As early as 1902, Correns used as Mendelian pairs, presence of coloring material and absence; also modification into yellow and no modification. In 1905, he extended somewhat this use of present and absent characters, k (keine) preceding the symbol of a character as a negative. Still he did not pretend to generalize the relation of dominance and recessiveness to be that of presence and absence. In 1903 (p. 146) de Vries stated that in very many cases Mendel's law held when one quality is active and the other latent, and that the active quality is dominant. His illustrations show that by activity he meant essentially presence, by latency absence from the visible soma. Bateson's third report (1906) applies presence and absence to several additional cases, and, at the International Genetics Conference of that year, Hurst developed the presence-and-absence hypothesis, favoring the view that the factor for absence is nothing at all, but finding that certain cases, such as Angora coat, offer a difficulty. At the same meeting I suggested that "a variation * * * that is due to abbreviation of the ontogenetic process, which depends on something having dropped out, will be recessive," a progressive variation dominant; and in 1908 I expressed the conclusion that "dominance in heredity appears when a stronger determiner meets a weaker determiner in the germ. The extreme case is that in which a strong determiner meets a determiner so weak as to be practically absent, as when a red flower is crossed with white." I suggested that in some cases of recessiveness of an apparent advanced condition, like Angora hair, the dominant factor is an inhibitor. In the last year or two the presence-and-absence theory has gained wide acceptance, but I still think the cases where there is dominance of the advanced condition over the less advanced—of the quantitatively well-developed over the quantitatively less well-developed—have not been sufficiently considered. In human hair-color any other hypothesis demands that there are many units in the higher grades of pigmentation and fewer in the lower grades and that the presence of the surplus factor in any other higher grade dominates over its absence in the next lower grade; but there is no evidence in human hair-color of distinct, discontinuous units in the common yellow-brown series. And, in ontogeny, the different grades of color form a continuous series whose development proceeds throughout early life and may even be stimulated to an advanced stage of darkening by disease. The cessation of color development may take place at any point, and this seems incompatible with the theory of unit-characters for the different grades of human hair-color. In the present paper, on the other hand, the characters dealt with are mostly unit-characters and their quantitative variations mostly heterozygotic. Even the case of the Silkie boot ([table 31], C) referred to in an earlier paper[14] as illustrating recessiveness of the less advanced condition proves, on further analysis, to be a case of heterozygotism. It seems highly probable that the future will show that many more advanced or progressive conditions are really due to one or more unit-characters not present in the less advanced condition. In that case it will appear that there is perfect accord in the two statements that the progressive condition and the "present" factor are dominant.
The definition of dominance on the ground of results meets at the outset with a difficulty the germ of which is observable in Mendel's cautious statement "ganz oder fast unverändert." Even Mendel observed that the hybrids between white-flowered and purple-red flowered peas have flowers less intensely colored than the darker parent. The experiments of the last seven years have shown that the "dominant" character is often very greatly changed—indeed, in extreme cases a blending of characters may occur—in the first generation. Correns (1900 b, p. 110) very early stated that in a certain set of crosses between good species the hybrids showed the character of both parents, only reduced, but in varying degrees. Bateson and Saunders (1902, p. 23) found in crossing two forms of Datura that—
Although the offspring resulting from a cross between any two of the forms employed are usually indistinguishable from the type which is dominant as regards the particular character crossed, yet in other cases the intensity of a dominant character may be more or less diminished either in particular individuals or in particular parts of one individual. In Tatula-Stramonium cross-breds the corolla is often paler in color than that of the dominant parent (as has already been noticed by Naudin), but even in the palest specimens the deep blue color of the unopened anthers leaves no doubt as to the presence of the dominant color element. * * * The occurrence of intermediate forms was also occasionally noticeable in the fruits. Among the large number of capsules examined, there were some of the mosaic type, in which part of the capsule was prickly and the remainder smooth, while others, suggesting a blend, were more or less prickly all over, but the prickles were much reduced in size, and often formed mere tubercles.
Bateson and Saunders further showed (1902, p. 123) that in the case of comb and extra-toe in poultry "the cross-bred may show some blending and * * * the intensity of the dominant character is often considerably reduced."
Correns (1905, p. 9) pointed out that there was known, even at that time, a complete series of cases at one extreme of which one determiner completely hindered the appearance of the other, while at the opposite end of the series the hybrid showed an intermediate condition, both determiners appearing with equal strength.
The following year, in my first report on Inheritance in Poultry, I laid great stress on the imperfection of dominance, and this phenomenon has become more striking and clear in the subsequent years, until in the present paper it is recognized as the key to the explanation of many apparently anomalous types of heredity.
The first case in the present work in which imperfection of dominance is considered is that of the hybrids between I and oo comb. Here median comb is mated with no-median. Each somatic cell of the hybrid—at least in the comb region—has only half the full determiner for median comb. The determiner is weakened, and so the median comb is imperfectly developed, namely, at the anterior end of its proper territory. The weakening varies much in degree in the heterozygote. The median comb may be reduced to 70 per cent of its normal length or it may not develop at all.
The second case of imperfection of dominance is that of polydactylism. Extra-toe mated to normal gives extra-toe in 73 per cent only of the offspring in the case of the Houdans. Any trace of 6 toes (on one or both feet) is found in only 12 per cent of the hybrid offspring from a 6-toed Silkie parent. Certainly dominance here is very like blending.
The third case of imperfection of dominance is that of syndactylism. No syndactyls were noticed in F1. My first conclusion was that syndactylism is recessive; but later studies have shown that it is dominant and that all matings of two syndactyl parents yield about 56 per cent syndactyl offspring.
Rumplessness gives an illustration of how dominance may be so weak as to be absent altogether; so that from F1 alone the erroneous conclusion is drawn that it is recessive; indeed, in one strain, only faint traces of the character made their appearance in successive generations.