A phenomenon sometimes termed irregularity of dominance has been investigated in a few cases. In certain breeds of poultry such as Dorkings there occurs an extra toe directed backwards like the hallux (cf. Fig. 15). In some families this character behaves as an ordinary dominant to the normal, giving the expected 3 : 1 ratio in F2. But in other families similarly bred the proportions of birds with and without the extra toe appear to be unusual. It has been shown that in such a family some of the birds without the extra toe may nevertheless transmit the peculiarity when mated with birds belonging to strains in which the extra toe never occurs. Though the external appearance of the bird generally affords some indication of the nature of the gametes which it is carrying, this is not always the case. Nevertheless we have reason to suppose that the character segregates in the gametes, though the nature of these cannot always be decided from the appearance of the bird which bears them.

Fowls' feet. On the right a normal and on the left one with an extra toe.

Scheme to illustrate the inheritance of horns in sheep. Heterozygous males shown dark with a white spot, heterozygous females light with a dark spot in the centre.

There are cases in which an apparent irregularity of dominance has been shown to depend upon another character, as in the experiments with sheep carried out by Professor Wood. In these experiments two breeds were crossed, of which one, the Dorset, is horned in both sexes, while the other, the Suffolk, is without horns in either sex. Whichever way the cross was made the resulting F1 generation was similar; the rams were horned, and

the ewes were hornless. In the F2 generation raised from these F1 animals both horned and hornless types appeared in both sexes but in very different proportions. While the horned rams were about three times as numerous as the hornless, this relation was reversed among the females, in which the horned formed only about one-quarter of the total. The simplest explanation of this interesting case is to suppose that the dominance of the horned character depends upon the sex of the animal—that it is dominant in the male but recessive in the female. A pretty experiment was devised for putting this view to the test. If it is true, equal numbers of gametes with and without the horned factor must be produced by the F1 ewes, while the factor should be lacking in all the gametes of the hornless F2 rams. A

hornless ram, therefore, put to a flock of F1 ewes should give rise to equal numbers of zygotes which are heterozygous for the horned character, and of zygotes in which it is completely absent. And since the heterozygous males are horned, while the heterozgyous females are hornless, we should expect from this mating equal numbers of horned and hornless rams, but only hornless ewes. The result of the experiment confirmed this expectation. Of the ram lambs 9 were horned and 8 were hornless, while all the 11 ewe lambs were completely destitute of horns.

Sheep