There can be little doubt that in many cases it is to the third category that the phenomena belong. An indication of the applicability of this reasoning will generally be found in the fact that in such “mule” forms the colour or the shape of the seeds will be recognizably peculiar and proper to the specimens themselves, as distinct from their parents, and we may safely anticipate that when those seeds are grown the plants will show some character which is recognizable as novel. The proof that the reasoning may apply can as yet only be got by finding that the forms in question cannot breed true even after successive selections, but constantly break up into the same series of forms[76].
This conception of the “mule” form, or “hybrid-character” as Mendel called it, though undeveloped, is perfectly clear in his work. He says that the dominant character may have two significations, it may be either a parental character or a hybrid-character, and it must be differentiated according as it appears in the one capacity or the other. He does not regard the character displayed by the hybrid, whether dominant or other, as a thing inherited from or transmitted by the pure parent at all, but as the peculiar function or property of the hybrid. When this conception has been fully understood and appreciated in all its bearings it will be found to be hardly less fruitful than that of the purity of the germ-cells.
The two parents are two—let us say—substances[77] represented by corresponding gametes. These gametes unite to form a new “substance”—the cross-bred zygote. This has its own properties and structure, just as a chemical compound has, and the properties of this new “substance” are not more strictly traceable to, or “inherited” from, those of the two parents than are those of a new chemical compound “inherited” from those of the component elements. If the case be one in which the gametes are pure, the new “substance” is not represented by them, but the compound is again dissociated into its components, each of which is separately represented by gametes.
The character of the cross-bred zygote may be anything. It may be something we have seen before in one or other of the parents, it may be intermediate between the two, or it may be something new. All these possibilities were known to Mendel and he is perfectly aware that his principle is equally applicable to all. The first case is his “dominance.” That he is ready for the second is sufficiently shown by his brief reference to time of flowering considered as a character (p. [65]). The hybrids, he says, flower at a time almost exactly intermediate between the flowering times of the parents, and he remarks that the development of the hybrids in this case probably happens in the same way as it does in the case of the other characters[78].
That he was thoroughly prepared for the third possibility appears constantly through the paper, notably in the argument based on the Phaseolus hybrids, and in the statement that the hybrid between talls and dwarfs is generally taller than the tall parent, having increased height as its “hybrid-character.”
All this Professor Weldon has missed. In place of it he offers us the sententia that no one can expect to understand these phenomena if he neglect ancestry. This is the idle gloss of the scribe, which, if we erase it not thoroughly, may pass into the text.
Enough has been said to show how greatly Mendel’s conception of heredity was in advance of those which pass current at the present day; I have here attempted the barest outline of the nature of the “hybrid-character,” and I have not sought to indicate the conclusions that we reach when the reasoning so clear in the case of the hybrid is applied to the pure forms and their own characters.
In these considerations we reach the very base on which all conceptions of heredity and variation must henceforth rest, and that it is now possible for us to attempt any such analysis is one of the most far-reaching consequences of Mendel’s principle. Till two years ago no one had made more than random soundings of this abyss.
I have briefly discussed these possibilities to assist the reader in getting an insight into Mendel’s conceptions. But in dealing with Professor Weldon we need not make this excursion; for his objection arising from the absence of uniform regularity in dominance is not in point.
The soundness of Mendel’s work and conclusions would be just as complete if dominance be found to fail often instead of rarely. For it is perfectly certain that varieties can be chosen in such a way that the dominance of one character over its antagonist is so regular a phenomenon that it can be used in the way Mendel indicates. He chose varieties, in fact, in which a known character was regularly dominant and it is because he did so that he made his discovery[79]. When Professor Weldon speaks of the existence of fluctuation and diversity in regard to dominance as proof of a “grave discrepancy” between Mendel’s facts and those of other observers[80], he merely indicates the point at which his own misconceptions began.