sulphate to act as an activator, a pigmentary substance is thrown down. The colour of this substance is that of the pigment in the skin or hairs of the animal used. Miss Durham interprets her results as indicating that the skin of these pigmented animals normally secretes one or more tyrosinases. The same result was obtained from the skins of some unhatched chickens. The skins of albinoes gave no results.
Not only have such results been obtained with sponges, Insects, cephalopods, birds and mammals, but Em. Bourquelot and G. Bertrand have shown that certain fungi, the tissues of which, when exposed to the air by injury, become immediately coloured, do so owing to the action of tyrosinase upon one or more chromogenous substances present in the plant. We may conceive, then, that a pigmented animal owes its colour to the power that certain tissues of its body possess to secrete both tyrosinases and chromogenic substances. And the period at which this process is most active is at birth, or preceding it or immediately succeeding it. In spite of the inquiry being only in its initial stages, there is already good evidence to believe that Cuénot’s theory is correct, and that an albino is an individual whose skin lacks the power to secrete either the ferment or the chromogen. It forms one but not both of these substances.
A moment’s consideration, however, will show that, while an albino may be an individual in which one or more of the complementary bodies of pigmentation are absent, a pigmented animal is something more than an individual which carries all the factors necessary for the development of colour. For it must be borne in mind that animals are not only coloured but the colour is arranged in a more or less definite pattern. The wild mouse, rat and rabbit are self-coloured, but the domesticated forms include various piebald patterns, such as spotted forms among mice, and the familiar black and white hooded and dorsal-striped pattern of some tame rats.
Colour, therefore, must be correlated with some determinant (determining factor) for pattern, and it cannot, therefore, exist alone in an animal’s coat. And we must conceive that each kind of pattern—the self, the spotted, the striped, the hooded and all others—has its own special determinant. Given the presence of all the necessary determinants for the development of pigment in a mammal’s coat, some or all of the hairs may bear this pigment according to the pattern determinants, or absence of pattern determinants, which the cells of the hair papillae carry. And this brings us to the question as to whether in a piebald animal the pigmented hairs are in any way different from the pigmentless or white hairs. No adequate investigation of this subject has yet been made, but some observations made by the author of this article, on the piebald black and white rat, show that differences connected with the microscopic structure exist.
There is thus evidence that colour is correlated with other factors which determine pattern. And this leads to the inquiry as to whether albinoes ever exhibit evidence that they carry the pattern determinants in the absence of those for pigmentation. For it is to be expected a priori that, since albinoes were derived from pigmented progenitors and may at any time appear, side by side with pigmented brothers, in a litter from pigmented parents, they would be carrying the pattern determinants of some one or other of their pigmented ancestors. Now we know, from the numerous experiments in heredity which have resulted since the rediscovery of Mendel’s principles, that an individual may carry a character in one of two conditions. It may be carried as a somatic character, when it will be visible in the body tissues, or it may be carried as a gametic character, and its presence can only then be detected in subsequent generations, by adequately devised breeding tests.
With regard to pattern, the evidence is now clear that albinoes may carry the determinants in both these ways. So far as they are carried gametically, i.e. by the sex-cells, it has been shown by Cuénot and G. M. Allen for mice, by C. C. Hurst for rabbits, and by L. Doncaster and G. P. Mudge for rats, that in a cross between a coloured individual of known gametic purity and an albino, the individuals of the progeny in either the first or second, or both generations, may differ, and that the difference in some cases wholly depends upon the albino used. It has been shown that the individuals in such an offspring may bear patterns which never occurred in the ancestry of the coloured parent, but did in that of the albino; and, moreover, if the same coloured parent be mated with another individual, either albino or coloured, that their offspring may never contain members bearing such patterns. The particular pattern will only appear when the coloured parent is mated with the particular albino. And yet the albino itself shows no somatic pattern or pigment. So clear is the evidence on this point that any one adequately acquainted at first hand with the phenomena, by employing an albino of known gametic structure and mating it with a coloured individual, also of known gametic constitution, could predict the result.
With respect to albinoes carrying pattern as a visible somatic character, i.e. in the body cells, no definite evidence has as yet been published. But W. Haacke has described a single albino rat, in which he states that the hairs of the shoulder and mid-dorsal regions were of a different texture from those of the rest of the body. And it is possible that this albino, had it developed colour, would have been of the piebald pattern. But the author of this article has quite recently reared some albinoes in which the familiar shoulder hood and dorsal stripe of the piebald rat is perfectly obvious, in spite of the absence of the slightest pigmentation. The hairs which occupy the region which in the pigmented individual is black, are longer, thinner and more widely separated than those in the regions which are white. As a result of this, the pink skin is quite visible where these hairs occur, but elsewhere it is invisible. Thus these albinoes exhibit a pattern of pink skin similar in form with the black pattern of the piebald rat. Moreover, some of the albinoes possess these particular “pattern” hairs all over the body and obviously such individuals are carrying the self pattern. There are other details into which we cannot here enter, but which support the interpretation put upon these facts, i.e. that these particular albinoes are carrying in the soma the pattern determinants simultaneously with the absence of some of the factors for pigmentation.
Not only do albinoes thus carry the determinants for pattern, but it has been known for some time that they also carry gametically, but never visible somatically, the determinants for either the ferment or the chromogen for one or more colours. L. Cuénot was the first to show this for albino mice. He was able by appropriate experiments to demonstrate that when an albino is derived (extracted) from a coloured ancestry, and is then crossed with a coloured individual, both the colour of the pigmented parent and of the pigmented ancestry of the albino may appear among the individuals of the offspring.
Immediately subsequent to Cuénot, G. M. Allen in America demonstrated the same fact upon the same species of rodents. C. C. Hurst, more recently, has shown that albino rabbits. whether pure bred for eight generations at least, or extracted from pigmented parents, may carry the determinants for black or for black and grey. In this latter case the determinants for black are carried by separate gametes from those carrying grey, and the two kinds of sex-cells exist in approximately equal numbers. This is likewise true of albino mice when they carry the determinants for more than one colour.
Since Hurst’s work, L. Doncaster and G. P. Mudge have both shown that albino rats also carry in a latent condition the determinants for black or grey. The experiments of the latter author show that, if a gametically pure black rat be crossed with an albino derived from a piebald black and white ancestry, all the offspring in successive litters will be black; but if the same black parent be crossed with albinoes extracted from parents of which One or both are grey, then both grey and black members will appear in the successive litters.
The proportions in which the various coloured individuals appear are approximately those demanded by the Mendelian principle of gametic purity and segregation. Cuénot and Hurst have also shown that when albinoes of one colour extraction are crossed with albinoes of another colour extraction the segregation of the colour determinants in the gametogenesis of the albinoes takes place in precisely the same way that it does in the
