It is known[[26]] that there is no difference in structure between the upper and lower halves of the pulvini of leaves, sufficient to account for their upward or downward movements. In this respect cotyledons offer an unusually good opportunity for comparing the structure of the two halves; for the cotyledons of Oxalis Valdiviana rise vertically at night, whilst those of O. rosea sink vertically; yet when sections of their pulvini were made, no clear difference could be detected between the corresponding halves of this organ in the two species which move so differently. With O. rosea, however, there were rather more cells in the lower than in the upper half, but this was likewise the case in one specimen of O. Valdiviana. the cotyledons of both species (3½ mm. in length) were examined in the morning whilst extended horizontally, and the upper surface of the pulvinus of O. rosea was then wrinkled transversely, showing that it was in a state of compression, and this might have been expected, as the cotyledons sink at night; with O. Valdiviana it was the lower surface which was wrinkled, and its cotyledons rise at night.
[26] Pfeffer, ‘Die Period. Bewegungen,’ 1875, p. 157.
Trifolium is a natural genus, and the leaves of all the species seen by us are pulvinated; so it is with the cotyledons of T. subterraneum and strictum, which stand vertically at night; whereas those of T. resupinatum exhibit not a trace of a pulvinus, nor of any nocturnal movement. This was ascertained by measuring the distance between the tips of the cotyledons of four seedlings at mid-day and at night. In this species, however, as in the others, the first-formed leaf, which is simple or not trifoliate, rises up and sleeps like the terminal leaflet on a mature plant.
In another natural genus, Oxalis, the cotyledons of O. Valdiviana, rosea, floribunda, articulata, and sensitiva are pulvinated, and all move at night into an upward or downward vertical position. In these several species the pulvinus is seated close to the blade of the cotyledon, as is the usual rule with most plants. Oxalis corniculata (var. Atro-purpurea) differs in several respects; the cotyledons rise at night to a very variable amount, rarely more than 45°; and in one lot of seedlings (purchased under the name of O. tropaeoloides, but certainly belonging to the above variety) they rose only from 5° to 15° above the horizon. The pulvinus is developed imperfectly and to an extremely variable degree, so that apparently it is tending towards abortion. No such case has hitherto, we believe, been described. It is coloured green from its cells containing chlorophyll; and it is seated nearly in the middle of the petiole, instead of at the upper end as in all the other species. The nocturnal movement is effected partly by its aid, and partly by the growth of the upper part of the petiole as in the case of plants destitute of a pulvinus. From these several reasons and from our having partially traced the development of the pulvinus from an early age, the case seems worth describing in some detail.
When the cotyledons of O. corniculata were dissected out of a seed from which they would soon have naturally emerged, no trace of a pulvinus could be detected; and all the cells forming the short petiole, 7 in number in a longitudinal row, were of nearly equal size. In seedlings one or two days old, the pulvinus was so indistinct that we thought at first that it did not exist; but in the middle of the petiole an ill-defined transverse zone of cells could be seen, which were much shorter than those both above and below, although of the same breadth with them. They presented the appearance of having been just formed by the transverse division of longer cells; and there can be little doubt that this had occurred, for the cells in the petiole which had been dissected out of the seed averaged in length 7 divisions of the micrometer (each division equalling .003 mm.), and were a little longer than those forming a well-developed pulvinus, which varied between 4 and 6 of these same divisions. After a few additional days the ill-defined zone of cells becomes distinct, and although it does not extend across the whole width of the petiole, and although the cells are of a green colour from containing chlorophyll, yet they certainly constitute a pulvinus, which as we shall presently see, acts as one. These small cells were arranged in longitudinal rows, and varied from 4 to 7 in number; and the cells themselves varied in length in different parts of the same pulvinus and in different individuals. In the accompanying figures, A and B (Fig. 64), we have views of the epidermis[[27]] in the middle part of the petioles of two seedlings, in which the pulvinus was for this species well developed. They offer a striking contrast with the pulvinus of O. rosea (see former Fig. 63), or of O. Valdiviana. With the seedlings, falsely called O. tropaeoloides, the cotyledons of which rise very little at night, the small cells were still fewer in number and in parts formed a single transverse row, and in other parts short longitudinal rows of only two or three. Nevertheless they sufficed to attract the eye, when the whole petiole was viewed as a transparent object beneath the microscope. In these seedlings there could hardly be a doubt that the pulvinus was becoming rudimentary and tending to disappear; and this accounts for its great variability in structure and function.
[27] Longitudinal sections show that the forms of the epidermic cells may be taken as a fair representation of those constituting the pulvinus.
Fig. 64. Oxalis corniculata: A and B the almost rudimentary pulvini of the cotyledons of two rather old seedlings, viewed as transparent objects. Magnified 50 times.
In the following Table some measurements of the cells in fairly well-developed pulvini of O. corniculata are given:—
Seedling 1 day old, with cotyledon 2.3 mm. in length. Divisions of Micrometer.[[28]] Average length of cells of pulvinus..................................................6 to 7 Length of longest cell below the pulvinus..................................... 13 Length of longest cell above the pulvinus...................................... 20
Seedling 5 days old, cotyledon 3.1 mm. in length, with the pulvinus quite distinct. Average length of cells of pulvinus.................................................. 6 Length of longest cell below the pulvinus..................................... 22 Length of longest cell above the pulvinus...................................... 40