A FAMILY WITH ORTHODACTYLY
Fig. 18.—Squares denote males and circles females, as is usual in the charts compiled by eugenists; black circles or squares denote affected individuals. A1 had all fingers affected in the way shown in Fig. 17; B2 had all but one finger affected; C2 had all but one finger affected; D2 had all fingers affected; D3 has all but forefingers affected. The family here shown is a branch, found by F. N. Duncan, of a very large family first described by Harvey Cushing, in which this abnormality has run for at least seven generations. It is an excellent example of an inherited defect due to a single Mendelian factor.
One of the best attested single characters in human heredity is brachydactyly, "short-fingerness," which results in a reduction in the length of the fingers by the dropping out of one joint. If one lumps together all the cases where any effect of this sort is found, it is evident that normals never transmit it to their posterity, that affected persons always do, and that in a mating between a normal and an affected person, all the offspring will show the abnormality. It is a good example of a unit character.
But its effect is by no means confined to the fingers. It tends to affect the entire skeleton, and in a family where one child is markedly brachydactylous, that child is generally shorter than the others. The factor for brachydactyly evidently produces its primary effect on the bones of the hand, but it also produces a secondary effect on all the bones of the body.
Moreover, it will be found, if a number of brachydactylous persons are examined, that no two of them are affected to exactly the same degree. In some cases only one finger will be abnormal; in other cases there will be a slight effect in all the fingers; in other cases all the fingers will be highly affected. Why is there such variation in the results produced by a unit character? Because, presumably, in each individual there is a different set of modifying factors or else a variation in the factor. It has been found that an abnormality quite like brachydactyly is produced by abnormality in the pituitary gland. It is then fair to suppose that the factor which produces brachydactyly does so by affecting the pituitary gland in some way. But there must be many other factors which also affect the pituitary and in some cases probably favor its development, rather than hindering it. Then if the factor for brachydactyly is depressing the pituitary, but if some other factors are at the same time stimulating that gland, the effect shown in the subject's fingers will be much less marked than if a group of modifying factors were present which acted in the same direction as the brachydactyly factor,—to perturb the action of the pituitary gland.
This illustration is largely hypothetical; but there is no room for doubt that every factor produces more than a single effect. A white blaze in the hair, for example, is a well-proved unit factor in man; the factor not only produces a white streak in the hair, but affects the pigmentation of the skin as well, usually resulting in one or more white spots on some part of the body. It is really a factor for "piebaldism."
For the sake of clear thinking, then, the idea of a unit character due to some unit determiner or factor in the germ-plasm must be given up, and it must be recognized that every visible character of an individual is the result of numerous factors, or differences in the germ-plasm. Ordinarily one of these produces a more notable contribution to the end-product than do the others; but there are cases where this statement does not appear to hold good. This leads to the conception of multiple factors.
In crossing a wheat with brown chaff and one with white chaff, H. Nilsson-Ehle (1909) expected in the second hybrid generation to secure a ratio of 3 brown to 1 white. As a fact, he got 1410 brown and 94 white, a ratio of 15:1. He interpreted this as meaning that the brown color in this particular variety was due not to one factor, but to two, which were equivalent to each other, and either one of which would produce the same result alone as would the two acting together. In further crossing red wheat with white, he secured ratios which led him to believe that the red was produced by three independent factors, any one of which would produce red either alone or with the other two. A. and G. Howard later corroborated this work,[48] but showed that the three factors were not identical: they are qualitatively slightly different, although so closely similar that the three reds look alike at first sight. E. M. East has obtained evidence from maize and G. H. Shull from shepherd's-purse, which bears out the multiple factor hypothesis.
