without M and the ova containing F. In other words we must on this view suppose that fertilisations between certain forms of gametes, even if they can occur, are incapable of giving rise to zygotes with the capacity for further development. If we admit this supposition, the scheme just given will cover such cases as those of the currant moth and the fowl, equally as well as that of the pomace fly. In the former there is repulsion between either the grossulariata factor and F, or else between the pigment inhibitor factor and F, while in the latter there is repulsion between the factor for red eye and M.
Scheme to illustrate the probable mode of inheritance of colour-blindness. The dark signs represent affected individuals. A black dot in the centre denotes an unaffected female who is capable of transmitting the condition to her sons.
Whatever the merits or demerits of such a scheme it certainly does offer an explanation of a peculiar form of sex limited inheritance in man. It has long been a matter of common knowledge that colour-blindness is much more common among men than among women, and also that unaffected women can transmit it to their sons. At first sight the case is not unlike that of the sheep, where the horned character is apparently dominant in the male but recessive in the female. The hypothesis that the colour-blind condition is due to the presence of an extra factor as compared with the normal, and that a single dose of it will produce
colour-blindness in the male but not in the female, will cover a good many of the observed facts (cf. Fig. 26). Moreover, it serves to explain the remarkable fact that all the sons of colour-blind women are also colour-blind. For a woman cannot be colour-blind unless she is homozygous for the colour-blind factor, in which case all her children must get a single dose of it even if she marries a normal male. And this is sufficient to produce colour-blindness in the male, though not in the female.
But there is one notable difference in this case as compared with that of the sheep. When crossed with pure hornless ewes the heterozygous horned ram transmits the horned character to half his male offspring (cf. p. [71]). But the heterozygous colour-blind man does not behave altogether like a sheep, for he apparently does not transmit the colour-blind condition to any of his male offspring. If, however, we suppose that the colour-blind factor is repelled by the factor for maleness, the amended scheme will cover the observed facts. For, denoting the colour-blind factor by X, the gametes produced by the colour-blind male are of two sorts only, viz. Mfx and mfX. If he marries a normal woman (Ffmmxx), the spermatozoa Mfx unite with ova fmx to give normal males, while the spermatozoa mfX unite with ova Fmx to give females which are heterozygous for the colour-blind factor. These daughters are themselves normal, but transmit the condition to about half their sons.
The attempt to discover a simple explanation of the nature of sex has led us to assume that certain combinations between gametes are incapable of giving rise to zygotes which can develop further. In the various cases hitherto considered there is no reason to suppose that anything of the sort occurs, or that the different gametes are otherwise than completely fertile one with another. One peculiar case, however, has been known for several years in which some of the gametes are apparently incapable of uniting to produce offspring. Yellow in the mouse is dominant to agouti, but hitherto a homozygous yellow has never been met with. The yellows from families where only yellows and agoutis occur produce, when bred together, yellows and agoutis in the ratio 2 : 1. If it were an ordinary Mendelian case the ratio should be 3 : 1, and one out of every three yellows so bred should be homozygous and give only yellows when crossed with agouti. But Cuénot and others have shown that all of the yellows are heterozygous, and when crossed with agoutis give both yellows and agoutis. We are led, therefore, to suppose that an ovum carrying the yellow factor is unproductive if fertilised by a spermatozoon which also bears this factor. In this way alone does it seem possible to explain the deficiency of yellows and the absence of homozygous ones in the families arising from the mating of yellows together. At present, however, it remains the only definite instance among animals in which we have
grounds for assuming that anything in the nature of unproductive fertilisation takes place.[[8]]
If we turn from animals to plants we find a more complicated state of affairs. Generally speaking, the higher plants are hermaphrodite, both ovules and pollen grains occurring on the same flower. Some plants, however like most animals, are of separate sexes, a single plant bearing only male or female flowers. In other plants the separate flowers are either male or female, though both are borne on the same individual. In others, again, the conditions are even more complex, for the same plant may bear flowers of three kinds, viz. male, female, and hermaphrodite. Or it may be that these three forms occur in the same species but in different individuals—female and hermaphrodites in one species; males, females, and hermaphrodites in another. One case, however, must be mentioned as it suggests a possibility which we have not hitherto encountered. In the common English bryony (Bryonia dioica) the sexes are separate, some plants having only male and others only female flowers. In another European species, B. alba, both male and female flowers occur on the same plant. Correns crossed these two species reciprocally, and also fertilised B. dioica by its own male with the following results:—