CHAPTER VI. BOOTING .

Previous

The method of inheritance of the feathering on the feet of some poultry has already been made the subject of much study. Hurst (1905, p. 152) crossed booted and non-booted birds and bred the hybrids together. He concluded that "the Mendelian principles are at work in these aberrant phenomena, but are masked by something not yet perceived." My own conclusion (1906, p. 72) was: "Booting is dominant, but usually imperfectly so." A more extended study has been desirable.

Booting is variable in amount. To indicate its degree I have had recourse to an artificial scale. I recognize 11 grades, running from 0 to 10. The grade 0 implies no feathers whatsoever. Grade 10 implies heavy booting extending over the front half of the shank. Grade 5 implies an extent of only half of the maximum, i. e., the outer front quarter of the shank. Intermediate grades indicate intermediate extension of the feathered area.

A. TYPES OF BOOTING.

The races of booted poultry used have been as follows: First, bantam Cochins of two varieties; second, a bantam Dark Brahma; and third, the Silkie. In my representatives of the first two groups, but particularly in the Dark Brahma, the amount of booting is variable. In one type the outer third of the shank in the newly hatched chick is covered by strong, heavy, specialized feathers, directed outward, while the middle and inner thirds are covered by smaller, finer, imbricating feathers sparsely placed and resembling reduced contour-feathers. In most individuals the transition from the one kind to the other is gradual, while in others it is sharp, and in a few the outer third only of the shank is feathered. In the Silkies, which the standard poultry books describe as being more sparsely feathered on the shank,[8] the outer zone of feathers is the only one developed; and, occasionally, as table 31 shows, even these feathers may be lacking. We have thus two types to distinguish—the extended (Cochin, Brahma) type and the restricted type.

B. NORMAL VARIABILITY.

To appreciate the results of hybridizing we must first examine the variability of pure-blooded races. This is done in table 31.

Table 31.Distribution of boot-grades in the offspring of Cochin, Dark Brahma, and Silkie parents.

[A] Determination made on embryo chicks.
A. OFFSPRING OF COCHIN PARENTS.
Pen No. Mother. Father. Grades of boot in offspring.
No. Boot-
grade.
No. Boot-
grade.
0 1 2 3 4 5 6 7 8 9 10 Average.
848 2297 10 545 10 ... ... ... 1 ... ... 1 ... ... 1 18 9.43
776 2574 10 2732 8 ... ... ... ... ... ... ... ... 3 2 6 9.27
848 2300 8 545 10 ... ... ... ... ... ... ... ... 1 2 5 9.25
776 2570 6 2732 8 ... ... ... ... ... ... 1 1 ... 11 1 8.71
848 2075 9 545 10 ... ... ... ... ... 1 1 ... ... ... 4 8.50
776 2072 6 2732 8 ... ... ... ... ... ... 1 ... 4 2 2 8.44
758 130 6 545 10 ... ... ... ... ... 1 1 1 3 9 ... 8.20
776 2073 6 2732 8 ... ... ... .... 1 2 ... 2 2 10 1 8.00
776 2300 6 2732 8 ... ... ... ... 1 ... 1 3 6 5 2 8.00
758 131 10 545 10 ... ... ... ... ... 1 ... 4 6 1 1 7.96
776 2297 6 2732 8 ... ... ... 1 ... 1 ... 3 6 6 2 7.95
776 1132 3 2732 8 ... ... ... 1 1 1 1 3 6 8 ... 7.57
776 2937 7 2732 8 ... ... ... ... ... ... 1 3 3 1 ... 7.50
776 2299 7 2732 8 ... ... 1 ... ... 1 1 4 7 3 1 7.44
Totals (199) ... ... 1 3 3 8 9 24 47 61 43 8.24
B. OFFSPRING OF DARK BRAHMA PARENTS.
[All individuals have sprung from No. 121 ? (boot of grade 9) and No. 122 ? (boot of grade 6).]
816 2030 6 122 6 ... ... ... ... ... ... ... ... ... 1 3 9.8
703 2030 6 122 6 ... ... ... ... ... ... 4 2 0 3 6 8.3
816 121 6 122 6 ... ... ... ... ... 1 3 1 2 4 5 8.3
816 5979 6 122 6 ... ... ... ... ... ... 1 0 2 ... ... 7.3
816 2353 5 122 6 ... ... ... ... [A]1 1 1 0 1 0 2 7.1
816 5835 5 122 6 ... ... ... [A]1 0 1 2 ... ... 1 3 6.5
816 5840 5 122 6 ... ... ... [A]1 ... ... 1 ... ... ... 1 6.3
703 2353 5 122 6 ... ... ... ... 1 1 3 ... 1 ... ... 5.8
Totals (61) ... ... ... 2 2 4 15 3 6 9 20 7.62
C. OFFSPRING OF SILKIE PARENTS.
734 468 4 774 3 ... ... 1 2 ... ... 1 1 ... ... ... 4.20
734 1002 3 774 3 ... ... 1 4 ... 1 3 ... ... ... ... 4.11
734 841 (?) 774 3 ... ... ... ... 2 ... ... ... ... ... ... 4.00
815 7434 7 774 3 ... ... ... ... 2 ... ... ... ... ... ... 4.00
734 773 1 774 3 ... ... ... 2 2 ... ... ... ... ... ... 3.50
734 680 1 774 3 ... ... ... 2 ... ... ... ... ... ... ... 3.00
734 405a 1 774 1 3 ... ... 1 3 1 ... ... ... ... ... 3.00
815 499 2 774 3 1 1 3 ... ... 2 ... 1 ... ... ... 3.00
734 499 2 774 3 1 1 5 2 2 1 ... ... ... ... ... 2.50
734 500 1 774 3 2 1 2 3 ... ... ... ... ... ... ... 1.75
815 773 1 774 3 4 1 3 ... ... ... ... ... ... ... ... 1.25
815 500 1 774 3 1 1 ... ... ... ... ... ... ... ... ... 0.50
815 496 3 774 3 1 ... ... ... ... ... ... ... ... ... ... 0.00
Totals(68) 10 5 16 18 9 4 4 2 ... ... ... 2.72
SUMMARY.
Races. Grades of boot in offspring, reduced to percentages.
0 1 2 3 4 5 6 7 8 9 10 Average.
Cochins. ... ... 0.5 1.5 1.5 4.0 4.5 12.1 23.6 30.7 21.6 8.24
Dark Brahmas. ... ... ... 3.3 3.3 6.6 24.6 4.9 9.8 14.8 32.8 7.62
Silkie. 14.8 7.4 23.5 26.5 13.2 5.9 5.9 2.9 ... ... ... 2.72

An inspection of table 31 shows that, in respect to booting, the Cochins and Dark Brahmas are clearly closely related to each other. Owing to smaller numbers and to other circumstances that will be discussed later, the results are less regular in the Dark Brahma offspring, but in both the range is from 2 or 3 upward to 10, with a great preponderance in grades above 5. In the Silkies, on the other hand, the greatest frequency is found in grades below 5. This difference is correlated with a difference of the parents, for the commonest grades of the parents of the Cochins are between 6 and 10, of the Dark Brahmas between 5 and 9, and of the Silkies between 1 and 3. These results suggest that the Silkie is typically heterozygous in boot, producing 25 per cent recessives (boot of grade 4-7) and 75 per cent dominant (0, 1) and heterozygous (2, 3). We shall see that this hypothesis receives support from all Silkie matings.

