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This movement, which shortens the tendrils and renders them elastic, commences in half a day, or in a day or two after their extremities have caught some object. There is no such movement in any leaf- climber, with the exception of an occasional trace of it in the petioles of Tropaeolum tricolorum. On the other hand, the tendrils of all tendril-bearing plants, contract spirally after they have caught an object with the following exceptions. Firstly, Corydalis claviculata, but then this plant might be called a leaf-climber. Secondly and thirdly, Bignonia unguis with its close allies, and Cardiospermum; but their tendrils are so short that their contraction could hardly occur, and would be quite superfluous. Fourthly, Smilax aspera offers a more marked exception, as its tendrils are moderately long. The tendrils of Dicentra, whilst the plant is young, are short and after attachment only become slightly flexuous; in older plants they are longer and then they contract spirally. I have seen no other exceptions to the rule that tendrils, after clasping with their extremities a support, undergo spiral contraction. When, however, the tendril of a plant of which the stem is immovably fixed, catches some fixed object, it does not contract, simply because it cannot; this, however, rarely occurs. In the common Pea the lateral branches alone contract, and not the central stem; and with most plants, such as the Vine, Passiflora, Bryony, the basal portion never forms a spire.

I have said that in Corydalis claviculata the end of the leaf or tendril (for this part may be indifferently so called) does not contract into a spire. The branchlets, however, after they have wound round thin twigs, become deeply sinuous or zigzag. Moreover the whole end of the petiole or tendril, if it seizes nothing, bends after a time abruptly downwards and inwards, showing that its outer surface has gone on growing after the inner surface has ceased to grow. That growth is the chief cause of the spiral contraction of tendrils may be safely admitted, as shown by the recent researches of H. de Vries. I will, however, add one little fact in support of this conclusion.

If the short, nearly straight portion of an attached tendril of Passiflora gracilis, (and, as I believe, of other tendrils,) between the opposed spires, be examined, it will be found to be transversely wrinkled in a conspicuous manner on the outside; and this would naturally follow if the outer side had grown more than the inner side, this part being at the same time forcibly prevented from becoming curved. So again the whole outer surface of a spirally wound tendril becomes wrinkled if it be pulled straight. Nevertheless, as the contraction travels from the extremity of a tendril, after it has been stimulated by contact with a support, down to the base, I cannot avoid doubting, from reasons presently to be given, whether the whole effect ought to be attributed to growth. An unattached tendril rolls itself up into a flat helix, as in the case of Cardiospermum, if the contraction commences at the extremity and is quite regular; but if the continued growth of the outer surface is a little lateral, or if the process begins near the base, the terminal portion cannot be rolled up within the basal portion, and the tendril then forms a more or less open spire. A similar result follows if the extremity has caught some object, and is thus held fast.

The tendrils of many kinds of plants, if they catch nothing, contract after an interval of several days or weeks into a spire; but in these cases the movement takes place after the tendril has lost its revolving power and hangs down; it has also then partly or wholly lost its sensibility; so that this movement can be of no use. The spiral contraction of unattached tendrils is a much slower process than that of attached ones. Young tendrils which have caught a support and are spirally contracted, may constantly be seen on the same stem with the much older unattached and uncontracted tendrils. In the Echinocystis I have seen a tendril with the two lateral branches encircling twigs and contracted into beautiful spires, whilst the main branch which had caught nothing remained for many days straight. In this plant I once observed a main branch after it had caught a stick become spirally flexuous in 7 hrs., and spirally contracted in 18 hrs. Generally the tendrils of the Echinocystis begin to contract in from 12 hrs. to 24 hrs. after catching some object; whilst unattached tendrils do not begin to contract until two or three or even more days after all revolving movement has ceased. A full-grown tendril of Passiflora quadrangularis which had caught a stick began in 8 hrs. to contract, and in 24 hrs. formed several spires; a younger tendril, only two-thirds grown, showed the first trace of contraction in two days after clasping a stick, and in two more days formed several spires. It appears, therefore, that the contraction does not begin until the tendril is grown to nearly its full length. Another young tendril of about the same age and length as the last did not catch any object; it acquired its full length in four days; in six additional days it first became flexuous, and in two more days formed one complete spire. This first spire was formed towards the basal end, and the contraction steadily but slowly progressed towards the apex; but the whole was not closely wound up into a spire until 21 days had elapsed from the first observation, that is, until 17 days after the tendril had grown to its full length.

