bow it slightly and thus detect any movement; but there was none. Some time before the last revolution was half performed, the lower part of the internode ceased to move.
A few more remarks will complete all that need be said about this internode. It moved during five days; but the more rapid movements, after the performance of the third revolution, lasted during three days and twenty hours. The regular revolutions, from the ninth to thirty-sixth inclusive, were effected at the average rate of 2 hrs. 31 m.; but the weather was cold, and this affected the temperature of the room, especially during the night, and consequently retarded the rate of movement a little. There was only one irregular movement, which consisted in the stem rapidly making, after an unusually slow revolution, only the segment of a circle. After the seventeenth revolution the internode had grown from 1¾ to 6 inches in length, and carried an internode 1⅞ inch long, which was just perceptibly moving; and this carried a very minute ultimate internode. After the twenty-first revolution, the penultimate internode was 2½ inches long, and probably revolved in a period of about three hours. At the twenty-seventh revolution the lower and still moving internode was 8⅜, the penultimate 3½, and the ultimate 2½ inches in length; and the inclination of the whole shoot was such, that a circle 19 inches in diameter was swept by it. When the movement ceased, the lower internode was 9 inches, and the penultimate 6 inches in length; so that, from the twenty-seventh to thirty-seventh revolutions inclusive, three internodes were at the same time revolving.
The lower internode, when it ceased revolving, became upright and rigid; but as the whole shoot was left to grow unsupported, it became after a time bent into a nearly horizontal position, the uppermost and growing internodes still revolving at the extremity, but of course no longer round the old central point of the supporting stick. From the changed position of the centre of gravity of the extremity, as it revolved, a slight and slow swaying movement was given to the long horizontally projecting shoot; and this movement I at first thought was a spontaneous one. As the shoot grew, it hung down more and more, whilst the growing and revolving extremity turned itself up more and more.
With the Hop we have seen that three internodes were at the same time revolving; and this was the case with most of the plants observed by me. With all, if in full health, two internodes revolved; so that by the time the lower one ceased to revolve, the one above was in full action, with a terminal internode just commencing to move. With Hoya carnosa, on the other hand, a depending shoot, without any developed leaves, 32 inches in length, and consisting of seven internodes (a minute terminal one, an inch in length, being counted), continually, but slowly, swayed from side to side in a semicircular course, with the extreme internodes making complete revolutions. This swaying movement was certainly due to the movement of the lower internodes, which, however, had not force sufficient to swing the whole shoot round the central supporting stick. The case of another Asclepiadaceous plant, viz., Ceropegia Gardnerii, is worth briefly giving. I allowed the top to grow out almost horizontally to the length of 31 inches; this now consisted of three long internodes, terminated by two short ones. The whole revolved in a course opposed to the sun (the reverse of that of the Hop), at rates between 5 hrs. 15 m. and 6 hrs. 45 m. for each revolution. The extreme tip thus made a circle of above 5 feet (or 62 inches) in diameter and 16 feet in circumference, travelling at the rate of 32 or 33 inches per hour. The weather being hot, the plant was allowed to stand on my study-table; and it was an interesting spectacle to watch the long shoot sweeping this grand circle, night and day, in search of some object round which to twine.
If we take hold of a growing sapling, we can of course bend it to all sides in succession, so as to make the tip describe a circle, like that performed by the summit of a spontaneously revolving plant. By this movement the sapling is not in the least twisted round its own axis. I mention this because if a black point be painted on the bark, on the side which is uppermost when the sapling is bent towards the holder’s body, as the circle is described, the black point gradually turns round and sinks to the lower side, and comes up again when the circle is completed; and this gives the false appearance of twisting, which, in the case of spontaneously revolving plants, deceived me for a time. The appearance is the more deceitful because the axes of nearly all twining-plants are really twisted; and they are twisted in the same direction with the spontaneous revolving movement. To give an instance, the internode of the Hop of which the history has been recorded, was at first, as could be seen by the ridges on its surface, not in the least twisted; but when, after the 37th revolution, it had grown 9 inches long, and its revolving movement had ceased, it had become twisted three times round its own axis, in the line of the course of the sun; on the other hand, the common Convolvulus, which revolves in an opposite course to the Hop, becomes twisted in an opposite direction.
