[12] ‘Arbeiten des bot. Instit. Würzburg,’ vol. i. 1873, p. 403.
The above movement in the bean was repeatedly seen by us; but our observations were made chiefly on Phaseolus multiflorus, the cotyledons of which are likewise hypogean. Some seedlings with well-developed radicles were first immersed in a solution of permanganate of potassium; and, judging from the changes of colour (though these were not very clearly defined), the hypocotyl is about .3 inch in length. Straight, thin, black lines of this length were now drawn from the bases of the short petioles along the hypocotyls of 23 germinating seeds, which were pinned to the lids of jars, generally with the hilum downwards, and with their radicles pointing to the centre of the earth. After an interval of from 24 to 48 hours the black lines on the hypocotyls of 16 out of the 23 seedlings became distinctly curved, but in very various degrees (namely, with radii between 20 and 80 mm. on Sachs’ cyclometer) in the same relative direction as shown at B in Fig. 59. As geotropism will obviously tend to check this curvature, seven seeds were allowed to germinate with proper precautions for their growth in a klinostat,[[13]] by which means geotropism was eliminated. The position of the hypocotyls was observed during four successive days, and they continued to bend towards the hilum and lower surface of the seed. On the fourth day they were deflected by an average angle of 63° 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.
[13] 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.
Fig. 59. Vicia faba: germinating seeds, suspended in damp air: A, with radicle growing perpendicularly downwards; B, the same bean after 24 hours and after the radicle has curved itself; r. radicle; h, short hypocotyl; e, epicotyl appearing as a knob in A and as an arch in B; p, petiole of the cotyledon, the latter enclosed within the seed-coats.
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 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. 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[[14]] In all these seedlings the hypocotyl was enlarged or swollen.
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.
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.
[14] In Pachira aquatica, as described by Mr. R. I. Lynch (‘Journal Linn. Soc. Bot.’ vol. xvii. 1878, p. 147), one of the hypogean cotyledons is of immense size; the other is small and soon falls off; the pair do not always stand opposite. In another and very different water-plant, ‘Trapa natans’, one of the cotyledons, filled with farinaceous matter, is much larger than the other, which is scarcely visible, as is stated by Aug. de Candolle, ‘Physiologie Veg.’ tom. ii. p. 834, 1832.
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. With Abronia arenaria there is a similar rudiment, which in one specimen was only 1/100th and in another 1/60th inch in length; it ultimately appeared as if seated halfway down the hypocotyl. In both these species the hypocotyl is so much enlarged, especially at a very early age, that it might almost be called a corm. The lower end forms a heel or projection, the use of which will hereafter be described.