It is clear that Mendel realized (b) as a possibility, for he says DR was fertilised with the pure forms to test the composition of its egg-cells, but the reciprocal crosses were made to test the composition of the pollen of the hybrids. Readers familiar with the literature will know that both Gärtner and Wichura had in many instances shown that the offspring of crosses in the form (a × b) ♀ × c ♂ were less variable than those of crosses in the form a ♀ × (b × c) ♂, &c. This important fact in many cases is observed, and points to differentiation of characters occurring frequently among the male gametes when it does not occur or is much less marked among the maternal gametes. Mendel of course knew this, and proceeded to test for such a possibility, finding by the result that differentiation was the same in the gametes of both sexes[155].
Of hypotheses (b) and (c) the results of recrossing with the two pure forms dispose; and we can suggest no hypothesis but (a) which gives an acceptable account of the facts.
It is the purity of the “extracted” recessives and the “extracted” dominants—primarily the former, as being easier to recognize—that constitutes the real proof of the validity of Mendel’s principle.
Using this principle we reach immediately results of the most far-reaching character. These theoretical deductions cannot be further treated here—but of the practical use of the principle a word may be said. Where-ever there is marked dominance of one character the breeder can at once get an indication of the amount of trouble he will have in getting his cross-bred true to either dominant or recessive character. He can only thus forecast the future of the race in regard to each such pair of characters taken severally, but this is an immeasurable advance on anything we knew before. More than this, it is certain that in some cases he will be able to detect the “mule” or heterozygous forms by the statistical frequency of their occurrence or by their structure, especially when dominance is absent, and sometimes even in cases where there is distinct dominance. With peas, the practical seedsman cares, as it happens, little or nothing for those simple characters of seed-structure, &c. that Mendel dealt with. He is concerned with size, fertility, flavour, and numerous similar characters. It is to these that Laxton (invoked by Professor Weldon) primarily refers, when he speaks of the elaborate selections which are needed to fix his novelties.
We may now point tentatively to the way in which some even of these complex cases may be elucidated by an extension of Mendel’s principle, though we cannot forget that there are other undetected factors at work.
The value of the appeal to Ancestry.
But it may be said that Professor Weldon’s appeal to ancestry calls for more specific treatment. When he suggests ancestry as “one great reason” for the different properties displayed by different races or individuals, and as providing an account of other special phenomena of heredity, he is perhaps not to be taken to mean any definite ancestry, known or hypothetical. He may, in fact, be using the term “ancestry” merely as a brief equivalent signifying the previous history of the race or individual in question. But if such a plea be put forward, the real utility and value of the appeal to ancestry is even less evident than before.
Ancestry, as used in the method of Galton and Pearson, means a definite thing. The whole merit of that method lies in the fact that by it a definite accord could be proved to exist between the observed characters and behaviour of specified descendants and the ascertained composition of their pedigree. Professor Weldon in now attributing the observed peculiarities of Telephone &c. to conjectural peculiarities of pedigree—if this be his meaning—renounces all that had positive value in the reference to ancestry. His is simply an appeal to ignorance. The introduction of the word “ancestry” in this sense contributes nothing. The suggestion that ancestry might explain peculiarities means no more than “we do not know how peculiarities are to be explained.” So Professor Weldon’s phrase “peas of probably similar ancestral history[156]” means “peas probably similar”; when he speaks of Mendel having obtained his results with “a few pairs of plants of known ancestry[157],” he means “a few pairs of known plants” and no more; when he writes that “the law of segregation, like the law of dominance appears to hold only for races of particular ancestry[158],” the statement loses nothing if we write simply “for particular races.” We all know—the Mendelian, best of all—that particular races and particular individuals may, even though indistinguishable by any other test, exhibit peculiarities in heredity.
But though on analysis those introductions of the word “ancestry” are found to add nothing, yet we can feel that as used by Professor Weldon they are intended to mean a great deal. Though the appeal may be confessedly to ignorance, the suggestion is implied that if we did know the pedigrees of these various forms we should then have some real light on their present structure or their present behaviour in breeding. Unfortunately there is not the smallest ground for even this hope.
As Professor Weldon himself tells us[159], conclusions from pedigree must be based on the conditions of the several ancestors; and even more categorically (p. 244), “The degree to which a parental character affects offspring depends not only upon its development in the individual parent, but on its degree of development in the ancestors of that parent.” [My italics.] Having rehearsed this profession of an older faith Professor Weldon proceeds to stultify it in his very next paragraph. For there he once again reminds us that Telephone, the mongrel pea of recent origin, which does not breed true to seed characters, has yet manifested the peculiar power of stamping the recessive characters on its cross-bred offspring, though pure and stable varieties that have exhibited the same characters in a high degree for generations have not that power. As we now know, the presence or absence of a character in a progenitor may be no indication whatever as to the probable presence of the character in the offspring; for the characters of the latter depend on gametic and not on zygotic differentiation.