Are Factors Changed Through Selection?
As we have seen, the variation that we find from individual to individual is due in part to the environment; this can generally be demonstrated. Other differences in an
ordinary population are recognized as due to different genetic (hereditary) combinations. No one will dispute this statement. But is all the variability accounted for in these two ways? May not a factor itself fluctuate? Is it not a priori probable that factors do fluctuate? Why, in a word, should we regard factors as inviolate when we see that everything else in organisms is more or less in amount? I do not know of any a priori reason why a factor may not fluctuate, unless it is, as I like to think, a chemical molecule. We are, however, dealing here not with generalities but with evidence, and there are three known methods by means of which it has been shown that variability, other than environmental or recombinational, is not due to variability in a factor, nor to various "potencies" possessed by the same factors.
(1) By making the stock uniform for all of its factors—chief factors and modifiers alike. Any change in such a stock produced by selection would then be due to a change in one or more of the factors themselves. Johannsen's experiment is an example of this sort.
Fig. 83 a. Drosophila ampelophila with truncate wings.
(2) The second method is one that is
capable of demonstrating that the effects of selection are actually due to modifiers. It has been worked out in our laboratory, chiefly by Muller, and used in a particular case to demonstrate that selection produced its effect by isolating modifying factors. For example, a mutant type called truncate appeared, characterized by shorter wings, usually square at the end, (fig. 83a). The wings varied from those of normal length to wings much shorter (fig. 83b). For three years the mutant stock was
bred from individuals having the shorter wings until at last a stock was obtained in which some of the individuals had wings much shorter than the body. By means of linkage experiments it was shown that at least three factors were present that modified the wings. These were isolated by means of their linkage relations, and their mutual influence on the production of truncate wings was shown.