The spiral contraction of tendrils is quite independent of their power of spontaneously revolving, for it occurs in tendrils, such as those of Lathyrus grandiflorus and Ampelopsis hederacea, which do not revolve. It is not necessarily related to the curling of the tips round a support, as we see with the Ampelopsis and Bignonia capreolata, in which the development of adherent discs suffices to cause spiral contraction. Yet in some cases this contraction seems connected with the curling or clasping movement, due to contact with a support; for not only does it soon follow this act, but the contraction generally begins close to the curled extremity, and travels downwards to the base. If, however, a tendril be very slack, the whole length almost simultaneously becomes at first flexuous and then spiral. Again, the tendrils of some few plants never contract spirally unless they have first seized hold of some object; if they catch nothing they hang down, remaining straight, until they wither and drop off: this is the case with the tendrils of Bignonia, which consist of modified leaves, and with those of three genera of the Vitaceae, which are modified flower-peduncles. But in the great majority of cases, tendrils which have never come in contact with any object, after a time contract spirally. All these facts taken together, show that the act of clasping a support and the spiral contraction of the whole length of the tendril, are phenomena not necessarily connected.

The spiral contraction which ensues after a tendril has caught a support is of high service to the plant; hence its almost universal occurrence with species belonging to widely different orders. When a shoot is inclined and its tendril has caught an object above, the spiral contraction drags up the shoot. When the shoot is upright, the growth of the stem, after the tendrils have seized some object above, would leave it slack, were it not for the spiral contraction which draws up the stem as it increases in length. Thus there is no waste of growth, and the stretched stem ascends by the shortest course. When a terminal branchlet of the tendril of Cobaea catches a stick, we have seen how well the spiral contraction successively brings the other branchlets, one after the other, into contact with the stick, until the whole tendril grasps it in an inextricable knot. When a tendril has caught a yielding object, this is sometimes enveloped and still further secured by the spiral folds, as I have seen with Passiflora quadrangularis; but this action is of little importance.

A far more important service rendered by the spiral contraction of the tendrils is that they are thus made highly elastic. As before remarked under Ampelopsis, the strain is thus distributed equally between the several attached branches; and this renders the whole far stronger than it otherwise would be, as the branches cannot break separately. It is this elasticity which protects both branched and simple tendrils from being torn away from their supports during stormy weather. I have more than once gone on purpose during a gale to watch a Bryony growing in an exposed hedge, with its tendrils attached to the surrounding bushes; and as the thick and thin branches were tossed to and fro by the wind, the tendrils, had they not been excessively elastic, would instantly have been torn off and the plant thrown prostrate. But as it was, the Bryony safely rode out the gale, like a ship with two anchors down, and with a long range of cable ahead to serve as a spring as she surges to the storm.

When an unattached tendril contracts spirally, the spire always runs in the same direction from tip to base. A tendril, on the other hand, which has caught a support by its extremity, although the same side is concave from end to end, invariably becomes twisted in one part in one direction, and in another part in the opposite direction; the oppositely turned spires being separated by a short straight portion. This curious and symmetrical structure has been noticed by several botanists, but has not been sufficiently explained. {35} It occurs without exception with all tendrils which after catching an object contract spirally, but is of course most conspicuous in the longer tendrils. It never occurs with uncaught tendrils; and when this appears to have occurred, it will be found that the tendril had originally seized some object and had afterwards been torn free. Commonly, all the spires at one end of an attached tendril run in one direction, and all those at the other end in the opposite direction, with a single short straight portion in the middle; but I have seen a tendril with the spires alternately turning five times in opposite directions, with straight pieces between them; and M. Leon has seen seven or eight such alternations. Whether the spires turn once or more than once in opposite directions, there are as many turns in the one direction as in the other. For instance, I gathered ten attached tendrils of the Bryony, the longest with 33, and the shortest with only 8 spiral turns; and the number of turns in the one direction was in every case the same (within one) as in the opposite direction.

The explanation of this curious little fact is not difficult. I will not attempt any geometrical reasoning, but will give only a practical illustration. In doing this, I shall first have to allude to a point which was almost passed over when treating of Twining-plants. If we hold in our left hand a bundle of parallel strings, we can with our right hand turn these round and round, thus imitating the revolving movement of a twining plant, and the strings do not become twisted. But if we hold at the same time a stick in our left hand, in such a position that the strings become spirally turned round it, they will inevitably become twisted. Hence a straight coloured line, painted along the internodes of a twining plant before it has wound round a support, becomes twisted or spiral after it has wound round. I painted a red line on the straight internodes of a Humulus, Mikania, Ceropegia, Convolvulus, and Phaseolus, and saw it become twisted as the plant wound round a stick. It is possible that the stems of some plants by spontaneously turning on their own axes, at the proper rate and in the proper direction, might avoid becoming twisted; but I have seen no such case.

In the above illustration, the parallel strings were wound round a stick; but this is by no means necessary, for if wound into a hollow coil (as can be done with a narrow slip of elastic paper) there is the same inevitable twisting of the axis. When, therefore, a free tendril coils itself into a spire, it must either become twisted


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