Mendel's principles of heredity: A defence

by William Bateson; Gregor Mendel

I. The Mendelian Principle of Purity of Germ-Cells and the Laws of Heredity Based on Ancestry.

Professor Weldon’s article is entitled “Mendel’s Laws of Alternative Inheritance in Peas.” This title expresses the scope of Mendel’s work and discovery none too precisely and even exposes him to distinct misconception.

To begin with, it says both too little and too much. Mendel did certainly determine Laws of Inheritance in peas—not precisely the laws Professor Weldon has been at the pains of drafting, but of that anon. Having done so, he knew what his discovery was worth. He saw, and rightly, that he had found a principle which must govern a wide area of phenomena. He entitles his paper therefore “Versuche über Pflanzen-Hybriden,” or, Experiments in Plant-Hybridisation.

Nor did Mendel start at first with any particular intention respecting Peas. He tells us himself that he wanted to find the laws of inheritance in hybrids, which he suspected were definite, and that after casting about for a suitable subject, he found one in peas, for the reasons he sets out.

In another respect the question of title is much more important. By the introduction of the word “Alternative” the suggestion is made that the Mendelian principle applies peculiarly to cases of “alternative” inheritance. Mendel himself makes no such limitation in his earlier paper, though perhaps by rather remote implication in the second, to which the reader should have been referred. On the contrary, he wisely abstains from prejudicial consideration of unexplored phenomena.

To understand the significance of the word “alternative” as introduced by Professor Weldon we must go back a little in the history of these studies. In the year 1897 Galton formally announced the Law of Ancestral Heredity referred to in the Introduction, having previously “stated it briefly and with hesitation” in Natural Inheritance, p. 134. In 1898 Professor Pearson published his modification and generalisation of Galton’s Law, introducing a correction of admitted theoretical importance, though it is not in question that the principle thus restated is fundamentally not very different from Galton’s[54]. It is an essential part of the Galton-Pearson Law of Ancestral Heredity that in calculating the probable structure of each descendant the structure of each several ancestor must be brought to account.

Professor Weldon now tells us that these two papers of Galton and of Professor Pearson have “given us an expression for the effects of blended inheritance which seems likely to prove generally applicable, though the constants of the equations which express the relation between divergence from the mean in one generation, and that in another, may require modification in special cases. Our knowledge of particulate or mosaic inheritance, and of alternative inheritance, is however still rudimentary, and there is so much contradiction between the results obtained by different observers, that the evidence available is difficult to appreciate.”

But Galton stated (p. 401) in 1897 that his statistical law of heredity “appears to be universally applicable to bi-sexual descent.” Pearson in re-formulating the principle in 1898 made no reservation in regard to “alternative” inheritance. On the contrary he writes (p. 393) that “if Mr Galton’s law can be firmly established, it is a complete solution, at any rate to a first approximation, of the whole problem of heredity,” and again (p. 412) that “it is highly probable that it [this law] is the simple descriptive statement which brings into a single focus all the complex lines of hereditary influence. If Darwinian evolution be natural selection combined with heredity, then the single statement which embraces the whole field of heredity must prove almost as epoch-making as the law of gravitation to the astronomer[55].”

As I read there comes into my mind that other fine passage where Professor Pearson warns us

“There is an insatiable desire in the human breast to resume in some short formula, some brief statement, the facts of human experience. It leads the savage to ‘account’ for all natural phenomena by deifying the wind and the stream and the tree. It leads civilized man, on the other hand, to express his emotional experience in works of art, and his physical and mental experience in the formulae or so-called laws of science[56].”

No naturalist who had read Galton’s paper and had tried to apply it to the facts he knew could fail to see that here was a definite advance. We could all perceive phenomena that were in accord with it and there was no reasonable doubt that closer study would prove that accord to be close. It was indeed an occasion for enthusiasm, though no one acquainted with the facts of experimental breeding could consider the suggestion of universal application for an instant.

But two years have gone by, and in 1900 Pearson writes[57] that the values obtained from the Law of Ancestral Heredity

“seem to fit the observed facts fairly well in the case of blended inheritance. In other words we have a certain amount of evidence in favour of the conclusion: That whenever the sexes are equipotent, blend their characters and mate pangamously, all characters will be inherited at the same rate,”

or, again in other words, that the Law of Ancestral Heredity after the glorious launch in 1898 has been home for a complete refit. The top-hamper is cut down and the vessel altogether more manageable; indeed she looks trimmed for most weathers. Each of the qualifications now introduced wards off whole classes of dangers. Later on (pp. 487–8) Pearson recites a further list of cases regarded as exceptional. “All characters will be inherited at the same rate” might indeed almost be taken to cover the results in Mendelian cases, though the mode by which those results are arrived at is of course wholly different.

Clearly we cannot speak of the Law of Gravitation now. Our Tycho Brahe and our Kepler, with the yet more distant Newton, are appropriately named as yet to come[58].

