We have seen in the last paragraph that the Y comb with only 10 per cent to 0 per cent median element has germ-cells bearing median comb as truly as the Y comb containing 70 per cent to 80 per cent median element, but we have also seen that in the latter case there is an excess of single-combed progeny. We have now to inquire whether, in general, there is a close relation between the proportion of median element in the comb of the parents and the percentage of single-combed offspring. These relations are brought out in the lower half of table 9.

Table 9.—Frequency of the different proportions of single element in the combs of offspring of parents having the average proportion of median element given in the column at the left.

Y combs.
Offspring.
0102030405060708090Total.
Parents 021543465108167
102153039242049
205421042120131
30817810922123083127
4097427391846265163
50752153213483511159
601072221914475115169
709240167284111109
80......111161211639
90...21003038926
Total90543120321417924019062939
All types of combs in offspring.
Number ofIYNon-median.
offspring.No.P. ct.No.P. ct.No.P. ct.
Parents0146422067463725
1099252549502525
2073223031432027
302496125127516124
403097324163537323
503299328159487723
6036812033169467921
702328035109474318
80104424039382322
9075385126341115
Total198459630.093947.344922.7

The proportion of single-combed offspring in the total filial population is 30.0 per cent, a departure of such magnitude from the expected 25 per cent as to arrest our attention. Further inspection of table 9 shows that the excess of single-combed offspring is found only in the lower half of the series. When the percentage of median element in the parents is under 50 the proportions of I, Y, and no-median combs are as 25.5 per cent, 49.8 per cent, 24.7 per cent, or close to expectation; but when the percentage is 50 or over the proportions are, on the average, 33.6 per cent, 45.2 per cent, and 21.2 per cent, a wide departure from expectation, 1108 individuals being involved. An examination of table 9 shows, moreover, that the proportion of offspring with single comb rises steadily as the proportion of the median element in the parentage increases from 50 per cent. The meaning of this fact is at present obscure, but the suspicion is awakened that, while the "cup comb" and the more deeply split combs are typical heterozygotes the slightly split combs are a complex of 2 or more units, one of which is "single comb." But that this is not the explanation follows for two reasons: first, that even in the F1 generation slightly split combs are obtained, and, second, that the offspring of the cup combs are much more variable than those of slightly split combs (70 to 90 per cent median). What is strikingly true is that, from 50 per cent up, as the proportion of the median element in the parents increases the percentage of single-combed offspring rises.

The matter may be looked at in another light. Median comb is dominant over its absence. Typically, we should expect F1 to show a single comb; the Y comb that we actually get is a heterozygous condition due to the failure of the median comb to dominate completely. Typically we should expect F2 to reveal 75 per cent single combs, of which 1 in 3 is homozygous and 2 in 3 are heterozygous. Owing to the failure of single comb always to dominate completely in the heterozygotes, we expect to find some of the 75 per cent with the Y comb. When in the parents dominance has been very incomplete in the heterozygote (as is the case in the 0 per cent to 40 per cent median-combed parents) we find it so in the offspring also and all heterozygotes show a Y comb of some type. But when in the parents dominance has been strong in the heterozygote (50 per cent to 90 per cent) it is so in the offspring also and only a part of the heterozygotes show the Y comb; the others show the single comb and thus swell the numbers of the single-combed type. The only objection to this explanation is found in the reduction in the percentages of the no-median type. Thus, adding together the homozygous and heterozygous median-combed offspring and comparing with the non-median-combed, we find these ratios:

Parental per cent 0-405060708090
Ratio75.3 : 24.7 76 : 23 79 : 21 82 : 18 78 : 22 85 : 15

There is a great deviation from 25 per cent in the "non-median" offspring of the 90 per cent parents, but in this particular case the total number of offspring is not large, and the deviation has a greater chance of being accidental. Altogether this explanation of the varying per cents of single comb on the ground of inheritance of varying potency in dominance seems best to fit the facts of the case.

From the foregoing facts and considerations we may conclude that the Y comb represents imperfect dominance of median over no-median comb; that there is a fluctuation in the potency of the dominance, so that the proportion of the median element varies from 0 to over 90 per cent; that the more potent the dominance of median element is in any parents the more complete will be the dominance in the offspring and the smaller will be the percentage of imperfectly dominant, or Y-combed, offspring. Dominance varies quantitatively and the degree of dominance is inheritable.

The index of heredity may be readily obtained in the familiar biometric fashion from table 9. This I have calculated and found to be 0.301 ± 0.002. This agrees with Pearson's theoretical coefficient of correlation between offspring and parent. The index is larger than it would otherwise be because it is measured with an average of the parents and these parents assortatively mated. But this instance is, in any case, an interesting example of strong inheritance of a quantitative variation.

What, it may be asked, is the relation of these facts to the general principle that inheritance is through the gametes? Why, when a gamete with the median element unites with a gamete without that element, does the zygote develop a soma that in some cases shows a nine-tenths median and sometimes a one-tenth median element? We have seen that the Y comb is a heterozygous form due to imperfection of dominance of the median element; but why this variation in the perfection of the median element? This is probably a piece of the question, why any dominance at all. We find, in general, that the determiner of a well-developed organ dominates in the zygote over the determiner of a slightly developed condition of that organ or its obsolete condition. It is as though there were in the zygote an interaction between the strong and the weak form of the determiner, and the strong won; but sometimes the victory is imperfect. In the specific case of comb the interaction between median and no-median leads to a modification, weakening, or imperfection of the median element, and this weakening varies in degree. Sometimes the weakening is inappreciable—when the comb is essentially single; sometimes it is great, and the result is a comb in which the median element is reduced to one-half; sometimes, finally, the determiner of median comb is so completely weakened by its dilution with "no-median" as not to be able to develop, and we have the cup comb with only a trace of the median element. Nevertheless, such a cup comb is heterozygous and produces both single-combed and Polish-combed germ-cells. Thus the variation in the extent of the median comb seems to point to variations in relative potency of the median comb over its absence.