Formation of pollen within the ovules.—This has now been recorded in two instances by Mr. S. J. A. Salter in Passiflora cærulea and in P. palmata,[185] and by the author in Rosa arvensis.[186]
Fig. 99.—Pollen within the ovule of Passiflora (after Salter).
In the case of the passion-flower there were various malformations in the ovaries, which were all more or less split open at the distal end, indicating a tendency towards dialysis. The pollen-bearing ovules were borne on the edges of these ovaries, and presented various intermediate conditions between anthers and ovules, commencing at the distal extremity of the carpel with a bi-lobed anther, and passing in series to the base of the ovary, an antheroid body of ovule-like form, a modified ovule containing pollen, an ovule departing from a perfectly natural condition only in the development of a few grains of pollen in its nucleus, and, finally, a perfect, normal ovule.
In the flowers of the Rose the stamens exhibited almost every conceivable gradation between their ordinary form and that of the carpels, while some of the ovules contained pollen in greater or less abundance. Speaking generally, the most common state of things in these flowers was the occurrence on the throat of the calyx, in the position ordinarily occupied by the stamens, and sometimes mingled with those organs, of twisted, ribbon-like filaments, which bore about the centre one or more pendulous, anatropous ovules on their margins. Immediately above the latter organs were the anther-lobes, more or less perfectly developed, and surmounting these a long style, terminating in a fringed, funnel-shaped stigma. Sometimes the ovules were perfect, at other times the nucleus protruded through the foramen, while in a third set the nucleus was included within the tegument, the ovules having in all respects their natural external conformation, containing, however, not only pollen-grains, but also a layer of those peculiar spheroidal cells, including a fibrous deposit, which are among the normal constituents of the anther. In one case, where the coat of the ovule was imperfect, and allowed the nucleus to protrude, the pollen was evidently contained within the central mass of the structure. In this instance the fibrous cells were not detected, these being only found in cases where the investment of the ovule was perfect; and hence it seems likely that the fibrous cells were part of the coat of the ovule, while the pollen was formed within the nucleus. In no case was any trace of embryo sac to be seen.
The main interest, as Mr. Salter remarks, in these cases is physiological; so far as structure alone is concerned, there does not appear any reason why pollen-grains should not be developed in any portion of the plant; the mother cells in which the pollen is formed not differing, to all outward appearance, from any other cells, unless it be in size.
The fundamental unity of construction in all the organs of plants could hardly be better illustrated than by these cases; while, in spite of their exceptional nature, they must be of great interest physiologically, as showing the wide limits of possible variation which thus may even involve the sex, "for an ovule to develop pollen within its interior," says Mr. Salter, "is equivalent to an ovum in an animal being converted into a capsule of spermatozoa. It is a conversion of germ into sperm, the most complete violation of individuality and unity of sex. * * * * The occurrence of an antheroid ovule and a normal ovule on the same carpellary leaf realises the simplest and the most absolute form of hermaphroditism."
It must, however, be remarked that the term substitution would be preferable to conversion. There is, at present, no evidence to show that the germinal vesicles were present in these cases; on the other hand, it seems most probable that they were not, so that the presence of the pollen-cells must be considered as simply adventitious. It can hardly be that they were, in the first instance, germinal vesicles, which, in course of time, became so modified as to assume the appearance of pollen-grains. Between the nucleus of the ovule and the tubercle of cellular tissue constituting the primordial anther, there is little or no difference, so that it may be said that, for a time, there is no distinction of sex in the nascent flower, but as development goes on, the difference becomes perceptible. It cannot at present be stated what precise circumstances induce the one mass to form mother-cells and pollen-grains, and the other to develop an embryo sac and germinal vesicles. Position and external circumstances may have some indirect effect, and it may, perhaps, be significant that in all the instances of polliniferous ovules, the ovular structures have been exposed on an open carpel or otherwise, in place of being confined within the cavity of a closed ovary, as under ordinary circumstances. Even among Conifers the ovuligerous scales are so closely packed that there is little or no exposure of the ovules. But, apart from all speculative notions as to the relation between the structure and functions of the anther and of the ovule respectively, and of the possibility or the reverse of parthenogenesis, it will clearly be necessary in any future alleged occurrence of the latter phenomenon to ascertain whether any or all of the apparent ovules are, or are not, anthers in disguise.
Homomorphic flowers of "Compositæ."—In a large section of the Compositæ there is, as is well known, a distinction between the florets of the "disc" and those of the "ray," the latter being ligulate, the former tubular.