But the truth is that even in 1898 such a comparison was scarcely happy. Not to mention moderns, these high hopes had been finally disposed of by the work of the experimental breeders such as Kölreuter, Knight, Herbert, Gärtner, Wichura, Godron, Naudin, and many more. To have treated as non-existent the work of this group of naturalists, who alone have attempted to solve the problems of heredity and species—Evolution, as we should now say—by the only sound method—experimental breeding—to leave out of consideration almost the whole block of evidence collected in Animals and Plants—Darwin’s finest legacy as I venture to declare—was unfortunate on the part of any exponent of Heredity, and in the writings of a professed naturalist would have been unpardonable. But even as modified in 1900 the Law of Ancestral Heredity is heavily over-sparred, and any experimental breeder could have increased Pearson’s list of unconformable cases by as many again.

But to return to Professor Weldon. He now repeats that the Law of Ancestral Heredity seems likely to prove generally applicable to blended inheritance, but that the case of alternative inheritance is for the present reserved. We should feel more confidence in Professor Weldon’s exposition if he had here reminded us that the special case which fitted Galton’s Law so well that it emboldened him to announce that principle as apparently “universally applicable to bi-sexual descent” was one of alternative inheritance—namely the coat-colour of Basset-hounds. Such a fact is, to say the least, ominous. Pearson, in speaking (1900) of this famous case of Galton’s, says that these phenomena of alternative inheritance must be treated separately (from those of blended inheritance)[59], and for them he deduces a proposed “law of reversion,” based of course on ancestry. He writes, “In both cases we may speak of a law of ancestral heredity, but the first predicts the probable character of the individual produced by a given ancestry, while the second tells us the percentages of the total offspring which on the average revert to each ancestral type[60].”

With the distinctions between the original Law of Ancestral Heredity, the modified form of the same law, and the Law of Reversion, important as all these considerations are, we are not at present concerned.

For the Mendelian principle of heredity asserts a proposition absolutely at variance with all the laws of ancestral heredity, however formulated. In those cases to which it applies strictly, this principle declares that the cross-breeding of parents need not diminish the purity of their germ-cells or consequently the purity of their offspring. When in such cases individuals bearing opposite characters, A and B, are crossed, the germ-cells of the resulting cross-bred, AB, are each to be bearers either of character A or of character B, not both.

Consequently when the cross-breds breed either together or with the pure forms, individuals will result of the forms AA, AB, BA, BB[61]. Of these the forms AA and BB, formed by the union of similar germs, are stated to be as pure as if they had had no cross in their pedigree, and henceforth their offspring will be no more likely to depart from the A type or the B type respectively, than those of any other originally pure specimens of these types.

Consequently in such examples it is not the fact that each ancestor must be brought to account as the Galton-Pearson Law asserts, and we are clearly dealing with a physiological phenomenon not contemplated by that Law at all.

Every case therefore which obeys the Mendelian principle is in direct contradiction to the proposition to which Professor Weldon’s school is committed, and it is natural that he should be disposed to consider the Mendelian principle as applying especially to “alternative” inheritance, while the law of Galton and Pearson is to include the phenomenon of blended inheritance. The latter, he tells us, is “the most usual case,” a view which, if supported by evidence, might not be without value.

It is difficult to blame those who on first acquaintance concluded Mendel’s principle can have no strict application save to alternative inheritance. Whatever blame there is in this I share with Professor Weldon and those whom he follows. Mendel’s own cases were almost all alternative; also the fact of dominance is very dazzling at first. But that was two years ago, and when one begins to see clearly again, it does not look so certain that the real essence of Mendel’s discovery, the purity of germ-cells in respect of certain characters, may not apply also to some phenomena of blended inheritance. The analysis of this possibility would take us to too great length, but I commend to those who are more familiar with statistical method, the consideration of this question: whether dominance being absent, indefinite, or suppressed, the phenomena of heritages completely blended in the zygote, may not be produced by gametes presenting Mendelian purity of characters. A brief discussion of this possibility is given in the Introduction, p. [31].

Very careful inquiry would be needed before such a possibility could be negatived. For example, we know that the Laws based on Ancestry can apply to alternative inheritance; witness the case of the Basset-hounds. Here there is no simple Mendelian dominance; but are we sure there is no purity of germ-cells? The new conception goes a long way and it may well reach to such facts as these.

But for the present we will assume that Mendel’s principle applies only to certain phenomena of alternative inheritance, which is as far as our warrant yet runs.

No close student of the recent history of evolutionary thought needs to be told what the attitude of Professor Weldon and his followers has been towards these same disquieting and unwelcome phenomena of alternative inheritance and discontinuity in variation. Holding at first each such fact for suspect, then treating them as rare and negligible occurrences, he and his followers have of late come slowly to accede to the facts of discontinuity a bare and grudging recognition in their scheme of evolution[62].

Therefore on the announcement of that discovery which once and for all ratifies and consolidates the conception of discontinuous variation, and goes far to define that of alternative inheritance, giving a finite body to what before was vague and tentative, it is small wonder if Professor Weldon is disposed to criticism rather than to cordiality.

We have now seen what is the essence of Mendel’s discovery based on a series of experiments of unequalled simplicity which Professor Weldon does not venture to dispute.