Part 33

The numbers 54 : 96 : 45 approach the ratio 1 : 2 : 1 so nearly that there can be no doubt we have here a simple case of Mendelian laws, operating without definite dominance, but rather with blending.

When Laxton speaks of the “remarkably fine but unfixable pea Evolution” we now know for the first time exactly what the phenomenon meant. It, like the “Giant Lavender,” was a “mule” form, not represented by germ-cells, and in each year arose by “self-crossing.”

This is only one case among many similar ones seen in the Chinese Primrose. In others there is no doubt that more complex factors are at work, the subdivision of compound characters, and so on. The history of the “Giant Lavender” goes back many years and is not known with sufficient precision for our purposes, but like all these forms it originated from crossings among the old simple colour varieties of sinensis.

VI. THE ARGUMENT BUILT ON EXCEPTIONS.

So much for the enormous advance that the Mendelian principles already permit us to make. But what does Professor Weldon offer to substitute for all this? Nothing.

Professor Weldon suggests that a study of ancestry will help us. Having recited Tschermak’s exceptions and the great irregularities seen in the Telephone group, he writes:

“Taking these results together with Laxton’s statements, and with the evidence afforded by the Telephone group of hybrids, I think we can only conclude that segregation of seed-characters is not of universal occurrence among cross-bred peas, and that when it does occur, it may or may not follow Mendel’s law.”

Premising that when pure types are used the exceptions form but a small part of the whole, and that any supposed absence of “segregation” may have been variation, this statement is perfectly sound. He proceeds:--

“The law of segregation, like the law of dominance, appears therefore to hold only for races of particular ancestry [my italics]. In special cases, other formulae expressing segregation have been offered, especially by De Vries and by Tschermak for other plants, but these seem as little likely to prove generally valid as Mendel’s formula itself.

“The fundamental mistake which vitiates all work based upon Mendel’s method is the neglect of ancestry, and the attempt to regard the whole effect upon offspring, produced by a particular parent, as due to the existence in the parent of particular structural characters; while the contradictory results obtained by those who have observed the offspring of parents identical in certain characters show clearly enough that not only the parents themselves, but their race, that is their ancestry, must be taken into account before the result of pairing them can be predicted.”

In this passage the Mendelian view is none too precisely represented. I should rather have said that it was from Mendel, first of all men, that we have learnt not to regard the effects produced on offspring “as due to the existence in the parent of particular structural characters.” We have come rather to disregard the particular structure of the parent except in so far as it may give us a guide as to the nature of its gametes.

This indication, if taken in the positive sense--as was sufficiently shown in considering the significance of the “mule” form or “hybrid-character”--we now know may be absolutely worthless, and in any unfamiliar case is very likely to be so. Mendel has proved that the inheritance from individuals of identical ancestry may be entirely different: that from identical ancestry, without new variation, may be produced three kinds of individuals (in respect of each pair of characters), namely, individuals capable of transmitting one type, or another type, or both: moreover that the statistical relations of these three classes of individuals to each other will in a great number of cases be a definite one: and of all this he shows a complete account.

Professor Weldon cannot deal with any part of this phenomenon. He does little more than allude to it in passing and point out exceptional cases. These he suggests a study of ancestry will explain.

As a matter of fact a study of ancestry will give little guide--perhaps none--even as to the probability of the phenomenon of dominance of a character, none as to the probability of normal “purity” of germ-cells. Still less will it help to account for fluctuations in dominance, or irregularities in “purity.”

Ancestry and Dominance.

In a series of astonishing paragraphs (pp. 241–2) Professor Weldon rises by gradual steps, from the exceptional facts regarding occasional dominance of green colour in Telephone to suggest that the whole phenomenon of dominance may be attributable to ancestry, and that in fact one character has no natural dominance over another, apart from what has been created by selection of ancestry. This piece of reasoning, one of the most remarkable examples of special pleading to be met with in scientific literature, must be read as a whole. I reproduce it entire, that the reader may appreciate this curious effort. The remarks between round parenthetical marks are Professor Weldon’s, those between crotchets are mine.

“Mendel treats such characters as yellowness of cotyledons and the like as if the condition of the character in two given parents determined its condition in all their subsequent offspring. Now it is well known to breeders, and is clearly shown in a number of cases by Galton and Pearson, that the condition of an animal does not as a rule depend upon the condition of any one pair of ancestors alone, but in varying degrees upon the condition of all its ancestors in every past generation, the condition in each of the half-dozen nearest generations having a quite sensible effect. Mendel does not take the effect of differences of ancestry into account, but considers that any yellow-seeded pea, crossed with any green-seeded pea, will behave in a certain definite way, whatever the ancestry of the green and yellow peas may have been. (He does not say this in words, but his attempt to treat his results as generally true of the characters observed is unintelligible unless this hypothesis be assumed.) The experiments afford no evidence which can be held to justify this hypothesis. His observations on cotyledon colour, for example, are based upon 58 cross-fertilised flowers, all of which were borne upon ten plants; and we are not even told whether these ten plants included individuals from more than two races.