Inside of any part of this table it appears that, on the whole, as the average grade of the boot in the progeny diminishes that of the parentage diminishes, although the correlation is by no means perfect. Thus the average of the parental grades in the first part of table 31, A (which is arranged in descending order of the averages of the offspring) is 8.5; in the lower half, 7.4. The average of parental grades in the upper half of table 31, B is 6.4; in the lower half 5.5. In table 31, C the grades are 2.9 and 2.3, respectively. This correlation indicates, without exactly measuring, heredity in grade of booting.

Table 32 shows the results of crosses between Cochins (high grade of boot) and Silkies (low grade).

Table 32.Distribution of boot-grades between a high and low grade of boot in parents.

HIGH AND LOW GRADE OF BOOT IN PARENTS.
Pen No. Mother. Father. Grade of boot in offspring.
No. Gen. Races. Gra. No. Gen. Race. Gra. 0 1 2 3 4 5 6 7 8 9 10 Aver-
age.
851 5567 P Bl. × Bf. C. 9 P 7526 Silkie. 3 ... ... ... ... 2 ... ... ... 3 3 5 8.15
851 3410 P Do. 9 P 7526 Do. 3 ... ... ... ... ... 4 3 2 1 6 1 7.29
851 6956 P Do. 8 7526 P Do. 3 ... ... ... ... 3 3 ... 2 2 ... 5 7.13
851 2073 P Do. 7 7526 P Do. 3 ... 1 ... 1 1 ... 1 1 1 3 2 6.91
851 2299 P Do. 7 7526 P Do. 3 ... ... ... ... 2 2 1 1 ... ... 3 6.78
851 840 P Bf. C. 10 7526 P Do. 3 ... ... ... ... 1 ... 1 ... ... 1 ... 6.33
851 1002 P Do. 8 7526 P Do. 3 ... ... ... 3 1 2 1 2 4 1 1 6.27
815 131 P Bk. C. 10 774 P Do. 4 ... ... ... 3 1 1 2 2 1 1 2 6.23
851 841 P Bf. C. 10 7526 P Do. 3 ... ... ... ... 1 ... 1 ... 1 ... ... 6.00
851 838 P Do. 8 7526 P Do. 3 ... ... ... 4 2 4 3 ... ... 2 2 5.65
Totals (116) 0 1 0 11 14 16 13 10 13 17 21 6.77

So far as the average grade of boot in offspring goes, this table stands between that of the Cochins (table 31, A) and that of the Silkies (table 31, C). But what is especially striking is the apparent dimorphism revealed in the line of totals. There is one (empirical) mode at 10, corresponding with that of the Cochins, and a second clear mode at 5, corresponding to that of the Silkies. If we assume the Cochin to be homozygous in boot (RR) and the Silkie to be heterozygous in boot, then we can interpret the high mode as extracted recessives, the median mode as heterozygotes.

C. RESULTS OF HYBRIDIZATION.

We have next to consider the nature of the inheritance when one parent belongs to an unbooted race, the other to a booted one (table 33).

Table 33.Distribution of boot-grades in the F1 generation of booted × non-booted parents.

A. COCHIN CROSSES.
Pen No. Mother. Father. Grade of boot in offspring.
No. Gen. Races. Gra. No. Gen. Races. Gra. 0 1 2 3 4 5 6 7 8 9 10 Aver-
age.
773 1334 P W. Legh. 0 836 P Bl. Coch. 10 ... ... ... 3 1 1 1 1 ... 2 ... 5.44
773 193 P Do. 0 836 P Do. 10 ... 1 2 6 8 7 4 2 ... ... ... 4.27
773 1366 P Do. 0 836 P Do. 10 ... ... ... 2 5 2 1 ... ... ... ... 4.20
773 127 P Do. 0 836 P Do. 10 ... ... 3 10 9 12 4 ... ... ... ... 4.11
773 692 P W. Legh. (R) 0 836 P Do. 10 ... ... ... 10 3 2 ... ... ... ... ... 3.47
774 2075 P Coch. 8 1431 P W. Legh. (R) 0 6 1 1 ... 1 ... ... ... ... ... ... 0.78
Totals (111) 6 2 6 31 27 24 10 3 0 2 0 3.91
B. DARK BRAHMA CROSSES.
727 Y P D. Br. 10 381 P Houd. 0 ... ... ... ... 2 3 2 1 2 ... ... 5.80
727 121 P Do. 10 381 P Do. 0 1 ... ... 1 1 5 4 ... ... ... ... 4.67
823 2030 P Do. 7 3858 P M × P 0 ... ... 5 16 15 4 1 2 ... ... ... 3.67
823 Y P Do. 8 3858 P Do. 0 ... ... 1 7 6 2 ... ... ... ... ... 3.56
838 3814 P W. Legh. 0 122 P D. Br. 6 ... 2 2 6 6 1 1 ... ... ... ... 3.28
838 202 P Min. 0 122 P Do. 6 ... ... 2 5 3 ... ... ... ... ... ... 3.10
838 71 P W. Legh. 0 122 P Do. 6 ... ... ... 1 ... ... ... ... ... ... ... 3.00
838 3832 P Do. 0 122 P Do. 6 1 1 ... 1 1 2... ... ... ... ... ... 3.00
838 10 P Do. 0 122 P Do. 6 ... 1 ... 3 1 ... ... ... ... ... ... 2.80
816 121 P D. Br. 9 4912 P M × P 0 ... ... 8 4 1 1 ... ... ... ... ... 2.64
816 5838 P Do. 9 4912 P Do. 0 ... ... 5 5 1 ... ... ... ... ... ... 2.64
838 5418 P W. L., Min. 0 122 P D. Br. 6 1 1 3 3 1 1 ... ... ... ... ... 2.50
816 5979 P D. Br. 6 4912 P M × P 0 4 3 4 7 4 1 1 ... ... ... ... 2.46
816 2353 P Do. 5 4912 P Do. 0 ... 2 2 4 1 ... ... ... ... ... ... 2.44
816 5977 P Do. 4 4912 P Do. 0 ... 3 2 1 ... 1 ... ... ... ... ... 2.14
816 5835 P Do. 5 4912 P Do. 0 3 5 5 8 3 ... ... ... ... ... ... 2.12
816 5840 P Do. 5 4912 P Do. 0 5 1 3 4 1 ... ... ... ... ... ... 1.64
823 6626 P Do. 2 3858 P Do. 0 1 10 2 2 ... ... ... ... ... ... ... 1.33
816 5980 P Do. 5 4912 P Do. 0 5 8 1 5 ... ... ... ... ... ... ... 1.32
Totals (268) 21 37 45 83 47 21 9 3 2 0 0 2.84
C. SILKIE CROSSES.
774 777 P Silkie. 8 1176 P W. Legh. 0 3 ... 1 1 1 ... ... ... ... ... ... 1.50
744 681 P Do. 5 1176 P Do. 0 11 2 1 1 1 1 ... ... ... ... ... 0.94
744 469 P Do. 1 1176 P Do. 0 11 3 ... ... ... ... ... ... ... ... ... 0.21
Totals (37) 25 5 2 2 2 1 0 0 0 0 0 0.76
SUMMARY.
Crosses. Grades of boot in offspring, reduced to percentages.
0 1 2 3 4 5 6 7 8 9 10 Aver-
age.
Cochin. 5.4 1.8 5.4 28.0 24.3 21.6 9.0 2.7 0.0 1.8 ... 3.91
Brahma. 7.8 13.8 16.8 31.0 17.5 7.8 3.4 1.1 0.7 ... ... 2.84
Silkie. 67.6 13.5 5.4 5.4 5.4 2.7 ... ... ... ... ... 0.76

An inspection of Table 33, which gives the distribution of grades of boot in the offspring constituting the first hybrid generation, might well lead to the conclusion that inheritance is here of a blending nature, or that, if either condition is dominant, it is the booted one, as suggested in my report of 1906. On this hypothesis the offspring with no boot illustrate imperfection of dominance, and one would say that, in booting, dominance is very imperfect.

