The first step in the classification of substantive variations is therefore to determine which are due to the addition of new elements or factors, and which are produced by the omission of old ones. A priori there is no valid criterion by which this can be known, and actual experiments in analytical breeding can alone provide the knowledge required. Some very curious results have by this method been obtained, which throw an altogether unexpected light on these problems. For example, in order that the remarkable development of mesoblastic black pigment characteristic of the Silky Fowl should be developed, it is practically certain that two distinct variations from such a type as Gallus bankiva must have occurred. I assume, as is reasonable, that G. bankiva has genetic properties similar to those of the Brown Leghorn breed which has been used in the experiments which Mr. Punnett and I have conducted. Gallus bankiva was not available but the Brown Leghorn agrees with it very closely in colouration, and probably in the general physiology of its pigmentation. Setting aside the various structural differences between the two breeds, the Silky is immediately distinguished from the Leghorn by the fact that the skin of the whole body including that of the face and comb appears to be of a deep purplish colour. The face and comb of the Leghorn are red and the skin of the body is whitish yellow. On examination it is found that the purple colour of the Silky is in reality due to the distribution of a deep black pigment in the mesoblastic membranes throughout the body. The somatopleura, the pleura, pia mater, the dermis, and in most organs the connective tissue and the sheaths of the blood-vessels, are thus impregnated with black. No such pigmentation exists in the Leghorn. As the result of an elaborate series of experimental matings we have proved that the distinction between the Leghorn and the Silky consists primarily in the fact that the Silky possesses a pigment-producing factor, P, which is not present in the Leghorn.

This variation must undoubtedly have been one of addition. But besides this there is another difference of an altogether dissimilar nature; for the Brown Leghorn possesses a factor which has the power of partially or completely restricting the operation of the pigment-producing factor, P. Moreover in respect of this pigment-restricting factor which we may call D, the sexes of the Brown Leghorn differ, for the male is homozygous or DD, but the female is heterozygous, Dd. Thus in order that the black-skinned breed could be evolved from such a type as a Brown Leghorn it must be necessary both that P should be added and that D should drop out. We have not the faintest conception of the process by which either of these events have come to pass, but there is no reasonable doubt that in the evolution of the Silky fowl they did actually happen.

We may anticipate that numerous interdependences of this kind will be discovered.

Before any indisputable progress can be made with the problem of evolution it is necessary that we should acquire some real knowledge of the genesis of that class of phenomena which formed the subject of the last chapter. So long as the process of division remains entirely mysterious we can form no conception even of the haziest sort as to the nature of living organisms, or of the proximate causes which determine their forms, still less can we attempt any answer to those remoter questions of origin and destiny which form the subject of the philosopher's contemplation. It is in no spirit of dogmatism that I have ventured to indicate the direction in which I look for a solution, though I have none to offer. It may well be that before any solution is attained, our knowledge of the nature of unorganised matter must first be increased. For a long time yet we may have to halt, but we none the less do well to prepare ourselves to utilise any means of advance that may be offered, by carefully reconnoitering the ground we have to traverse. The real difficulty which blocks our progress is ignorance of the nature of division, or to use the more general term, of repetition.

Let us turn to the more familiar problem of the causes of variation. Now since variation consists as much in meristic change as in alteration in substance or material, there is one great range of problems of causation from which we are as yet entirely cut off. We know nothing of the causation of division, and we have scarcely an observation, experiment or surmise touching the causes by which the meristic processes may be altered.

Of the way in which variations in the substantive composition of organisms are caused we have almost as little real evidence, but we are beginning to know in what such variations must consist. These changes must occur either by the addition or loss of factors.

We must not lose sight of the fact that though the factors operate by the production of enzymes, of bodies on which these enzymes can act, and of intermediary substances necessary to complete the enzyme-action, yet these bodies themselves can scarcely be themselves genetic factors, but consequences of their existence. What then are the factors themselves? Whence do they come? How do they become integral parts of the organism? Whence, for example, came the power which is present in a White Leghorn of destroying—probably reducing—the pigment in its feathers? That power is now a definite possession of the breed, present in all its germ-cells, male and female, taking part in their symmetrical divisions, and passed on equally to all as much as is the protoplasm or any other attribute of the breed. From the body of the bird the critical and efficient substance could in all likelihood be isolated by suitable means, just as the glycogen of the liver can be. But even when this extraction has been accomplished and the reducing body isolated, we shall know no more than we did before respecting the mode by which the power to produce it was conferred on the fowl, any more than we know how the walls of its blood-vessels acquired the power to form a fibrin-ferment.

It is when the scope of such considerations as this are fully grasped that we realise the fatuousness of the conventional treatment which the problem of the causes of variation commonly receives. Environmental change, chemical injury, differences in food supply, in temperature, in moisture, or the like have been proposed as "causes." Admitting as we must do, that changes may be produced—usually inhibitions of development—by subjecting living things to changes in these respects, how can we suppose it in the smallest degree likely that very precise, new, and adaptative powers can be conferred on the germs by such treatment? Reports of positive genetic consequences observed comparable with those I have mentioned, become from time to time current. We should I think regard them with the gravest doubt. Few, so far as I am aware, have ever been confirmed, though clear and repeated confirmation should be demanded before we suffer ourselves at all to build upon such evidence. In a subsequent chapter some of these cases will be considered in detail.

In no class of cases would the transmission of an acquired character superficially appear so probable as in those where power of resisting the attack of a pathogenic organism is acquired in the lifetime of the zygote. The possession of such a power is moreover a distinction comparable with those which differentiate varieties and species. It is due to the development in the blood of specific substances which pervade the whole fluid. This development is exactly one of those "appropriate responses to stimuli" which naturalists who incline to regard adaptation as a direct consequence of an environmental influence might most readily invoke as an illustration of their views. And yet all evidence is definitely unfavourable to the suggestion of an inheritance of the acquired power of resistance. Such change as can be perceived in the virulence of the attacks on successive generations may be most easily regarded as due to the extermination of the more susceptible strains, and perhaps in some measure to variation in the invading organisms themselves, an "acquired character" of quite different import.

The specific "anti-body" may have been produced in response to the stimulus of disease, but the power to produce it without this special stimulus is not included in the germ-cells any more than a pigment. All that they bear is the power to produce the anti-bodies when the stimulus is applied.