Fig. 172.—Twisted stem of Dipsacus fullonum.
G. Franc[358] was one of the first to notice this twisting in Galium, and M. Duchartre,[359] in mentioning a similar instance, gives the following explanation of the appearance which will be found to apply to most of these cases. In the normal stem of Galium Mollugo the branches are opposite in each verticil and crossed in the two successive ones. The stem is four-angled, each angle having a nerve. Each of these nerves, springing from the origin of a branch in one whorl, terminates in the interval which separates the point of origin of the two branches in the whorl next above it. In the deformed stem one of the nerves corresponds to the insertion of a branch, its neighbour is in the adjoining vacant space; hence it results that four nerves correspond to two branches and to two consecutive interspaces, and hence the analogy between a single normal internode provided with its two branches and its four nerves. What confirms this inference is that the nerve, which begins at the point of origin of a branch, after making one spiral turn round the stem, terminates in the interval that separates the two following branches, just as in a branch of the normal stem it ends in the upper whorl between the two next branches. The torsion, then, in this Galium caused the separation of the two opposite branches of the same verticil, and placed them one above another, and this being reproduced in all the whorls, all the branches come to be arranged on the same longitudinal line. The leaves are susceptible of the same explanation; they are inserted in groups of three or four in one arc round the origin of each branch. In the malformation each series or group of four leaves, with its central branch, is equivalent to half a whorl of the natural plant with its axillary branch. In other words, the malformation consists in a torsion of the stem, which separates each whorl into two distinct halves; these half-whorls, with their axillary branches, are placed on a single longitudinal series one above another. This case is quoted at some length, as it is an admirable example of a very common form of malformation in these plants.
In some parts of Holland where madder is cultivated a similar deformation is particularly frequent. The leaves, however, are not always grouped in the way in which they were described by M. Duchartre, but more commonly form a single continuous line; when arranged in leaf-whorls it generally happens that some of the leaves are turned downwards, while others are erect. It has been said that this condition occurs particularly frequently in plants growing in damp places. It is certainly true that spiral torsion of the stem is specially frequent in the species of Equisetum, most of which grow in such spots. In these plants either the whole of the upper part of the stem is thus twisted, or a portion only: thus Reinsch[360] cites a case in Equisetum Telmateia, where the upper and lower portions of the stem were normal, while the intermediate portion was twisted spirally. In this instance the whorl next beneath the spiral had twenty-eight branchlets, and that immediately above it thirty. Along the course of the spire there were two hundred and three; dividing this latter number by the mean of the two preceding, it was seen that the spire included the constituents of seven ordinary verticils.
Fig. 173.—Stem of Galium spirally twisted. From a specimen communicated by Mr. Darwin.
Here also may be mentioned a curious bamboo, the stem of which is preserved in the British Museum, and in which the internodes, on the exterior, and the corresponding diaphragms and cavities within are spiral or oblique in direction.
The root is also subject to the same malformation, the inducing cause being usually some obstruction to downward growth, as when a plant has been grown in a small pot, and becomes, as gardeners say, pot-bound.