The explanation given by Mr Darwin of the evil effects of self-fertilization, viz. the want of sufficient differentiation in the sexual elements[33], would apply with far greater force to cases of parthenogenesis.
In the production of fresh individuals, two circumstances are obviously favourable to the species. (1) That the maximum number possible of fresh individuals should be produced, (2) That the individuals should be as vigorous as possible. Sexual differentiation (even in hermaphrodites) is clearly very inimical to the production of the maximum number of individuals. There can be little doubt that the ovum is potentially capable of developing by itself into a fresh individual, and therefore, unless the absence of sexual differentiation was very injurious to the vigour of the progeny, parthenogenesis would most certainly be a very constant occurrence; and, on the analogy of the arrangements in plants to prevent self-fertilization, we might expect to find some contrivance both in animals and in plants to prevent the ovum developing by itself without fertilization. If my view about the polar cells is correct, the formation of these bodies functions as such a contrivance.
Reproduction by budding or fission has probably arisen as a means of increasing the number of individuals produced, so that the coexistence of asexual with sexual reproduction is to be looked on as a kind of compromise for the loss of the power of rapid reproduction due to the absence of parthenogenesis. In the Arthropoda and Rotifera the place of budding has been taken by parthenogenesis, which may be a frequent, though not always a necessary occurrence, as in various Branchiopoda (Apus, Limnadia, etc.) and Lepidoptera (Psyche helix, etc.); or a regular occurrence for the production of one sex, as in Bees, Wasps, Nematus, etc.; or an occurrence confined to a certain stage in the cycle of development in which all the individuals reproduce their kind parthenogenetically, as in Aphis, Cecidomyia, Gall Insects (Neuroterus, etc.), Daphnia[34].
On my hypothesis the possibility of parthenogenesis, or at any rate its frequency, in Arthropoda and Rotifera is possibly due to the absence of polar cells. In the case of all animals, so far as is known to me, fertilization of the ovum occasionally occurs[35], but there are instances in the vegetable kingdom where so-called parthenogenesis appears to be capable of recurring for an indefinite period. One of the best instances appears to be that of Cœlebogyne, an introduced exotic Euphorbiaceous plant which regularly produces fertile seeds although a male flower never appears. The recent researches of Strasburger have however shewn that in Cœlebogyne and other parthenogenetic flowering plants, embryos are formed by the budding and subsequent development of cells belonging to the ovule. This being the case, it is impossible to assert of these plants that they are really parthenogenetic, for the embryos contained in the seed of a flower which has certainly not been fertilized, may have been formed, not by the development of the ovum, but by budding from the surrounding tissue of the ovule.
The above view with reference to the nature of the polar bodies is not to be regarded as forming more than an hypothesis.
Impregnation of the Ovum.
A far greater amount of certainty has been attained as to the effects of impregnation than as to the changes of the germinal vesicle which precede this, and there appears, moreover, to be a greater uniformity in the series of resulting phenomena.
