The phenomenon is not confined to woody plants, but has been met with in chicory, in Antirrhinum, and other herbaceous species.
It is very difficult in some cases to separate these instances of irregular torsion from those in which the twisting takes place in a more or less regular spiral direction. In the former case the fibres of the plant are only indirectly involved, but in the latter the fibres themselves are coiled spirally from right to left, or vice versâ (spiral torsion), while not unfrequently both conditions may be met with at the same time.
The leaves also are subject to similar deformities, of which a notable illustration has been recorded in the case of the date palm, Phœnix dactylifera, originally observed by Goethe, and figured and described by Jaeger;[355] the leaves are folded and twisted in every direction, in consequence of the fibrous band or cord which surrounds the leaves, and which generally breaks as the leaflets increase in size, remaining from some cause or other unbroken, and thus serving to restrain the growth.
A similar irregularity of growth occurs, not unfrequently, in the case of crocus leaves, when in the course of their growth, as they push their way through the soil, their progress becomes checked either by a stone or even by frost.
Spiral torsion.—Growth in a spiral direction, and the arrangement of the various organs of the plant in a spiral manner, are among the most common of natural phenomena in plants.[356] Fibres are coiled spirally in the minute vessels of flowering plants, and are not wholly wanting even among fungi. The leaf-organs are very generally spirally arranged; the leaf-stalks are often so twisted as to bring leaves on one plane which otherwise would occupy several. In the leaf itself we have a spiral twist taking place constantly in Alstrœmeria, in Avena, and other plants. A similar tendency is manifested in the flower-stalks, as in Cyclamen and Vallisneria, and the whole inflorescence, as in Spiranthes. Even the bark and wood of trees is often disposed spirally. This is very noticeable in some firs, and in the bark of the sweet chestnut (Castanea), of Thuja occidentalis, and other trees. The knaurs or excrescences which are sometimes found on the roots or stems of trees afford other illustrations of this universal tendency. These bodies consist of a number of embryo buds, which, from some cause or other, are incapable of lengthening. On examination every rudimentary or undeveloped bud may be seen to be surrounded by densely crowded fibres arranged spirally.
The axes of nearly all twining plants are themselves twisted, and twisted in a direction corresponding to the spontaneous revolving movement exhibited by these plants, as in the hop, the convolvulus, passion flower, &c., the degree of twisting being dependent to a great extent on the roughness of the surface around which the stem twines[357].
Considered as an exceptional occurrence, it occurs frequently in certain plants, and, when it affects the stem or branches, necessarily causes some changes in the arrangement of the parts attached to them; thus, spiral torsion of the axial organs is generally accompanied by displacement of the leaves, whorled leaves becoming alternate, and opposite or whorled leaves becoming arranged on one side of the stem only. Frequently also this condition is associated with fasciation, or, at least, with a distended or dilated state. An illustration of this in Asparagus has been figured at p. 14.
Very often the leaves are produced in a spiral line round the stem, as in a specimen of Dracocephalum speciosum described and figured by C. Morren. The leaves of this plant are naturally rectiserial and decussate, but, in the twisted stem the leaves were curviserial, and arranged according to the 5/13 plan. Now, referring to the ordinary notation of alternate leaves, we shall have the first leaf covered by the fifth, with two turns of the spiral; since decussate leaves result from two conjugate lines, the formula will be necessarily 2/5. The fraction 5/13 hence comes regularly into the 2/5 series (2/5, 3/8, 5/13). Thus, the leaves in assuming a new phyllotaxy, take one quite analogous to the normal one.
One of the most curious instances that have fallen under the writer's own observation occurred in the stem of Dipsacus fullonum. (See 'Proceedings of the Linnean Society,' March 6, 1855, vol. ii, p. 370). The stem was distended, and hollow, and twisted on itself; its fibres, moreover, were arranged in an oblique or spiral direction; the branches or leaf-stalks, which usually are arranged in an opposite and decussate manner, were, in this case, disposed in a linear series, one over the other, following the line of curvature of the stem. When the course of the fibres was traced from the base of one of the stalks, upward around the stem, a spiral was found to be completed at the base of the second stalk, above that which was made the starting point. Now, if opposite leaves depend on the shortened condition of the internode between the two leaves, then, in the teazel-stem just described, each turn of the spiral would represent a lengthened internode; and, if the fibres of this specimen could be untwisted, and made to assume the vertical direction, and, at the same time, the internodes were shortened, the result would be the opposition of the branches and the decussation of the pairs; this explanation is borne out by the similar twisting which takes place so frequently in the species of Galium and other Rubiaceæ.