However plausible such an interpretation might appear when based on the first hybrid generation alone, it becomes untenable when subsequent generations are taken into account, as we shall see later. The hypothesis breaks down completely in the second hybrid generation and we are forced to the opposite hypothesis, namely, that the clean-shanked condition is dominant. Such an hypothesis would seem, at first, to contravene the principle enunciated in my report of 1906 that the more progressive condition is dominant over the less progressive condition, or absence. But such is not necessarily the fact. We have no right to assume that presence of boot is the new character. The rest of the body of poultry (save the head) is covered with feathers. If the foot is not it must be because there is something in the skin of the foot that inhibits the development of feathers there. And this inhibiting factor is dominant over its absence.

Table 33 shows that the Silkie crosses yield an exceptionally high per cent of the dominant clear-footed condition. This is additional evidence that the Silkies are DR, and so this cross produces 50 per cent of pure extracted dominants in addition to 50 per cent of heterozygotes in booting.

To get further light on the nature of inheritance of booting we pass to the examination of the second hybrid generation (table 34).

In the case of Silkies, which throw 67.6 per cent clean-shanked progeny in F1, we find in F2 only about 60 per cent clean-shanked. This diminution is, of course, due to the extraction of some pure booted recessives, which draw from the proportion of clean shanks.

In the case of the Cochins and Dark Brahmas, expectation, with perfect dominance, is that 75 per cent of the offspring shall be clean-shanked. Since dominance is imperfect (as shown by the occurrence of many booted birds in F1) we should look for an actual failure to reach so large a proportion, but we are hardly prepared for the result that in most of the F2 crosses of Cochins and Brahmas less than 25 per cent of the offspring are clean-shanked. In 4 pens the average is only 10 to 12 per cent, and in one only 2 per cent of the offspring fail to develop feathers on the feet. What shall we say of such a case as the last? The history of the father (No. 666) is absolutely certain; his mother was No. 121, the original Dark Brahma female, with a boot of grade 9 and a record in her immediate progeny that indicates perfect purity of booting in her germ-cells. His father was a White Leghorn with clean shanks and without a suspicion of having such antipodal blood as the Asiatic in his ancestry. No. 666 is certainly heterozygous in boot, if boot is a single unit. The hens with which No. 666 were mated were clearly heterozygous, as is known not only from their ancestry, but also from their behavior when mated with another cock, No. 254, in which case they threw 12 per cent non-booted offspring. If now both parents are heterozygous they must produce 25 per cent recessives. This is the fact that forces us to conclude that clean shank is not recessive, but dominant and due to an inhibitor that frequently fails to dominate. In table 31 the two recessive varieties, mated inter se, produce no featherless shanks; the feathers grow freely as they do over the rest of the body. Some of the Silkies of table 31, however, are really heterozygous, with the dominant inhibitor not showing; consequently they throw a large proportion of non-booted offspring. In F1, as table 33 shows, the heterozygous offspring have a reduced boot and perfect dominance—complete inhibition of boot—in from 6 to 68 per cent. Dominance is most complete in the Silkies, where, the feathering being feeble, the inhibitor has, as it were, less to do in overcoming it. In F2 the expected 75 per cent dominant is approached in the case of the Silkies (62 per cent and 59 per cent, respectively), but inhibition is very imperfect in the Cochin and Brahma crosses, being reduced to between 25 and 2 per cent. More proof that boot is due to the absence of a factor rather than to its presence is found in this generation. If absence of boot is recessive, then, combined with imperfection of dominance, at least 25 per cent of the offspring should be recessive and probably a much larger proportion. The results in table 34 are absolutely incompatible with this hypothesis, since, in one case, there are only 2 per cent that can not develop boot. Two extracted clean-footed birds sometimes throw boot and sometimes not, and this result is to be expected on the hypothesis that clean-footedness is dominant, but two heavily booted birds can not transmit the boot inhibitor.

Table 34.Distribution of boot-grade in the F2 generation of booted × non-booted poultry.

COCHIN CROSSES.
Pen No. Mother. Father. Offspring.
No. Gen. Races. Grade. No. Gen. Races. Grade. Boot
present.
Boot
slight.
Boot
absent.
P. ct.
absent.
650 170 F1 Bl. Coch. × Wh. Legh Pr. 265 F1 Bl. Coch. × Wh. Legh. Pr. 19 2 2 8.7
650 263 F1 Do. Pr. 265 F1 Do. Pr. 36 2 2 5.0
650 278 F1 Do. Pr. 265 F1 Do. Pr. 26 4 4 11.8
650 361 F1 Do. Pr. 265 F1 Do. Pr. 24 2 9 25.7
650 364 F1 Do. Pr. 265 F1 Do. Pr. 39 5 3 6.4
Totals (179) 144 15 20 11.1
654 602 F1 Wh. Legh. × Bf. Coch Pr. 704 F1 Wh. Legh. × Bf. Coch Pr. 11 4 5 25.0
654 828 F1 Do. Pr. 704 F1 Do. Pr. 7 11 0 0.0
654 640 F1 Do. Pr. 704 F1 Do. Pr. 13 2 3 16.7
654 696 F1 Do. Pr. 704 F1 Do. Pr. 8 5 8 38.1
654 767 F1 Do. Pr. 704 F1 Do. Pr. 3 1 3 42.9
654 697 F1 Do. Pr. 704 F1 Do. Pr. 4 3 6 46.2
Totals (97) 46 26 25 25.8

TABLE 34.—Distribution of boot-grade in the F2 generation of booted × non-booted poultry—Continued.

