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, Sphaerostemma, 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. {11} 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. {12}
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.