THE CIRCUMNUTATION OF CLIMBING PLANTS.
The simplest case of modified circumnutation is that offered by climbing plants, with the exception of those which climb by the aid of motionless hooks or of rootlets: for the modification consists chiefly in the greatly increased amplitude of the movement. This would follow either from greatly increased growth over a small length, or more probably from moderately increased growth spread over a considerable length of the moving organ, preceded by turgescence, and acting successively on all sides. The circumnutation of climbers is more regular than that of ordinary plants; but in almost every other respect there is a close similarity between their movements, namely, in their tendency to describe ellipses directed successively to all points of the compass—in their courses being often interrupted by zigzag lines, triangles, loops, or small ellipses—in the rate of movement, and in different species revolving once or several times within the same length of time. In the same internode, the movements cease first in the lower part and then slowly upwards. In both sets of cases the movement may be modified in a closely analogous manner by geotropism and by heliotropism; though few climbing plants are heliotropic. Other points of similarity might be pointed out.
That the movements of climbing plants consist of ordinary circumnutation, modified by being increased in amplitude, is well exhibited whilst the plants are very young; for at this early age they move like other seedlings, but as they grow older their movements gradually increase without undergoing any other change. That this power is innate, and is not excited by any external agencies, beyond those necessary for growth and vigour, is obvious. No one doubts that this power has been gained for the sake of enabling climbing plants to ascend to a height, and thus to reach the light. This is effected by two very different methods; first, by twining spirally round a support, but to do so their stems must be long and flexible; and, secondly, in the case of leaf-climbers and tendril-bearers, by bringing these organs into contact with a support, which is then seized by the aid of their sensitiveness. It may be here remarked that these latter movements have no relation, as far as we can judge, with circumnutation. In other cases the tips of tendrils, after having been brought into contact with a support, become developed into little discs which adhere firmly to it.
We have said that the circumnutation of climbing plants differs from that of ordinary plants chiefly by its greater amplitude. But most leaves circumnutate in an almost vertical plane, and therefore describe very narrow ellipses, whereas the many kinds of tendrils which consist of metamorphosed leaves, make much broader ellipses or nearly circular figures; and thus they have a far better chance of catching hold of a support on any side. The movements of climbing plants have also been modified in some few other special ways. Thus the circumnutating stems of Solnanum dulcamara can twine round a support only when this is as thin and flexible as a string or thread. The twining stems of several British plants cannot twine round a support when it is more than a few inches in thickness; whilst in tropical forests some can embrace thick trunks;[[1]] and this great difference in power depends on some unknown difference in their manner of circumnutation. The most remarkable special modification of this movement which we have observed is in the tendrils of Echinocystis lobata; these are usually inclined at about 45° above the horizon, but they stiffen and straighten themselves so as to stand upright in a part of their circular course, namely, when they approach and have to pass over the summit or the shoot from which they arise. If they had not possessed and exercised this curious power, they would infallibly have struck against the summit of the shoot and been arrested in their course. As soon as one of these tendrils with its three branches begins to stiffen itself and rise up vertically, the revolving motion becomes more rapid; and as soon as it has passed over the point of difficulty, its motion coinciding with that from its own weight, causes it to fall into its previously inclined position so quickly, that the apex can be seen travelling like the hand of a gigantic clock.
[1] ‘The Movements and Habits of Climbing Plants,’ p. 36.
A large number of ordinary leaves and leaflets and a few flower-peduncles are provided with pulvini; but this is not the case with a single tendril at present known. The cause of this difference probably lies in the fact, that the chief service of a pulvinus is to prolong the movement of the part thus provided after growth has ceased; and as tendrils or other climbing-organs are of use only whilst the plant is increasing in height or growing, a pulvinus which served to prolong their movements would be useless.
It was shown in the last chapter that the stolons or runners of certain plants circumnutate largely, and that this movement apparently aids them in finding a passage between the crowded stems of adjoining plants. If it could be proved that their movements had been modified and increased for this special purpose, they ought to have been included in the present chapter; but as the amplitude of their revolutions is not so conspicuously different from that of ordinary plants, as in the case of climbers, we have no evidence on this head. We encounter the same doubt in the case of some plants which bury their pods in the ground. This burying process is certainly favoured by the circumnutation of the flower-peduncle; but we do not know whether it has been increased for this special purpose.