DARK BRAHMA CROSSES.
Pen No. Mother. Father. Offspring.
No. Gen. Races. Grade. No. Gen. Races. Grade. Boot
present.
Boot
slight.
Boot
absent.
P. ct.
absent.
608 384 F1 Wh. Legh. × Dk. Brah. Pr. 409 F1 Wh. Legh. × Dk. Brah. Pr. 36 5 3 6.8
608 248 F1 Do. Pr. 409 F1 Do. Pr. 32 5 4 9.8
608 249 F1 Do. Pr. 409 F1 Do. Pr. 39 11 13 20.6
608 395 F1 Do. Pr. 409 F1 Do. Pr. 20 11 10 24.4
608 385 F1 Do. Pr. 409 F1 Do. Pr. 20 6 14 35.0
Totals (229) 147 38 44 19.2
659 762 F1 Wh. Legh. × Dk. Brah. Pr. 375 F1 Wh. Legh. × Dk. Brah. Pr. 18 4 1 4.4
659 503 F1 Do. Pr. 375 F1 Do. Pr. 23 6 2 6.5
659 382 F1 Do. Pr. 375 F1 Do. Pr. 10 2 1 7.7
659 250 F1 Do. Pr. 375 F1 Do. Pr. 33 7 5 11.1
659 737 F1 Do. Pr. 375 F1 Do. Pr. 19 2 3 12.5
659 387 F1 Do. Pr. 375 F1 Do. Pr. 16 6 4 15.4
Totals (162) 119 27 16 9.9
655 720 F1 Wh. Legh. × Dk. Brah. Pr. 666 F1 Wh. Legh. × Dk. Brah. Pr. 5 2 ... 0.0
655 724 F1 Do. Pr. 666 F1 Do. Pr. 6 1 ... 0.0
655 728 F1 Do. Pr. 666 F1 Do. Pr. 3 1 ... 0.0
655 730 F1 Do. Pr. 666 F1 Do. Pr. 4 ... ... 0.0
655 732 F1 Do. Pr. 666 F1 Do. Pr. 9 ... ... 0.0
655 734 F1 Do. Pr. 666 F1 Do. Pr. 3 ... ... 0.0
655 761 F1 Do. Pr. 666 F1 Do. Pr. 6 2 ... 0.0
655 800 F1 Do. Pr. 666 F1 Do. Pr. 1 ... ... 0.0
655 721 F1 Do. Pr. 666 F1 Do. Pr. 9 1 1 9.1
Totals (54) 46 7 1 1.9
655 724 F1 Wh. Legh. × Dk. Brah. Pr. 254 F1 Wh. Legh. × Dk. Brah. Pr. 3 ... ... 0.0
655 734 F1 Do. Pr. 254 F1 Do. Pr. 12 1 ... 0.0
655 800 F1 Do. Pr. 254 F1 Do. Pr. 13 ... 1 7.1
655 720 F1 Do. Pr. 254 F1 Do. Pr. 12 ... 1 7.7
655 728 F1 Do. Pr. 254 F1 Do. Pr. 8 1 1 10.0
655 761 F1 Do. Pr. 254 F1 Do. Pr. 17 4 4 16.0
655 732 F1 Do. Pr. 254 F1 Do. Pr. 8 1 2 18.2
655 730 F1 Do. Pr. 254 F1 Do. Pr. 7 ... 2 22.2
655 721 F1 Do. Pr. 254 F1 Do. Pr. 9 ... 3 25.0
Totals (110) 89 7 14 12.7
632 742 F1 Min. × Dk. Brah. Pr. 637 F1 Min. × Dk. Brah. Pr. 4 1 0 0.0
632 690 F1 Do. Pr. 637 F1 Do. Pr. 27 6 1 2.9
632 631 F1 Do. Pr. 637 F1 Do. Pr. 32 11 2 4.4
632 618 F1 Do. Pr. 637 F1 Do. Pr. 35 8 2 4.4
632 700 F1 Do. Pr. 637 F1 Do. Pr. 18 3 2 8.7
632 703 F1 Do. Pr. 637 F1 Do. Pr. 14 11 3 10.7
632 743 F1 Do. Pr. 637 F1 Do. Pr. 22 2 3 11.1
632 599 F1 Do. Pr. 637 F1 Do. Pr. 23 8 4 11.4
632 524 F1 Do. Pr. 637 F1 Do. Pr. 18 6 5 17.2
632 576 F1 Do. Pr. 637 F1 Do. Pr. 14 9 6 20.7
632 638 F1 Do. Pr. 637 F1 Do. Pr. 8 2 6 37.5
Totals (316) 215 67 34 10.8
Pen
No.
Mother. Father. Boot-grade in offspring.
No. Gen. Races. Gr. No. Gen. Races. Gr. 0 1 2 3 4 5 6 7 8 9 10 Aver-
age.
P. ct.
absent.
801 2526 F1 Min. × Dk. Brah. 2 5399 F1 W. L. × Dr. Brah. 8 ... ... ... ... 1 ... ... 1 ... ... 1 7.0 0.0
801 2831 F1 Do. 4 5399 F1 Do. 8 1 1 1 4 1 7 2 2 2 ... 2 5.0 4.3
801 1892 F1 Do. 3 5399 F1 Do. 8 1 1 0 1 2 ... 1 ... 1 1 1 5.0 11.1
Totals (35) 2 2 1 5 4 7 3 3 3 1 4 5.2 5.71

Table 34.Distribution of boot-grade in the F2 generation of booted × non-booted poultry—Continued.

SILKIE CROSSES.
Pen
No.
Mother. Father. Boot-grade in offspring.
No. Gen. Races. Gr. No. Gen. Races. Gr. 0 1 2 3 4 5 6 7 8 9 10 Aver-
age.
P. ct.
absent.
709 1955 F1 Silkie × Spanish 5 1578 F1 Silkie × Spanish 0 5 1 2 1 1 1 1 0 0 0 0 1.92 41.7
753 1966 F1 Silkie × Min 0 2573 F1 Min. × Silkie 0 19 4 2 2 ... 2 2 2 1 ... ... 1.71 55.9
709 1963 F1 Silkie × Spanish 7 1578 F1 Silkie × Spanish 0 23 6 1 6 7 ... ... ... ... ... ... 1.26 53.5
753 2575 F1 Silkie × Min 0 2573 F1 Silkie × Min. 0 15 3 7 ... ... ... ... ... ... ... ... 0.68 60.0
753 2071 F1 Do. 0 2573 F1 Do. 0 23 4 6 ... ... ... ... ... ... ... ... 0.49 69.7
709 1453 F1 Do. 1 1578 F1 Silkie × Spanish 0 24 11 3 ... ... ... ... ... ... ... ... 0.45 63.2
709 2223 F1 Silkie × Spanish 0 1578 F1 Do. 0 32 7 3 ... ... ... ... ... ... ... ... 0.31 76.2
Totals (227) 141 36 24 9 8 3 3 2 1 0 0 0.87 62.2
830 4082 F1 Silkie × W. Legh 2 3947 F1 Silkie × W. Legh 1 11 8 ... 7 1 ... ... ... ... ... ... 1.22 40.7
830 4079 F1 Do. 0 3947 F1 Do. 1 18 7 6 3 ... ... ... ... ... ... ... 0.82 53.0
830 5379 F1 Do. 0 3947 F1 Do. 1 18 4 5 3 ... ... ... ... ... ... ... 0.77 60.0
830 4081 F1 Do. 0 3947 F1 Do. 1 24 6 10 1 ... ... ... ... ... ... ... 0.71 58.5
830 5374 F1 Do. 0 3947 F1 Do. 1 11 3 3 1 ... ... ... ... ... ... ... 0.67 61.1
830 3946 F1 Do. 0 3947 F1 Do. 1 19 1 ... ... 1 ... ... ... ... ... ... 0.24 90.5
Totals (170) 101 29 24 14 2 0 0 0 0 0 0 0.75 59.4

The distribution of table 35 is characterized by its large variability. Although the numbers are small, there are evidences of two modes, one between grades 3 and 6, and the other at from 8 to 10; these evidently correspond to the modes of the typical Silkie and the typical Cochin respectively or to DR and RR types of booting respectively. The distribution of table 35 is additional evidence of the heterozygous nature of the Silkie boot.

Table 35.Distribution of boot-grades in Silkie × Cochin crosses.