Hence it is not surprising that Hugo von Mohl (p. 105, 108, &c.) thought that the twisting of the axis caused the revolving movement; but it is not possible that the twisting of the axis of the Hop three times should have caused thirty-seven revolutions. Moreover, the revolving movement commenced in the young internode before any twisting of its axis could be detected. The internodes of a young Siphomeris and Lecontea revolved during several days, but became twisted only once round their own axes. The best evidence, however, that the twisting does not cause the revolving movement is afforded by many leaf-climbing and tendril-bearing plants (as Pisum sativum, Echinocystis lobata, Bignonia capreolata, Eccremocarpus scaber, and with the leaf-climbers, Solanum jasminoides and various species of Clematis), of which the internodes are not twisted, but which, as we shall hereafter see, regularly perform revolving movements like those of true twining-plants. Moreover, according to Palm (pp. 30, 95) and Mohl (p. 149), and Léon, [8] internodes may occasionally, and even not very rarely, be found which are twisted in an opposite direction to the other internodes on the same plant, and to the course of their revolutions; and this, according to Léon (p. 356), is the case with all the internodes of a certain variety of Phaseolus multiflorus. Internodes which have become twisted round their own axes, if they have not ceased to revolve, are still capable of twining round a support, as I have several times observed.
Mohl has remarked (p. 111) that when a stem twines round a smooth cylindrical stick, it does not become twisted. [9a] Accordingly I allowed kidney-beans to run up stretched string, and up smooth rods of iron and glass, one-third of an inch in diameter, and they became twisted only in that degree which follows as a mechanical necessity from the spiral winding. The stems, on the other hand, which had ascended ordinary rough sticks were all more or less and generally much twisted. The influence of the roughness of the support in causing axial twisting was well seen in the stems which had twined up the glass rods; for these rods were fixed into split sticks below, and were secured above to cross sticks, and the stems in passing these places became much twisted. As soon as the stems which had ascended the iron rods reached the summit and became free, they also became twisted; and this apparently occurred more quickly during windy than during calm weather. Several other facts could be given, showing that the axial twisting stands in some relation to inequalities in the support, and likewise to the shoot revolving freely without any support. Many plants, which are not twiners, become in some degree twisted round their own axes; [9b] but this occurs so much more generally and strongly with twining-plants than with other plants, that there must be some connexion between the capacity for twining and axial twisting. The stem probably gains rigidity by being twisted (on the same principle that a much twisted rope is stiffer than a slackly twisted one), and is thus indirectly benefited so as to be enabled to pass over inequalities in its spiral ascent, and to carry its own weight when allowed to revolve freely. [10]
I have alluded to the twisting which necessarily follows on mechanical principles from the spiral ascent of a stem, namely, one twist for each spire completed. This was well shown by painting straight lines on living stems, and then allowing them to twine; but, as I shall have to recur to this subject under Tendrils, it may be here passed over.
The revolving movement of a twining plant has been compared with that of the tip of a sapling, moved round and round by the hand held some way down the stem; but there is one important difference. The upper part of the sapling when thus moved remains straight; but with twining plants every part of the revolving shoot has its own separate and independent movement. This is easily proved; for when the lower half or two-thirds of a long revolving shoot is tied to a stick, the upper free part continues steadily revolving. Even if the whole shoot, except an inch or two of the extremity, be tied up, this part, as I have seen in the case of the Hop, Ceropegia, Convolvulus, &c., goes on revolving, but much more slowly; for the internodes, until they have grown to some little length, always move slowly. If we look to the one, two, or several internodes of a revolving shoot, they will be all seen to be more or less bowed, either during the whole or during a large part of each revolution. Now if a coloured streak be painted (this was done with a large number of twining plants) along, we will say, the convex surface, the streak will after a time (depending on the rate of revolution) be found to be running laterally along one side of the bow, then along the concave side, then laterally on the opposite side, and, lastly, again on the originally convex surface. This clearly proves that during the revolving movement the internodes become bowed in every direction. The movement is, in fact, a continuous self-bowing of the whole shoot, successively directed to all points of the compass; and has been well designated by Sachs as a revolving nutation.
