[12] ‘Proc. Acad. Nat. Sc. Philadelphia,’ June 16th, 1874, and July 23rd, 1875.
[13] See F. Elfving’s interesting paper in ‘Arbeiten Bot. Institut., in Würzburg,’ vol. ii. 1880, p. 489. Carl Kraus (Triesdorf) had previously observed (‘Flora,’ 1878, p. 324) that the underground shoots of Triticum repens bend vertically up when the parts above ground are removed, and when the rhizomes are kept partly immersed in water.
No explanation has hitherto been attempted of such cases as the foregoing,—namely, of secondary radicles growing vertically downwards, and of lateral shoots growing vertically upwards, after the amputation of the primary radicle or of the leading shoot. The following considerations give us, as we believe, the clue. Firstly, any cause which disturbs the constitution[[14]] is apt to induce reversion; such as the crossing of two distinct races, or a change of conditions, as when domestic animals become feral. But the case which most concerns us, is the frequent appearance of peloric flowers on the summit of a stem, or in the centre of the inflorescence,—parts which, it is believed, receive the most sap; for when an irregular flower becomes perfectly regular or peloric, this may be attributed, at least partly, to reversion to a primitive and normal type. Even the position of a seed at the end of the capsule sometimes gives to the seedling developed from it a tendency to revert. Secondly, reversions often occur by means of buds, independently of reproduction by seed; so that a bud may revert to the character of a former state many bud-generations ago. In the case of animals, reversions may occur in the individual with advancing age. Thirdly and lastly, radicles when they first protrude from the seed are always geotropic, and plumules or shoots almost always apogeotropic. If then any cause, such as an increased flow of sap or the presence of mycelium, disturbs the constitution of a lateral shoot or of a secondary radicle, it is apt to revert to its primordial state; and it becomes either apogeotropic or geotropic, as the case may be, and consequently grows either vertically upwards or downwards. It is indeed possible, or even probable, that this tendency to reversion may have been increased, as it is manifestly of service to the plant.
[14] The facts on which the following conclusions are founded are given in ‘The Variation of Animals and Plants under Domestication,’ 2nd edit. 1875. On the causes leading to reversion see chap. xii. vol. ii. and p. 59, chap. xiv. On peloric flowers, chap. xiii. p. 32; and see p. 337 on their position on the plant. With respect to seeds, p. 340. On reversion by means of buds, p. 438, chap. xi. vol. i.
A SUMMARY OF CHAPTER.
A part or organ may be called sensitive, when its irritation excites movement in an adjoining part. Now it has been shown in this chapter, that the tip of the radicle of the bean is in this sense sensitive to the contact of any small object attached to one side by shellac or gum-water; also to a slight touch with dry caustic, and to a thin slice cut off one side. The radicles of the pea were tried with attached objects and caustic, both of which acted. With Phaseolus multiflorus the tip was hardly sensitive to small squares of attached card, but was sensitive to caustic and to slicing. The radicles of Tropaeolum were highly sensitive to contact; and so, as far as we could judge, were those of Gossypium herbaceum, and they were certainly sensitive to caustic. The tips of the radicles of Cucurbita ovifera were likewise highly sensitive to caustic, though only moderately so to contact. Raphanus sativus offered a somewhat doubtful case. With Æsculus the tips were quite indifferent to bodies attached to them, though sensitive to caustic. Those of Quercus robur and Zea mays were highly sensitive to contact, as were the radicles of the latter to caustic. In several of these cases the difference in sensitiveness of the tip to contact and to caustic was, as we believe, merely apparent; for with Gossypium, Raphanus, and Cucurbita, the tip was so fine and flexible that it was very difficult to attach any object to one of its sides. With the radicles of Æsculus, the tips were not at all sensitive to small bodies attached to them; but it does not follow from this fact that they would not have been sensitive to somewhat greater continued pressure, if this could have been applied.
The peculiar form of sensitiveness which we are here considering, is confined to the tip of the radicle for a length of from 1 mm. to 1.5 mm. When this part is irritated by contact with any object, by caustic, or by a thin slice being cut off, the upper adjoining part of the radicle, for a length of from 6 or 7 to even 12 mm., is excited to bend away from the side which has been irritated. Some influence must therefore be transmitted from the tip along the radicle for this length. The curvature thus caused is generally symmetrical. The part which bends most apparently coincides with that of the most rapid growth. The tip and the basal part grow very slowly and they bend very little.
Considering the widely separated position in the vegetable series of the several above-named genera, we may conclude that the tips of the radicles of all, or almost all, plants are similarly sensitive, and transmit an influence causing the upper part to bend. With respect to the tips of the secondary radicles, those of Vicia faba, Pisum sativum, and Zea mays were alone observed, and they were found similarly sensitive.
In order that these movements should be properly displayed, it appears necessary that the radicles should grow at their normal rate. If subjected to a high temperature and made to grow rapidly, the tips seem either to lose their sensitiveness, or the upper part to lose the power of bending. So it appears to be if they grow very slowly from not being vigorous, or from being kept at too low a temperature; also when they are forced to germinate in the middle of the winter.
The curvature of the radicle sometimes occurs within from 6 to 8 hours after the tip has been irritated, and almost always within 24 h., excepting in the case of the massive radicles of Æsculus. The curvature often amounts to a rectangle,—that is, the terminal part bends upwards until the tip, which is but little curved, projects almost horizontally. Occasionally the tip, from the continued irritation of the attached object, continues to bend up until it forms a hook with the point directed towards the zenith, or a loop, or even a spire. After a time the radicle apparently becomes accustomed to the irritation, as occurs in the case of tendrils, for it again grows downwards, although the bit of card or other object may remain attached to the tip. It is evident that a small object attached to the free point of a vertically suspended radicle can offer no mechanical resistance to its growth as a whole, for the object is carried downwards as the radicle elongates, or upwards as the radicle curves upwards. Nor can the growth of the tip itself be mechanically checked by an object attached to it by gum-water, which remains all the time perfectly soft. The weight of the object, though quite insignificant, is opposed to the upward curvature. We may therefore conclude that it is the irritation due to contact which excites the movement. The contact, however, must be prolonged, for the tips of 15 radicles were rubbed for a short time, and this did not cause them to bend. Here then we have a case of specialised sensibility, like that of the glands of Drosera; for these are exquisitely sensitive to the slightest pressure if prolonged, but not to two or three rough touches.