Pen
No.
Mother. Father. Boot-grades in offspring.
No. Gen. Races. Gra. No. Gen. Races. Gra. 0 1 2 3 4 5 6 7 8 9 10 Aver-
age.
P. ct.
abs.
821 5925 F1 Silk. × Coch. 7 6095 F1 Silk. × Coch. 7 ... ... ... ... 1 ... ... 1 3 1 1 7.7 0.0
821 7408 F1 Do. 4 6095 F1 Do. 7 ... ... ... 1 2 2 3 ... 2 1 2 6.5 0.0
821 7413 F1 Do. 3 6095 F1 Do. 7 2 0 3 1 0 1 1 0 0 1 1 3.9 20.0
821 7416 F1 Do. 5 6095 F1 Do. 7 ... ... ... 3 1 0 4 0 3 3 2 6.8 0.0
821 7417 F1 Do. ... 6095 F1 Do. 7 ... ... ... ... ... ... ... 1 ... 1 4 9.3 0.0
821 7418 F1 Do. 4 6095 F1 Do. 7 ... ... 1 ... 2 1 1 1 1 ... 1 5.8 0.0
821 7423 F1 Do. 6 6095 F1 Do. 7 ... ... ... 1 ... 2 ... 2 2 ... 2 7.0 0.0
821 7428 F1 Do. ... 6095 F1 Do. 7 1 ... ... ... ... 1 ... ... 1 ... ... 4.3 33.3
821 7429 F1 Do. 8 6095 F1 Do. 7 ... ... ... 1 1 1 ... ... ... 1 1 6.2 0.0
Totals (77) 3 0 4 7 7 8 9 5 12 8 14 6.42 3.90
29 48

We are now in a position to consider the effect of back crosses (table 36). The contrast between the totals in tables 36 and 37 is very great. The strict Mendelian expectation is: in the DR × D crosses 50 per cent DD (clean-footed) and 50 per cent heterozygous, which, with imperfect dominance, might be expected to show foot-feathering. Actually about 46 per cent are clean-footed. In the DR × R crosses expectation is that 50 per cent certainly (the extracted recessives) and 50 per cent more possibly will have the shanks feathered, on account of imperfect dominance of the heterozygotes. Actually all have feathered feet. These statistics thus confirm the view of the dominance of the inhibiting factor. Were clean shank recessive, then the DR × R crosses must give 50 per cent clean-footed and probably over. The actual result, none clean-footed, is not in accord with the latter assumption.

Table 36.Distribution of boot-grade in DR × D (non-booted) crosses.

Pen
No.
Mother. Father. Boot-grade in offspring.
No. Gen. Races. Grade. No. Gen. Race. Grade. Present. Slight. Absent. Per cent.
present.
653 508 F1 Wh. Legh. × Bf. Coch. Pr. 117 P. Game. 0 3 4 6 46.2
653 508 F1 Do. Pr. 116 P. Do. 0 6 5 4 26.7
653 577 F1 R × Bf. Coch. 3 117 P. Do. 0 1 0 7 87.5
653 577 F1 Do. 3 116 P. Do. 0 1 3 2 33.3
653 587 F1 Do. 1 117 P. Do. 0 1 2 4 57.1
653 587 F1 Do. 1 116 P. Do. 0 3 3 2 25.0
653 635 F1 Do. 3 117 P. Do. 0 ... 1 6 85.7
653 635 F1 Do. 3 116 P. Do. 0 2 2 1 20.0
653 652 F1 Do. 5 117 P. Do. 0 5 8 4 23.5
653 652 F1 Do. 5 116 P. Do. 0 1 2 2 40.0
653 691 F1 Do. Pr. 117 P. Do. 0 2 2 1 20.0
653 705 F1 Do. 2 117 P. Do. 0 3 2 5 50.0
653 705 F1 Do. 2 116 P. Do. 0 1 1 5 71.4
653 713 F1 Do. Pr. 117 P. Do. 0 ... 0 4 100.0
653 713 F1 Do. Pr. 116 P. Do. 0 1 1 3 60.0
653 760 F1 Do. Pr. 117 P. Do. 0 2 2 6 60.0
653 760 F1 Do. Pr. 116 P. Do. 0 0 3 2 40.0
653 799 F1 Do. 3 117 P. Do. 0 2 0 3 60.0
Total (143) 34 42 67 46.9
661 635 F1 Bf. Coch. × Game. Pr. 466 P. Game. 0 1 ... 2 66.7
661 635 F1 Do. Pr. 428 P. Do. 0 2 ... 1 33.3
661 691 F1 Do. Pr. 466 P. Do. 0 2 ... 2 50.0
661 691 F1 Do. Pr. 428 P. Do. 0 2 ... 1 33.3
661 799 F1 Do. Pr. 466 P. Do. 0 3 ... 2 40.0
661 799 F1 Do. Pr. 428 P. Do. 0 4 ... 1 20.0
Total (23) 14 0 9 39.1
Grand Total (166) 48 42 76 45.8

Table 37.Distribution of boot-grade in DR × RR (booted) crosses.

[A]Pure-blooded Silkie assumed heterozygous to boot.
Pen
No.
Mother. Father. Boot-grade in offspring.
No. Gen. Race. Gr. No. Gen. Race. Gr. 0 1 2 3 4 5 6 7 8 9 10
851 838 P. Cochin. 8 7526 [A]F1 Silkie. 3 ... ... ... 3 2 4 3 ... ... 2 2
851 840 P. Do. 10 7526 F1 Do. 3 ... ... ... ... 1 ... 1 ... ... 1 ...
851 841 P. Do. 10 7526 F1 Do. 3 ... ... ... ... 1 ... 1 ... 1 ... ...
851 1002 P. Do. 8 7526 F1 Do. 3 ... ... ... 3 1 2 1 2 3 1 1
851 2073 P. Do. 7 7526 F1 Do. 3 ... ... 1 1 1 ... 1 1 1 3 2
851 2299 P. Do. 9 7526 F1 Do. 3 ... ... ... ... 2 2 1 1 ... ... 2
851 3410 P. Do. 9 7526 F1 Do. 3 ... ... ... ... ... 4 3 2 1 5 1
851 5567 P. Do. 9 7526 F1 Do. 3 ... ... ... ... 2 ... ... ... 3 3 5
851 6956 P. Do. 8 7526 F1 Do. 3 ... ... ... ... 3 3 ... 2 2 ... 5
Totals (99) 0 0 1 7 13 15 11 8 11 15 18

Numerous observations have been made upon the progeny of parents belonging to hybrid generations beyond the first. Owing to the extreme imperfection of dominance it is rarely possible to say with certainty from inspection whether a given bird has germ-cells dominant or recessive, or both, with reference to booting; only breeding enables us to make a decision. There is an exception, however, in the case of pure extracted recessives. They are distinguished by heavy booting and produce only booted offspring. I propose to give, in detail, the matings of these later generations and their progeny, the families being arranged in decreasing order of average grade of booting (table 38).

Table 38.Distribution of boot-grades in offspring of parents one or both of which belong to a hybrid generation beyond the first.

B = Brahma; C = Cochin; G = Game; L = Leghorn; M = Minorca; S = Silkie; Sp = Spanish; T = Tosa; WL = White Leghorn

>
3 5 2 7 5 6 2 4 ... 5.5
47 732 953 F2 M × B 3 2442 F2 M × B 6 Do. ... 2 2 3 8 9 5 3 7 6 ... 5.5
48 801 7528 F1 Do. 4 4562 F2 Do. 8 Do. ... ... ... 1 4 2 1 ... 1 ... 1 5.3
49 731 2116 F2 Do. 10 1249 F2 WL × B 7 R × R ... 1 1 1 2 3 0 2 2 1 ... 5.2
50 745 2115 F2 C × T 4 1258 F2 B × T 4 DR × DR ... ... ... 2 1 6 4 2 ... ... ... 5.2
51 801 6843 F2 B × T 3 4562 F2 Do. 8 DR × R ... ... ... ... 3 2 1 ... 1 ... ... 5.1
52 801 2831 F1 M × B 4 5399 F2 Do. 8 Do. 1 1 1 4 1 7 2 2 2 ... 2 5.0
53 801 1892 F1 Do. 3 5399 F2 Do. 8 Do. 1 1 ... 1 2 ... 1 0 1 1 1 5.0
54 801 7528 F1 Do. 4 4562 F2 Do. 8 Do. ... ... ... 1 2 1 1 ... 1 ... ... 5.0
55 731 1755 F2 WL × B 6 1249 F2 WL × B 7 R × R ... ... ... ... 2 1 4 1 2 ... ... 5.0
56 745 2513 F3 C × T 4 1258 F2 B × T 4 DR × DR ... ... ... ... 2 5 2 ... 3 1 ... 5.0
57 839 3950 F2 M × B 4 4348 F2 M × B 3 Do. ... 2 3 3 2 4 1 1 1 2 2 4.95
58 754 873 F2 B × T 3 871 F2 B × T 2 Do. 1 2 1 4 1 ... ... ... 8 ... ... 4.94
59 806 599 F2 M × B 3 5257 F2 M × B 7 DR × R ... ... 2 1 2 2 1 ... 1 ... ... Table 38.Distribution of boot-grades in offspring of parents one or both of which belong to a hybrid generation beyond the first—Continued.

