Since heredity lies at the basis of the breeder's work, it is evident that any contribution to a more exact knowledge of this subject must prove of service to him, and there is no doubt that he will be able to profit by Mendelian knowledge in the conduct of his operations. Indeed, as we shall see later, these ideas have already led to striking results in the raising of new and more profitable varieties. In the first place, heredity is a question of individuals. Identity of appearance is no sure guide to reproductive qualities. Two individuals similarly bred and indistinguishable in outward form may nevertheless behave entirely differently when bred from. Take, for instance, the family of sweet peas shown on Plate IV. The F2 generation here consists of seven distinct types, three sorts of purples, three sorts of reds, and whites. Let us suppose that our object is to obtain a true breeding strain of the pale purple picotee form. Now from the proportions in which they come we know that the dilute colour is due to the absence of the factor which intensifies the colour. Consequently the picotee cannot throw the
two deeper shades of red or purple. But it may be heterozygous for the purpling factor when it will throw the dilute red (tinged white), or it may be heterozygous for either or both of the two colour factors (cf. p. [44]), in which case it will throw whites. Of the picotees which come in such a family, therefore, some will give picotees, tinged whites, and whites, others will give picotees and tinged whites only, others will give picotees and whites only, while others, again, and these the least numerous, will give nothing but picotees. The new variety is already fixed in a certain definite proportion of the plants; in this particular instance in 1 out of every 27. All that remains to be done is to pick out these plants. Since all the picotees look alike, whatever their breeding capacity, the only way to do this is to save the seed from a number of such plants individually, and to raise a further generation. Some of them will be found to breed true. The variety is then established, and may at once be put on the market with full confidence that it will hereafter throw none of the other forms. The all-important thing is to save and sow the seed of separate individuals separately. However alike they look, the seed from different individuals must on no account be mixed. Provided that due care is taken in this respect no long and tedious process of selection is required for the fixation of any given variety. Every possible variety arising from a cross appears in the F2 generation if only a sufficient
number is raised, and of all these different varieties a certain proportion of each is already fixed. Heredity is a question of individuals, and the recognition of this will save the breeder much labour, and enable him to fix his varieties in the shortest possible time.
Such cases as these of the sweet pea throw a fresh light upon another of the breeder's conceptions, that of purity of type. Hitherto the criterion of a "pure-bred" thing, whether plant or animal, has been its pedigree, and the individual was regarded as more or less pure bred for a given quality according as it could show a longer or shorter list of ancestors possessing this quality. To-day we realise that this is not essential. The pure-bred picotee appears in our F2 family though its parent was a purple bicolor, and its remoter ancestors whites for generations. So also from the cross between pure strains of black and albino rabbits we may obtain in the F2 generation animals of the wild agouti colour which breed as true to type as the pure wild rabbit of irreproachable pedigree. The true test of the pure breeding thing lies not in its ancestry but in the nature of the gametes which have gone to its making. Whenever two similarly constituted gametes unite, whatever the nature of the parents from which they arose, the resulting individual is homozygous in all respects and must consequently breed true. In deciding questions of purity it is to the gamete, and not to ancestry, that our appeal must henceforth be made.
Improvement is after all the keynote to the breeder's operations. He is aiming at the production of a strain which shall combine the greatest number of desirable properties with the least number of undesirable ones. This good quality he must take from one strain, that from another, and that again from a third, while at the same time avoiding all the poor qualities that these different strains possess. It is evident that the Mendelian conception of characters based upon definite factors which are transmitted on a definite scheme must prove of the greatest service to him. For once these factors have been determined, their distribution is brought under control, and they can be associated together or dissociated at the breeder's will. The chief labour involved is that necessary for the determination of the factors upon which the various characters depend. For it often happens that what appears to be a simple character turns out when analysed to depend upon the simultaneous presence of several distinct factors. Thus the Malay fowl breeds true to the walnut comb, as does also the Leghorn to the single comb, and when pure strains are crossed all the offspring have walnut combs. At first sight it would be not unnatural to regard the difference as dependent upon the presence or absence of a single factor. Yet, as we have already seen, two other types of comb, the pea and the rose, make their appearance in the F2 generation. Analysis shows that the difference between the walnut
and the single is a difference of two factors, and it is not until this has been determined that we can proceed with certainty to transfer the walnut character to a single-combed breed. Moreover, in his process of analysis the breeder must be prepared to encounter the various phenomena that we have described under the headings of interaction of factors, coupling and repulsion, and the recognition of these phenomena will naturally influence his procedure. Or again, his experiments may show him that one of the characters he wants, like the blue of the Andalusian fowl, is dependent upon the heterozygous nature of the individual which exhibits it, and if such is the case he will be wise to refrain from any futile attempt at fixing it. If it is essential it must be built up again in each generation, and he will recognise that the most economical way of doing this is to cross the two pure strains so that all the offspring may possess the desired character. The labour of analysis is often an intricate and tedious business. But once done it is done once for all. As soon as the various factors are determined, upon which the various characters of the individual depend, as soon as the material to be made use of has been properly analysed, the production and fixation of the required combinations becomes a matter of simple detail.
An excellent example of the practical application of Mendelian principles is afforded by the experiments which Professor Biffen has recently carried out in Cambridge.
Taken as a whole English wheats compare favourably with foreign ones in respect of their cropping power. On the other hand, they have two serious defects. They are liable to suffer from the attacks of the fungus which causes rust, and they do not bake into a good loaf. This last property depends upon the amount of gluten present, and it is the greater proportion of this which gives to the "hard" foreign wheat its quality of causing the loaf to rise well when baked. For some time it was held that "hard" wheat with a high glutinous content could not be grown in the English climate, and undoubtedly most of the hard varieties imported for trial deteriorated greatly in a very short time. Professor Biffen managed to obtain a hard wheat which kept its qualities when grown in England. But in spite of the superior quality of its grain from the baker's point of view its cropping capacity was too low for it to be grown profitably in competition with English wheats. Like the latter, it was also subject to rust. Among the many varieties which Professor Biffen collected and grew for observation he managed to find one which was completely immune to the attacks of the rust fungus, though in other respects it had no desirable quality to recommend it. Now as the result of an elaborate series of investigations he was able to show that the qualities of heavy cropping capacity, "hardness" of grain, and immunity to rust can all be expressed in terms of Mendelian factors. Having once analysed his material
the rest was comparatively simple, and in a few years he has been able to build up a strain of wheat which combines the cropping capacity of the best English varieties with the hardness of the foreign kinds, and at the same time is completely immune to rust. This wheat has already been shown to keep its qualities unchanged for several years, and there is little doubt that when it comes to be grown in quantity it will exert an appreciable influence on wheat-growing in Great Britain.