As this movement is rather difficult to understand, it will be well to give an illustration. Take a sapling and bend it to the south, and paint a black line on the convex surface; let the sapling spring up and bend it to the east, and the black line will be seen to run along the lateral face fronting the north; bend it to the north, the black line will be on the concave surface; bend it to the west, the line will again be on the lateral face; and when again bent to the south, the line will be on the original convex surface. Now, instead of bending the sapling, let us suppose that the cells along its northern surface from the base to the tip were to grow much more rapidly than on the three other sides, the whole shoot would then necessarily be bowed to the south; and let the longitudinal growing surface creep round the shoot, deserting by slow degrees the northern side and encroaching on the western side, and so round by the south, by the east, again to the north. In this case the shoot would remain always bowed with the painted line appearing on the several above specified surfaces, and with the point of the shoot successively directed to each point of the compass. In fact, we should have the exact kind of movement performed by the revolving shoots of twining plants. [12]
It must not be supposed that the revolving movement is as regular as that given in the above illustration; in very many cases the tip describes an ellipse, even a very narrow ellipse. To recur once again to our illustration, if we suppose only the northern and southern surfaces of the sapling alternately to grow rapidly, the summit would describe a simple arc; if the growth first travelled a very little to the western face, and during the return a very little to the eastern face, a narrow ellipse would be described; and the sapling would be straight as it passed to and fro through the intermediate space; and a complete straightening of the shoot may often be observed in revolving plants. The movement is frequently such that three of the sides of the shoot seem to be growing in due order more rapidly than the remaining side; so that a semi-circle instead of a circle is described, the shoot becoming straight and upright during half of its course.
When a revolving shoot consists of several internodes, the lower ones bend together at the same rate, but one or two of the terminal ones bend at a slower rate; hence, though at times all the internodes are in the same direction, at other times the shoot is rendered slightly serpentine. The rate of revolution of the whole shoot, if judged by the movement of the extreme tip, is thus at times accelerated or retarded. One other point must be noticed. Authors have observed that the end of the shoot in many twining plants is completely hooked; this is very general, for instance, with the Asclepiadaceæ. The hooked tip, in all the cases observed by me, viz. in Ceropegia, Sphærostemma, Clerodendron, Wistaria, Stephania, Akebia, and Siphomeris, has exactly the same kind of movement as the other internodes; for a line painted on the convex surface first becomes lateral and then concave; but, owing to the youth of these terminal internodes, the reversal of the hook is a slower process than that of the revolving movement. [14] This strongly marked tendency in the young, terminal and flexible internodes, to bend in a greater degree or more abruptly than the other internodes, is of service to the plant; for not only does the hook thus formed sometimes serve to catch a support, but (and this seems to be much more important) it causes the extremity of the shoot to embrace the support much more closely than it could otherwise have done, and thus aids in preventing the stem from being blown away during windy weather, as I have many times observed. In Lonicera brachypoda the hook only straightens itself periodically, and never becomes reversed. I will not assert that the tips of all twining plants when hooked, either reverse themselves or become periodically straight, in the manner just described; for the hooked form may in some cases be permanent, and be due to the manner of growth of the species, as with the tips of the shoots of the common vine, and more plainly with those of Cissus discolor—plants which are not spiral twiners.
The first purpose of the spontaneous revolving movement, or, more strictly speaking, of the continuous bowing movement directed successively to all points of the compass, is, as Mohl has remarked, to favour the shoot finding a support. This is admirably effected by the revolutions carried on night and day, a wider and wider circle being swept as the shoot increases in length. This movement likewise explains how the plants twine; for when a revolving shoot meets with a support, its motion is necessarily arrested at the point of contact, but the free projecting part goes on revolving. As this continues, higher and higher points are brought into contact with the support and are arrested; and so onwards to the extremity; and thus the shoot winds round its support. When the shoot follows the sun in its revolving course, it winds round the support from right to left, the support being supposed to stand in front of the beholder; when the shoot revolves in an opposite direction, the line of winding is reversed. As each internode loses from age its power of revolving, it likewise loses its power of spirally twining. If a man swings a rope round his head, and the end hits a stick, it will coil round the stick according to the direction of the swinging movement; so it is with a twining plant, a line of growth travelling round the free part of the shoot causing it to bend towards the opposite side, and this replaces the momentum of the free end of the rope.