B = Brahma; C = Cochin; G = Game; L = Leghorn; M = Minorca; S = Silkie; Sp = Spanish; T = Tosa; WL = White Leghorn.

Serial
No.
Pen
No.
Mother. Father. Mating. Boot-grade in offspring.
No. Gen. Races. Gr. No. Gen. Races. Gr. 0 1 2 3 4 5 6 7 8 9 10 Av.
62 732 2407 F2 M × B 2 2442 F2 M × B 6 DR × R ... ... 1 3 4 1 1 2 ... 1 ... 4.69
63 701 894 F2 L × B 7 1898 F2 L × B 3 R × DR 1 1 2 8 6 1 2 2 4 2 ... 4.62
64 760 994 F2 B × T 3 1270 F2 B × T 3 DR × DR ... ... ... ... 4 2 1 ... ... ... ... 4.57
65 760 981 F2 Do. 3 1270 F2 Do. 3 Do. 1 ... 3 6 1 4 2 7 ... ... ... 4.54
66 701 1772 F2 L × B 6 1898 F2 L × B 3 R × DR ... ... ... 4 7 2 2 2 ... ... ... 4.47
67 839 3541 F1 M × B 6 4348 F2 M × B 3 DR × DR 4 1 4 ... ... 4 2 ... 2 1 2 4.30
68 842 1645 F2 Do. 2 4385 F2 Do. 4 Do. 3 2 6 5 6 6 3 0 2 4 1 4.29
69 770 2049 F2 L × B 3 926 F2 Do. 3 Do. 9 3 1 6 8 2 6 6 3 1 3 4.29
70 731 2577 F1.5 L × C 4 1249 F2 L × B 7 DR × R ... ... 2 2 3 2 2 1 ... ... ... 4.25
71 701 250 F1 L × B 3 1898 F2 Do. 3 DR × DR 3 3 5 8 12 10 10 6 1 ... ... 4.22
72 701 1335 F2 T × L × B 8 1898 F2 Do. 3 R × DR ... ... 1 9 6 6 4 1 ... ... ... 4.22
73 806 4767 F3 M × B 3 5257 F3 M × B 7 DR × R ... ... 1 ... 2 1 1 ... ... ... ... 4.20
74 740 1439 F2 C × L 2 1145 F2 C × L 3 DR × DR 3 ... 1 3 6 4 2 ... 2 1 ... 4.18
75 754 3126 F2 B × T 4 871 F2 B × T 3 Do. ... 2 5 11 7 10 5 0 2 1 ... 4.14
76 770 1645 F2 M × B 4 926 F2 M × B 3 Do. 3 2 1 9 5 5 2 2 3 1 ... 4.10
77 731 249 F1 L × B 3 1249 F2 L × B 7 DR × R 7 4 6 5 7 5 9 3 6 1 ... 4.08
78 732 703 F1 M × B 3 2442 F2 M × B 6 Do. 1 3 13 13 8 6 7 6 3 ... ... 4.07
79 770 720 F1 B × L 4 926 F2 Do. 3 DR × DR 6 1 3 9 5 4 5 1 4 1 1 4.05
80 732 2441 F2 M × B 0 2442 F2 Do. 6 DR × R ... 1 6 8 2 6 0 3 1 ... ... 4.00
81 760 1042 F2 B × T 3 1270 F2 B × T 2 DR × DR 2 3 3 9 3 5 8 2 0 1 ... 4.00
82 731 384 F1 L × B 4 1249 F2 L × B 7 DR × R 2 1 4 4 3 2 4 0 1 1 ... 3.82
83 814 4566 F2 B × T 2 3975 F2 B × T 9 Do. 1 4 2 4 3 2 1 ... ... ... ... 3.82
84 732 599 F1 M × B 3 2442 F2 M × B 6 Do. 6 5 23 10 5 3 4 5 8 3 1 3.78
85 770 761 F1 B × L 3 926 F2 Do. 3 DR × DR 7 3 5 3 7 7 2 6 1 1 ... 3.71
86 731 1770 F2 Do. 7 1249 F2 L × B 7 DR × R 1 ... 8 6 9 3 2 ... 2 ... ... 3.65
87 861 5165 F2 T × C 10 95 F1 T × C 5 R × DR ... ... ... 10 3 2 1 ... ... ... ... 3.63
88 754 3175 F2 B × T 2 871 F2 B × T 2 DR × DR 1 ... 2 1 3 4 ... ... ... ... ... 3.55
89 731 2102 F2 L × B 1 1249 F2 L × B 7 DR × R 1 0 4 2 4 1 1 1 ... ... ... 3.43
90 840 1755 F2 M × B 6 4177 F2 Do. 2 R × DR ... ... 6 7 7 3 1 ... ... ... ... 3.42
91 701 2576 F2 L × B 2 1898 F2 Do. 3 DR × DR 2 1 1 8 11 2 1 ... ... ... ... 3.35
92 842 2049 F1 Do. 3 4385 F2 M × B 4 Do. 11 1 2 8 5 3 1 0 2 2 2 3.35
93 754 853 F2 B × T 1 871 F2 B × T 3 Do. 2 3 4 6 4 1 6 ... ... ... ... 3.31
94 826 2652 F1 M × B 3 4093 F2 M × B 0 Do. 8 2 1 8 1 ... ... 1 2 3 ... 3.28
95 754 1052 F2 B × T 2 871 F2 B × T 2 Do. 3 ... 7 9 9 5 2 ... ... ... ... 3.26
96 701 965 F2 T × L × B 0 1898 F2 L × B 3 Do. 1 4 6 12 8 4 0 2 0 ... ... 3.19
97 732 1833 F2 M × B 1 2442 F2 M × B 6 DR × R 1 1 7 6 6 4 1 ... ... ... ... 3.19
98 732 631 F1 Do. 3 2442 F2 Do. 6 Do. 3 4 10 16 12 4 1 2 ... ... ... 3.08
99 754 862 F2 B × T 1 871 F2 B × T 2 DR × DR 1 5 10 17 10 4 1 ... ... ... ... 2.96
100 837 5641 F2 T × L × B 0 4288 F3 L × B 2 Do. 1 2 2 3 ... 2 ... 1 ... ... ... 2.91
101 840 3841 F2 L × B 0 4177 F2 Do. 2 D × DR 3 3 2 6 4 2 ... ... ... ... ... 2.86
102 701 721 F1 Do. 2 1898 F2 Do. 3 DR × DR 2 4 3 8 3 2 2 ... ... ... ... 2.83
103 839 3949 F2 Do. 4 4348 F2 Do. 3 Do. 1 2 ... ... 1 1 1 ... ... ... ... 2.83
104 840 732 F1 Do. 3 4177 F2 Do. 2 Do. 7 6 9 8 7 2 1 2 ... 1 ... 2.67
105 840 249 F1 Do. 3 4177 F2 Do. 2 Do. 7 3 5 6 2 9 ... ... ... ... ... 2.62
106 840 3916 F1.5 Do. 2 4177 F2 Do. 2 Do. 5 1 4 2 2 2 1 ... ... ... ... 2.29
107 842 4945 F2 M,L × B 1 4385 F2 M × B 4 Do. 9 3 6 5 1 2 4 ... ... ... ... 2.27
108 731 2595 F2 L × B 1 1249 F2 L × B 7 D × R 6 6 7 1 ... ... ... ... ... ... ... 2.15
109 840 5169 F2 Do. 3 4177 F2 Do. 2 DR × DR 6 2 5 5 2 2 ... ... ... ... ... 2.05
110 837 5667 F3 Do. 2 4288 F3 Do. 2 Do. 2 1 2 ... 1 1 ... ... ... ... ... 2.00
111 749 1355 F2 G × C 2 1854 F2 G(C × L) 0 DR × D ... 2 5 1 ... ... ... ... ... ... ... 1.87
112 824 3901 F2 M × S 1 5095 F2 M × S 1 DR × DR 17 3 2 3 ... 2 ... ... 1 2 ... 1.73
113 751 1254 F2 L × B 0 1139 F2 L × B 8 D × R 17 5 5 4 3 3 ... 1 ... ... ... 1.63
114 749 816 F1 Do. 2 1854 F2 G(C × L) 0 DR × D 6 7 3 5 1 ... ... ... ... ... ... 1.45
115 749 929 F2 G × C 0 1854 F2 Do. 0 D × D 8 3 1 ... ... ... 1 ... ... 1 ... 1.43
116 749 819 F1 L × B 1 1854 F1.5 G(C × L) 0 DR × D 9 3 5 3 ... ... ... ... ... ... ... 1.10
117 804 5099 F2 S × Sp 0 3823 F1 S × Sp 0 D × D 2 ... ... 1 ... ... ... ... ... ... ... 1.00
118 804 6043 F2 Do. 1 3823 F1 Do. 0 Do. ... 1 ... ... ... ... ... ... ... ... ... 1.00
119 817 5730 F1 L × Sp 0 3900 F2 Do. 1 D × DR 3 3 1 ... ... ... ... ... ... ... ... 0.71
120 817 4696 F1 Do. 0 3900 F2 Do. 1 Do. 9 7 3 ... ... ... ... ... ... ... ... 0.68
121 817 6046 F2 S × M 0 3900 F2 Do. 1 Do. 10 ... 3 ... ... ... ... ... ... In table 38 I have given in the section lying between that headed "Father" and that headed "Offspring" the "Matings." This column differs from the others of the table in not being, in general, based upon observation, but upon a sometimes complicated judgment. Of course, all of the F1 generation, where this generation occurs, may be taken as of DR composition; but the decision as to whether a given individual of F2 is a DR, an extracted dominant, or an extracted recessive is not always easy, because of the manifestation of imperfect dominance. But the assignments are by no means arbitrary. Taking the Brahma crosses, which are by far the most numerous, we see, from tables 31, B and 33, that those F2 individuals that have a boot of grade 6 or higher are almost certainly extracted recessives (which are equivalent to pure-bred Dark Brahmas). Those with a grade of 3 or even 4 and lower to 2 or even 1 are probably heterozygotes, while those with grade 0 and some of those with grade 1 are extracted dominants. In cases of doubt the distribution of grades in the offspring will give the deciding vote. In case the individual has been used as a parent in more than one mating the results in all the matings are taken into account, for the germinal constitution of an individual must be regarded as fixed at all times and in all matings. The assignment under "Matings" has, then, been made by the application of the above rules.

