Conversely the pistils in the flowers which open last sometimes abort; as when they are ready for fertilisation all the pollen has been shed. He further shows by means of a series of gradations amongst the Compositae, that a tendency from the causes just specified to produce either male or female florets, sometimes spreads to all the florets on the same head, and sometimes even to the whole plant (7/5. 'Ueber die Geschlechtsverhaltnisse bei den Compositen' 1869 page 89.); and in this latter case the species becomes dioecious. In those rare instances mentioned in the Introduction, in which some of the individuals of both monoecious and hermaphrodite plants are proterandrous, others being proterogynous, their conversion into a dioecious condition would probably be much facilitated, as they already consist of two bodies of individuals, differing to a certain extent in their reproductive functions.
Dimorphic heterostyled plants offer still more strongly marked facilities for becoming dioecious; for they likewise consist of two bodies of individuals in approximately equal numbers, and what probably is more important, both the male and female organs differ in the two forms, not only in structure but in function, in nearly the same manner as do the reproductive organs of two distinct species belonging to the same genus. Now if two species are subjected to changed conditions, though of the same nature, it is notorious that they are often affected very differently; therefore the male organs, for instance, in one form of a heterostyled plant might be affected by those unknown causes which induce abortion, differently from the homologous but functionally different organs in the other form; and so conversely with the female organs. Thus the great difficulty before alluded to is much lessened in understanding how any cause whatever could lead to the simultaneous reduction and ultimate suppression of the male organs in half the individuals of a species, and of the female organs in the other half, whilst all were subjected to exactly the same conditions of life.
That such reduction or suppression has occurred in some heterostyled plants is almost certain. The Rubiaceae contain more heterostyled genera than any other family, and from their wide distribution we may infer that many of them became heterostyled at a remote period, so that there will have been ample time for some of the species to have been since rendered dioecious. Asa Gray informs me that Coprosma is dioecious, and that it is closely allied through Nertera to Mitchella, which as we know is a heterostyled dimorphic species. In the male flowers of Coprosma the stamens are exserted, and in the female flowers the stigmas; so that, judging from the affinities of the above three genera, it seems probable that an ancient short-styled form bearing long stamens with large anthers and large pollen-grains (as in the case of several Rubiaceous genera) has been converted into the male Coprosma; and that an ancient long-styled form with short stamens, small anthers and small pollen-grains has been converted into the female form. But according to Mr. Meehan, Mitchella itself is dioecious in some districts; for he says that one form has small sessile anthers without a trace of pollen, the pistil being perfect; while in another form the stamens are perfect and the pistil rudimentary. (7/6. 'Proceedings of the Academy of Sciences of Philadelphia' July 28, 1868 page 183.) He adds that plants may be observed in the autumn bearing an abundant crop of berries, and others without a single one. Should these statements be confirmed, Mitchella will be proved to be heterostyled in one district and dioecious in another.
Asperula is likewise a Rubiaceous genus, and from the published description of the two forms of A. scoparia, an inhabitant of Tasmania, I did not doubt that it was heterostyled; but on examining some flowers sent me by Dr. Hooker they proved to be dioecious. The male flowers have large anthers and a very small ovarium, surmounted by a mere vestige of a stigma without any style; whilst the female flowers possess a large ovarium, the anthers being rudimentary and apparently quite destitute of pollen.