All the authors, except Palm and Mohl, who have discussed the spiral twining of plants, maintain that such plants have a natural tendency to grow spirally. Mohl believes (p. 112) that twining stems have a dull kind of irritability, so that they bend towards any object which they touch; but this is denied by Palm. Even before reading Mohl’s interesting treatise, this view seemed to me so probable that I tested it in every way that I could, but always with a negative result. I rubbed many shoots much harder than is necessary to excite movement in any tendril or in the foot-stalk of any leaf climber, but without any effect. I then tied a light forked twig to a shoot of a Hop, a Ceropegia, Sphærostemma, and Adhatoda, so that the fork pressed on one side alone of the shoot and revolved with it; I purposely selected some very slow revolvers, as it seemed most likely that these would profit most from possessing irritability; but in no case was any effect produced. [16] Moreover, when a shoot winds round a support, the winding movement is always slower, as we shall immediately see, than whilst it revolves freely and touches nothing. Hence I conclude that twining stems are not irritable; and indeed it is not probable that they should be so, as nature always economizes her means, and irritability would have been superfluous. Nevertheless I do not wish to assert that they are never irritable; for the growing axis of the leaf-climbing, but not spirally twining, Lophospermum scandens is, certainly irritable; but this case gives me confidence that ordinary twiners do not possess any such quality, for directly after putting a stick to the Lophopermum, I saw that it behaved differently from a true twiner or any other leaf-climber. [17]
The belief that twiners have a natural tendency to grow spirally, probably arose from their assuming a spiral form when wound round a support, and from the extremity, even whilst remaining free, sometimes assuming this form. The free internodes of vigorously growing plants, when they cease to revolve, become straight, and show no tendency to be spiral; but when a shoot has nearly ceased to grow, or when the plant is unhealthy, the extremity does occasionally become spiral. I have seen this in a remarkable manner with the ends of the shoots of the Stauntonia and of the allied Akebia, which became wound up into a close spire, just like a tendril; and this was apt to occur after some small, ill-formed leaves had perished. The explanation, I believe, is, that in such cases the lower parts of the terminal internodes very gradually and successively lose their power of movement, whilst the portions just above move onwards and in their turn become motionless; and this ends in forming an irregular spire.
When a revolving shoot strikes a stick, it winds round it rather more slowly than it revolves. For instance, a shoot of the Ceropegia, revolved in 6 hrs., but took 9 hrs. 30 m. to make one complete spire round a stick; Aristolochia gigas revolved in about 5 hrs., but took 9 hrs. 15 m. to complete its spire. This, I presume, is due to the continued disturbance of the impelling force by the arrestment of the movement at successive points; and we shall hereafter see that even shaking a plant retards the revolving movement. The terminal internodes of a long, much-inclined, revolving shoot of the Ceropegia, after they had wound round a stick, always slipped up it, so as to render the spire more open than it was at first; and this was probably in part due to the force which caused the revolutions, being now almost freed from the constraint of gravity and allowed to act freely. With the Wistaria, on the other hand, a long horizontal shoot wound itself at first into a very close spire, which remained unchanged; but subsequently, as the shoot twined spirally up its support, it made a much more open spire. With all the many plants which were allowed freely to ascend a support, the terminal internodes made at first a close spire; and this, during windy weather, served to keep the shoots in close contact with their support; but as the penultimate internodes grew in length, they pushed themselves up for a considerable space (ascertained by coloured marks on the shoot and on the support) round the stick, and the spire became more open. [18]
It follows from this latter fact that the position occupied by each leaf with respect to the support depends on the growth of the internodes after they have become spirally wound round it. I mention this on account of an observation by Palm (p. 34), who states that the opposite leaves of the Hop always stand in a row, exactly over one another, on the same side of the supporting stick, whatever its thickness may be. My sons visited a hop-field for me, and reported that though they generally found the points of insertion of the leaves standing over each other for a space of two or three feet in height, yet this never occurred up the whole length of the pole; the points of insertion forming, as might have been expected, an irregular spire. Any irregularity in the pole entirely destroyed the regularity of position of the leaves. From casual inspection, it appeared to me that the opposite leaves of Thunbergia alata were arranged in lines up the sticks round which they had twined; accordingly, I raised a dozen plants, and gave them sticks of various thicknesses, as well as string, to twine round; and in this case one alone out of the dozen had its leaves arranged in a perpendicular line: I conclude, therefore, Palm’s statement is not quite accurate.
The leaves of different twining-plants are arranged on the stem (before it has twined) alternately, or oppositely, or in a spire. In the latter case the line of insertion of the leaves and the course of the revolutions coincide. This fact has been well shown by Dutrochet, [19] who found different individuals of Solanum dulcamara twining in opposite directions, and these had their leaves in each case spirally arranged in the same direction. A dense whorl of many leaves would apparently be incommodious for a twining plant, and some authors assert that none have their leaves thus arranged; but a twining Siphomeris has whorls of three leaves.
If a stick which has arrested a revolving shoot, but has not as yet been encircled, be suddenly taken away, the shoot generally springs forward, showing that it was pressing with some force against the stick. After a shoot has wound round a stick, if this be withdrawn, it retains for a time its spiral form; it then straightens itself, and again commences to revolve. The long, much-inclined shoot of the Ceropegia