On the fourth day they were deflected by an average angle of 63o from a line perpendicular to the lower surface, and were therefore considerably more curved than the hypocotyl and radicle in the bean at B (Fig. 59), though in the same relative direction.

It will, we presume, be admitted that all leguminous plants with hypogean cotyledons are descended from forms which once raised their cotyledons above the ground in the ordinary manner; and in doing so, it is certain that their hypocotyls would have been abruptly arched, as in the case of every other dicotyledonous plant. This is especially clear in the case of Phaseolus, for out of five species, the seedlings of which we observed, namely, P. multiflorus, caracalla, vulgaris, Hernandesii and Roxburghii (inhabitants of the Old and New Worlds), the three last-named species have well-developed hypocotyls which break through the ground as arches. Now, if we imagine a seedling of the common bean or of P. multiflorus, to behave as its progenitors once did, the hypocotyl (h, Fig. 59), in whatever position the seed may have been buried, would become so much arched that the upper part would be doubled down parallel to the lower part; and

* An instrument devised by Sachs, consisting essentially of a slowly revolving horizontal axis, on which the plant under observation is supported: see 'Würzburg Arbeiten,' 1879, p. 209. [page 94]

this is exactly the kind of curvature which actually occurs in these two plants, though to a much less degree. Therefore we can hardly doubt that their short hypocotyls have retained by inheritance a tendency to curve themselves in the same manner as they did at a former period, when this movement was highly important to them for breaking through the ground, though now rendered useless by the cotyledons being hypogean. Rudimentary structures are in most cases highly variable, and we might expect that rudimentary or obsolete actions would be equally so; and Sachs' curvature varies extremely in amount, and sometimes altogether fails. This is the sole instance known to us of the inheritance, though in a feeble degree, of movements which have become superfluous from changes which the species has undergone.

Rudimentary Cotyledons.--A few remarks on this subject may be here interpolated. It is well known that some dicotyledonous plants produce only a single cotyledon; for instance, certain species of Ranunculus, Corydalis, Chaerophyllum; and we will here endeavour to show that the loss of one or both cotyledons is apparently due to a store of nutriment being laid up in some other part, as in the hypocotyl or one of the two cotyledons, or one of the secondary radicles.

Fig. 60. Citrus aurantium: two young seedlings: c, larger cotyledon; c', smaller cotyledon; h, thickened hypocotyl; r, radicle. In A the epicotyl is still arched, in B it has become erect. [page 95]

With the orange (Citrus aurantium) the cotyledons are hypogean, and one is larger than the other, as may be seen in A (Fig. 60). In B the inequality is rather greater, and the stem has grown between the points of insertion of the two petioles, so that they do not stand opposite to one another; in another case the separation amounted to one-fifth of an inch. The smaller cotyledon of one seedling was extremely thin, and not half the length of the larger one, so that it was clearly becoming rudimentary,* In all these seedlings the hypocotyl was enlarged or swollen.

Fig. 61. Abronia umbellata: seedling twice natural size: c cotyledon; c', rudimentary cotyledon; h, enlarged hypocotyl, with a heel or projection (h') at the lower end; r, radicle.

With Abronia umbellata one of the cotyledons is quite rudimentary, as may be seen (c') in Fig. 61. In this specimen it consisted of a little green flap, 1/84th inch in length, destitute of a petiole and covered with glands like those on the fully developed cotyledon (c). At first it stood opposite to the larger cotyledon; but as the petiole of the latter increased in length and grew in the same line with the hypocotyl (h), the rudiment appeared in older seedlings as if seated some way down the hypocotyl.

Charles Darwin

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