In the many other revolving and twining plants observed by me, I never but twice saw the movement reversed; once, and only for a short space, in Ipomoea jucunda; but frequently with Hibbertia dentata. This plant at first perplexed me much, for I continually observed its long and flexible shoots, evidently well fitted for twining, make a whole, or half, or quarter circle in one direction and then in an opposite direction; consequently, when I placed the shoots near thin or thick sticks, or perpendicularly stretched string, they seemed as if constantly trying to ascend, but always failed. I then surrounded the plant with a mass of branched twigs; the shoots ascended, and passed through them, but several came out laterally, and their depending extremities seldom turned upwards as is usual with twining plants. Finally, I surrounded a second plant with many thin upright sticks, and placed it near the first one with twigs; and now both had got what they liked, for they twined up the parallel sticks, sometimes winding round one and sometimes round several; and the shoots travelled laterally from one to the other pot; but as the plants grew older, some of the shoots twined regularly up thin upright sticks. Though the revolving movement was sometimes in one direction and sometimes in the other, the twining was invariably from left to right; [36] so that the more potent or persistent movement of revolution must have been in opposition to the course of the sun. It would appear that this Hibbertia is adapted both to ascend by twining, and to ramble laterally through the thick Australian scrub.
I have described the above case in some detail, because, as far as I have seen, it is rare to find any special adaptations with twining plants, in which respect they differ much from the more highly organized tendril-bearers. The Solanum dulcamara, as we shall presently see, can twine only round stems which are both thin and flexible. Most twining plants are adapted to ascend supports of moderate though of different thicknesses. Our English twiners, as far as I have seen, never twine round trees, excepting the honeysuckle (Lonicera periclymenum), which I have observed twining up a young beech-tree nearly 4½ inches in diameter. Mohl (p. 134) found that the Phaseolus multiflorus and Ipomoea purpurea could not, when placed in a room with the light entering on one side, twine round sticks between 3 and 4 inches in diameter; for this interfered, in a manner presently to be explained, with the revolving movement. In the open air, however, the Phaseolus twined round a support of the above thickness, but failed in twining round one 9 inches in diameter. Nevertheless, some twiners of the warmer temperate regions can manage this latter degree of thickness; for I hear from Dr. Hooker that at Kew the Ruscus androgynus has ascended a column 9 inches in diameter; and although a Wistaria grown by me in a small pot tried in vain for weeks to get round a post between 5 and 6 inches in thickness, yet at Kew a plant ascended a trunk above 6 inches in diameter. The tropical twiners, on the other hand, can ascend thicker trees; I hear from Drs. Thomson and Hooker that this is the case with the Butea parviflora, one of the Menispermaceæ, and with some Dalbergias and other Leguminosæ. [37] This power would be necessary for any species which had to ascend by twining the large trees of a tropical forest; otherwise they would hardly ever be able to reach the light. In our temperate countries it would be injurious to the twining plants which die down every year if they were enabled to twine round trunks of trees, for they could not grow tall enough in a single season to reach the summit and gain the light.
By what means certain twining plants are adapted to ascend only thin stems, whilst others can twine round thicker ones, I do not know. It appeared to me probable that twining plants with very long revolving shoots would be able to ascend thick supports; accordingly I placed Ceropegia Gardnerii near a post 6 inches in diameter, but the shoots entirely failed to wind round it; their great length and power of movement merely aid them in finding a distant stem round which to twine. The Sphærostemma marmoratum is a vigorous tropical twiner; and as it is a very slow revolver, I thought that this latter circumstance might help it in ascending a thick support; but though it was able to wind round a 6-inch post, it could do this only on the same level or plane, and did not form a spire and thus ascend.