Mendel, on the contrary, disregards the “condition of the character” in the parent altogether; but is solely concerned with the nature of the characters of the gametes.

“The many thousands of individuals raised from these ten plants afford an admirable illustration of the effect produced by crossing a few pairs of plants of known ancestry; but while they show this perhaps better than any similar experiment, they do not afford the data necessary for a statement as to the behaviour of yellow-seeded peas in general, whatever their ancestry, when crossed with green-seeded peas of any ancestry. [Mendel of course makes no such statement.]

“When this is remembered, the importance of the exceptions to dominance of yellow cotyledon-colour, or of smooth and rounded shape of seeds, observed by Tschermak, is much increased; because although they form a small percentage of his whole result, they form a very large percentage of the results obtained with peas of certain races. [Certainly.] The fact that Telephone behaved in crossing on the whole like a green-seeded race of exceptional dominance shows that something other than the mere character of the parental generation operated in this case. Thus in eight out of 27 seeds from the yellow Pois d’Auvergne ♀ × Telephone ♂ the cotyledons were yellow with green patches; the reciprocal cross gave two green and one yellow-and-green seed out of the whole ten obtained; and the cross Telephone ♀ × (yellow-seeded) Buchsbaum ♂ gave on one occasion two green and four yellow seeds.

Regarding this “exception” see p. 146.

“So the cross Couturier (orange-yellow) ♀ × the green-seeded Express ♂ gave a number of seeds intermediate in colour. (It is not clear from Tschermak’s paper whether all the seeds were of this colour, but certainly some of them were.) The green Plein le Panier [Fillbasket] ♀ × Couturier ♂ in three crosses always gave either seeds of colour intermediate between green and yellow, or some yellow and some green seeds in the same pod. The cross reciprocal to this was not made; but Express ♀ × Couturier ♂ gave 22 seeds of which four were yellowish green.

See p. 148.

“These facts show first that Mendel’s law of dominance conspicuously fails for crosses between certain races, while it appears to hold for others; and secondly that the intensity of a character in one generation of a race is no trustworthy measure of its dominance in hybrids. The obvious suggestion is that the behaviour of an individual when crossed depends largely upon the characters of its ancestors. When it is remembered that peas are normally self-fertilised, and that more than one named variety may be selected out of the seeds of a single hybrid pod, it is seen to be probable that Mendel worked with a very definite combination of ancestral characters, and had no proper basis for generalisation about yellow and green peas of any ancestry” [which he never made].

Where was that “logician,” the “consulting-partner,” when this piece of reasoning passed the firm?

Let us pause a moment before proceeding to the climax. Let the reader note we have been told of two groups of cases in which dominance of yellow failed or was irregular. (Why are not Gärtner’s and Seton’s “exceptions” referred to here?) In one of these groups Couturier was always one parent, either father or mother, and were it not for Tschermak’s own obvious hesitation in regard to his own exceptions (see p. 148), I would gladly believe that Couturier--a form I do not know--may be an exceptional variety. How Professor Weldon proposes to explain its peculiarities by reference to ancestry he omits to tell us. The Buchsbaum case is already disposed of, for on Tschermak’s showing, it is an unstable form.

Happily, thanks to Professor Weldon, we know rather more of the third case, that of Telephone, which, whether as father or mother, was frequently found by Tschermak to give either green, greenish, or patchwork-seeds when crossed with yellow varieties. It behaves, in short, “like a green-seeded pea of exceptional dominance,” as we are now told. For this dominant quality of Telephone’s greenness we are asked to account by appeal to its ancestry. May we not expect, then, this Telephone to be--if not a pure-bred green pea from time immemorial--at least as pure-bred as other green peas which do not exhibit dominance of green at all? Now, what is Telephone? Do not let us ask too much. Ancestry takes a lot of proving. We would not reject him “parce qu’il n’avait que soixante & onze quartiers, & que le reste de son arbre généalogique avait été perdu par l’injure du tems.”

But with stupefaction we learn from Professor Weldon himself that Telephone is the very variety which he takes as his type of a permanent and incorrigible mongrel, a character it thoroughly deserves.