In tables 39 to 43 there are grouped together the progeny from matings of the same sort, selecting from table 38 the crosses into which the Dark Brahma enters as the booted parent.

Table 39.RR × RR crosses from table 38.

Serial No. Boot-grade in offspring. Parental grades.
0 1 2 3 4 5 6 7 8 9 10 Avge. Female. Male. Average.
1 ... ... ... ... ... ... ... ... ... 10 15 9.6 7 9 8.0
2 ... ... ... ... ... ... ... ... ... 1 1 9.5 4 8 6.0
3 ... ... ... ... ... ... ... ... 1 3 4 9.4 5 9 7.0
4 ... ... ... ... ... ... ... ... 1 1 2 9.3 6 7 6.5
5 ... ... ... ... ... 1 ... ... 3 4 9 9.1 9 9 9.0
6 ... ... ... ... ... 1 1 1 ... 2 7 8.8 8 9 8.5
7 ... ... ... ... ... ... 1 1 1 1 4 8.8 2 9 5.5
8 ... ... ... ... ... ... 1 1 2 2 3 8.6 7 5 6.0
9 ... ... ... ... 1 ... ... 1 4 2 3 8.3 7 5 6.0
10 ... ... ... ... ... ... 1 2 4 3 ... 7.9 8 6 7.0
11 ... ... ... ... 1 1 ... 3 ... 6 1 7.8 5 5 5.0
13 ... ... ... ... ... 1 1 2 1 1 1 7.4 6 7 6.5
18 ... ... ... ... 1 ... 2 ... 2 ... 1 7.0 10 9 9.5
21 ... ... ... ... 3 7 7 7 7 5 2 6.8 9 6 7.5
25 ... ... ... ... ... 2 1 ... ... ... 1 6.5 7 7 7.0
37 ... ... ... 2 8 5 6 2 7 3 ... 6.0 8 6 7.0
39 ... ... ... 5 4 6 6 1 11 1 ... 5.9 8 8 8.0
41 ... ... 1 2 1 8 2 3 6 ... ... 5.8 9 7 8.0
49 ... 1 1 1 2 3 ... 2 2 1 ... 5.2 10 7 8.5
55 ... ... ... 2 1 4 1 2 ... ... ... 5.0 6 7 6.5
Totals (287) ... 1 2 12 22 39 30 28 53 46 54 7.25
Per cent. ... 0.3 0.7 4.2 7.7 13.6 10.5 9.8 18.5 16.0 18.8 ...

Table 40.DR × RR crosses from table 38.