As ferns differ so much in structure from phanerogamic plants, it may be worth while here to show that twining ferns do not differ in their habits from other twining plants. In Lygodium articulatum the two internodes of the stem (properly the rachis) which are first formed above the root-stock do not move; the third from the ground revolves, but at first very slowly. This species is a slow revolver: but L. scandens made five revolutions, each at the average rate of 5 hrs. 45 m.; and this represents fairly well the usual rate, taking quick and slow movers, amongst phanerogamic plants. The rate was accelerated by increased temperature. At each stage of growth only the two upper internodes revolved. A line painted along the convex surface of a revolving internode becomes first lateral, then concave, then lateral and ultimately again convex. Neither the internodes nor the petioles are irritable when rubbed. The movement is in the usual direction, namely, in opposition to the course of the sun; and when the stem twines round a thin stick, it becomes twisted on its own axis in the same direction. After the young internodes have twined round a stick, their continued growth causes them to slip a little upwards. If the stick be soon removed, they straighten themselves, and recommence revolving. The extremities of the depending shoots turn upwards, and twine on themselves. In all these respects we have complete identity with twining phanerogamic plants; and the above enumeration may serve as a summary of the leading characteristics of all twining plants.
The power of revolving depends on the general health and vigour of the plant, as has been laboriously shown by Palm. But the movement of each separate internode is so independent of the others, that cutting off an upper one does not affect the revolutions of a lower one. When, however, Dutrochet cut off two whole shoots of the Hop, and placed them in water, the movement was greatly retarded; for one revolved in 20 hrs. and the other in 23 hrs., whereas they ought to have revolved in between 2 hrs. and 2 hrs. 30 m. Shoots of the Kidney-bean, cut off and placed in water, were similarly retarded, but in a less degree. I have repeatedly observed that carrying a plant from the greenhouse to my room, or from one part to another of the greenhouse, always stopped the movement for a time; hence I conclude that plants in a state of nature and growing in exposed situations, would not make their revolutions during very stormy weather. A decrease in temperature always caused a considerable retardation in the rate of revolution; but Dutrochet (tom. xvii. pp. 994, 996) has given such precise observations on this head with respect to the common pea that I need say nothing more. When twining plants are placed near a window in a room, the light in some cases has a remarkable power (as was likewise observed by Dutrochet, p. 998, with the pea) on the revolving movement, but this differs in degree with different plants; thus Ipomoea jucunda made a complete circle in 5 hrs. 30 m.; the semicircle from the light taking 4 hrs. 80 m., and that towards the light only 1 hr. Lonicera brachypoda revolved, in a reversed direction to the Ipomoea, in 8 hrs.; the semicircle from the light taking 5 hrs. 23 m., and that to the light only 2 hrs. 37 m. From the rate of revolution in all the plants observed by me, being nearly the same during the night and the day, I infer that the action of the light is confined to retarding one semicircle and accelerating the other, so as not to modify greatly the rate of the whole revolution. This action of the light is remarkable, when we reflect how little the leaves are developed on the young and thin revolving internodes. It is all the more remarkable, as botanists believe (Mohl, p. 119) that twining plants are but little sensitive to the action of light.
I will conclude my account of twining plants by giving a few miscellaneous and curious cases. With most twining plants all the branches, however many there may be, go on revolving together; but, according to Mohl (p. 4), only the lateral branches of Tamus elephantipes twine, and not the main stem. On the other hand, with a climbing species of Asparagus, the leading shoot alone, and not the branches, revolved and twined; but it should be stated that the plant was not growing vigorously. My plants of Combretum argenteum and C. purpureum made numerous short healthy shoots; but they showed no signs of revolving, and I could not conceive how these plants could be climbers; but at last C. argenteum put forth from the lower part of one of its main branches a thin shoot, 5 or 6 feet in length, differing greatly in appearance from the previous shoots, owing to its leaves being little developed, and this shoot revolved vigorously and twined. So that this plant produces shoots of two kinds. With Periploca Græca (Palm, p. 43) the uppermost shoots alone twine. Polygonum convolvulus twines only during the middle of the summer (Palm, p. 43, 94); and plants growing vigorously in the autumn show no inclination to climb. The majority of Asclepiadaceæ are twiners; but Asclepias nigra only "in fertiliori solo incipit scandere subvolubili caule"...
