4. THE NATURE OF MENDEL'S DISCOVERY. The reconstruction has been undertaken in order to test the
plausibility of the view that Mendel's statements as to the course and procedure of his experimentation are to be taken as an entirely literal account, or whether, on the other hand, there is evidence that data have been a**embled from various sources, or the same data rediscussed from different standpoints in different sections of his account. There can, I believe, now be no doubt whatever that his report is to be taken entirely literally, and that his experiments were carried out in just the way and much in the order that they are recounted. The detailed reconstruction of his programme on this a**umption leads to no discrepancy whatever. A serious and almost inexplicable discrepancy has, however, appeared, in that in one series of results the numbers observed agree excellently with the two to one ratio, which Mendel himself expected, but differ significantly from what should have been expected had his theory been corrected to allow for the small size of his test progenies. To suppose that Mendel recognized this theoretical complication, and adjusted the frequencies supposedly observed to allow for it, would be to contravene the weight of the evidence supplied in detail by his paper as a whole. Although no explanation can be expected to be satisfactory, it remains a possibility among others that Mendel was deceived by some a**istant who knew too well what was expected. This possibility is supported by independent evidence that the data of most, if not all, of the experiments have been falsified so as to agree closely with Mendel's expectations.
The importance of the conclusion, if it is well established, that Mendel's statements are to be taken literally, lies in the inferences which flow from this view. First, that prior to the reported experiments Mendel was sufficiently aware of the independent inheritance of seven factors in peas to have chosen seven pairs of varieties, each pair differing only in a single factor. If it be thought that out of thirty-eight varieties he could not by deliberate choice have found the material for seven such crosses, it should be remembered also that at this stage he was choosing not only the varieties but, perhaps, also the factors to use in his experiment, and that he may have known of other factors in peas in addition to those with which his experiments are concerned, which, however, could not have been introduced without bringing in an undesirable complication.
Next, it appears that Mendel regarded the numerical frequency ratios, in which the laws of inheritance expressed themselves, simply as a ready method of demonstrating the truth of his factorial system, and that he was never much concerned to demonstrate either their exactitude or their consistency. It may be that the seed counts of 1859 were a revelation to him of the precision with whic h his system worked, and could be demonstrated; they may also possibly have given him an exaggerated impression of the precision with which the theoretical ratios should be verified, but from that moment it is clear, from the form his experiments took, that he knew very surely what to expect, and designed them as a demonstration for others rather than for his own enlightenment. That the hereditary contribution of the two parents might be unequal he did not seriously consider, although his first experiments provided splendid evidence on this important question, which it does not occur to him to present. It seems also not to have occurred to him that the inheritance of different factors might not be wholly independent. He a**erts independence for all his factors, but gives evidence for only three of them, and for these much less than he might have given. A feature such as linkage would have been a complication extraneous to his theory, as he conceived it, which he would only have taken seriously had the observations forced it under his notice.
The theoretical consequences of his system he had thought out thoroughly, and in this respect his thought is considerably in advance of that of the first generation of geneticists which followed his rediscovery. He pointe d out that n factors would give rise to 3n different genotypes, of which 2n would be capable of breeding true. He realized that even in intra-specific crosses n would be sufficiently great for these to be very large numbers, and that even more factors must be involved when crosses are made between different species, when minor in addition to major differences are considered. This understanding of the consequences of the factorial system contrasts sharply with many of the speculations of the earlier geneticists, such as that new species might be formed by the mutation of a single factor, or that the mimetic groups, found among bu*terflies and other insects, might be explained by the paucity of the genetic factors controlling the pattern and coloration of the wings. In these respects it has taken nearly a generation to rediscover Mendel's point of view.
Mendel seems also to have realized that the factorial system resolved one of the chief difficulties felt and discussed by Darwin, namely that, if the wide variation observable in cultivated plants were caused by the changed conditions and increased nourishment experienced on being brought into cultivation, then this cause of variation must continue to act, as Darwin had written, "for an improbably long time", since anciently cultivated species are not less but rather more variable than others. With segregating, heritable factors, on the other hand, the variability is easily explained by the preservation in culture of variants which, apart from man, would have been eliminated by natural selection. This, indeed, seems to have been Mendel's view (p. 351) : "It will be willingly granted that by cultivation the originationof new varieties is favoured, and that by men's labour many varieties are acquired which, under natural conditions, would be lost; but nothing justifies the a**umption that the tendency to the formation of varieties is so extraordinarily increased that the species speedily lose all stability, and their offspring diverge into an endless series of extremely variable forms. Were the change in the conditions the sole cause of variability you might expect that those cultivated plants which are grown forcenturies under almost identical conditions would again attain constancy. That, as is well known, is not the case, ..." The reflection of Darwin's thought is unmistakable, and Mendel's comment is extremely pertinent, though it seems to have been overlooked. He may at this time have read the Origin, but the point under discussion may equally have reached his notice at second hand.