Serial No. Boot-grade in offspring.
0 1 2 3 4 5 6 7 8 9 10 Average.
14 ... 2 1 ... 1 3 ... 1 3 6 5 7.2
15 ... ... ... ... 3 2 3 1 1 1 5 7.1
16 ... ... ... 1 4 1 2 7 6 1 ... 7.1
17 ... ... ... ... 1 ... ... 1 ... ... 1 7.0
20 ... ... ... 1 1 2 2 ... 1 1 3 6.9
22 ... ... ... 2 1 ... 1 ... ... 2 2 6.8
23 ... ... ... 2 2 2 1 ... ... 3 3 6.7
24 ... ... ... ... 1 ... ... 1 ... 1 ... 6.7
26 1 ... ... 1 4 6 9 4 4 6 3 6.5
27 ... ... ... 2 3 ... 3 1 1 4 1 6.5
28 ... ... 1 2 6 13 11 5 11 8 3 6.3
29 ... ... ... 2 2 5 2 1 1 3 2 6.3
30 ... ... ... 1 3 8 5 2 ... 2 4 6.3
31 ... ... ... 6 4 1 ... 1 1 1 6 6.2
32 ... ... ... 1 5 4 3 1 2 2 2 6.1
33 ... ... ... 5 6 11 12 7 4 9 ... 6.1
34 ... ... 1 ... ... ... 1 ... 2 ... ... 6.0
35 ... ... 1 ... ... 1 3 ... 1 1 ... 6.0
36 ... ... 2 3 3 ... 2 ... ... 5 2 6.0
40 ... 1 2 3 2 5 3 5 9 1 ... 5.8
42 ... ... 1 3 9 5 8 4 7 2 ... 5.7
43 ... ... ... ... 7 4 3 2 3 1 ... 5.7
44 ... ... ... 1 ... 1 ... ... ... 1 ... 5.7
45 ... ... 1 2 5 3 1 1 1 4 ... 5.6
46 ... ... 3 5 2 7 5 6 2 4 ... 5.5
47 ... 2 2 3 8 9 5 3 7 6 ... 5.5
48 ... ... ... 1 4 2 1 ... 1 ... 1 5.3
51 ... ... ... ... 3 2 1 ... 1 ... ... 5.1
52 1 1 1 4 1 7 2 2 2 ... 2 5.0
53 1 1 ... 1 2 ... 1 ... 1 1 1 5.0
54 ... ... ... 1 2 1 1 ... 1 ... ... 5.0
59 ... ... 2 1 2 2 1 ... 1 ... ... 4.9
60 ... ... 2 19 8 13 6 4 5 2 1 4.8
61 ... 1 1 1 1 ... 1 ... 1 1 ... 4.8
62 ... ... 1 3 4 1 1 2 ... 1 ... 4.7
63 1 1 2 8 6 1 2 2 4 2 ... 4.6
66 ... ... ... 4 7 2 2 2 ... ... ... 4.5
70 ... ... 2 2 3 2 2 1 ... ... ... 4.3
72 ... ... 1 9 6 6 4 1 ... ... ... 4.2
73 ... ... 1 ... 2 1 1 ... ... ... ... 4.2
77 7 4 6 5 7 5 9 3 6 1 ... 4.1
78 1 3 13 13 8 6 7 6 3 ... ... 4.1
80 ... 1 6 8 2 6 ... 3 1 ... ... 4.0
82 2 1 4 4 3 2 4 ... 1 1 ... 3.8
83 1 4 2 4 3 2 1 ... ... ... ... 3.8
84 6 5 23 10 5 3 4 5 8 3 1 3.8
86 1 ... 8 6 9 3 2 ... 2 ... ... 3.7
89 1 ... 4 2 4 1 1 1 ... ... ... 3.4
90 ... ... 6 7 7 3 1 ... ... ... ... 3.4
97 1 1 7 6 6 4 1 ... ... ... ... 3.2
98 3 4 10 16 12 4 1 2 ... ... ... 3.1
Total (1199) 27 32 117 181 200 172 142 88 105 87 48 5.04
Per cent. 2.3 2.7 9.8 15.1 16.7 14.3 11.9 7.3 8.8 7.2 4.0 ...

Table 41.DR × DD crosses.

Serial No. Boot-grade in offspring.
0 1 2 3 4 5 Average.
101 3 3 2 6 4 2 2.9
113 6 7 3 5 1 ... 1.5
116 9 3 5 3 ... ... 1.1
Total (62) 18 13 10 14 5 2 1.69
Per cent. 29.5 21.3 16.4 23.0 8.2 1.6 ...

Table 42.DR x DR crosses.

Serial No. Boot-grade in offspring.
0 1 2 3 4 5 6 7 8 9 10 Average.
19 ... ... ... ... 1 1 ... ... ... 1 ... 7.0
54 ... ... ... 2 1 6 4 2 ... ... ... 5.2
56 ... ... ... 2 5 2 ... 3 1 ... ... 5.0
57 ... 2 3 3 2 4 1 1 1 2 2 5.0
58 1 2 1 4 1 ... ... ... 8 ... ... 4.9
59 ... ... 2 1 2 2 1 ... 1 ... ... 4.9
64 ... ... ... ... 4 2 1 ... ... ... ... 4.6
65 1 ... 3 6 1 4 2 7 ... ... ... 4.5
67 4 1 4 ... ... 4 2 ... 2 1 2 4.3
68 3 2 6 5 6 6 3 ... 2 4 1 4.3
69 9 3 1 6 8 2 6 6 3 1 3 4.3
71 ... ... 2 2 3 2 2 1 ... ... ... 4.3
75 ... 2 5 11 7 10 5 ... 2 1 ... 4.1
76 3 2 1 9 5 5 2 2 3 1 ... 4.1
79 6 1 3 9 5 4 5 1 4 1 1 4.1
81 2 3 3 9 3 5 8 2 ... 1 ... 4.0
85 7 3 5 3 7 7 2 6 1 1 ... 3.7
88 1 ... 2 1 3 4 ... ... ... ... ... 3.6
91 2 1 1 8 11 2 1 ... ... ... ... 3.4
92 11 1 2 8 5 3 1 ... 2 2 2 3.4
93 2 3 4 6 4 1 6 ... ... ... ... 3.3
94 8 2 1 8 1 ... ... 1 2 3 ... 3.3
95 3 ... 7 9 5 2 ... ... ... ... ... 3.3
96 1 4 6 12 8 4 ... 2 ... ... ... 3.2
99 1 5 10 17 10 4 1 ... ... ... ... 3.0
100 1 2 2 3 ... 2 ... 1 ... ... ... 2.9
102 2 4 3 8 3 2 2 ... ... ... ... 2.8
103 1 2 ... ... 1 1 1 ... ... ... ... 2.8
104 7 6 9 8 7 2 1 2 ... 1 ... 2.7
105 7 3 5 6 2 9 ... ... ... ... ... 2.6
106 5 1 4 2 2 2 1 ... ... ... ... 2.3
107 9 3 6 5 1 2 4 ... ... ... ... 2.3
109 6 2 5 5 2 2 ... ... ... ... ... 2.1
110 2 1 2 ... 1 1 ... ... ... ... ... 2.0
Total (851) 105 61 108 178 127 109 62 37 32 20 12 3.59
Per cent. 12.3 7.2 12.7 20.9 14.9 12.8 7.3 4.4 3.8 2.3 1.4 ...

Table 43.DD x DD (Silkie crosses).

Serial No. Boot-grade in offspring.
0 1 2 3 Average.
117 2 ... ... 1 1.00
118 ... 1 ... ... 1.00
128 26 6 1 1 0.32
130 11 4 ... ... 0.27
131 18 2 1 ... 0.19
132 19 ... ... ... 0.0
133 33 ... ... ... 0.0
134 8 ... ... ... 0.0
135 19 ... ... ... 0.0
136 16 ... ... ... 0.0
Total (169) 152 13 2 2 0.14
Per cent. 89.9 7.7 1.2 1.2 ...

The significance of the data given in tables 39 to 43 is best brought out by summarizing them. Especially instructive is a comparison of the pure-bred with the hybrids. Since the data are most complete in the case of the Brahma crosses, these will be considered in most detail. So far as they go, the results with the Cochins and Silkies are entirely confirmatory.

Table 44 shows clearly, first, that there are families of two booted parents that never fail to produce booted offspring. There is, however, even in pure-bred booted races, a marked variability in the grade of booting, extending from 3 (or 4) to 10. The significance of this variability must be left for future investigations. There is in the least boot, as it were, an extension of the field of activity of the feather-inhibiting factor that is always present on the hinder aspect of the shank, so that it interferes with the development of feathers on the inner face of the shank also. In the first hybrid generation all somatic cells are hybrid. The feather inhibitor is present in the skin of the shank, but its strength is diluted by the presence in the same cells of a protoplasm devoid of the inhibiting property. Consequently, the prevailing grade of the boot falls from 6 (or 10) to 3. Despite the dilution, inhibition is complete in about 8 per cent of the offspring (grade 0); in about 10 per cent of the offspring the inhibiting factor is so weak that the boot develops as in the pure-blooded Brahma. When, as a result of inbreeding F1's, the feather-inhibiting factor is eliminated from certain offspring, and such full-feathered birds are bred together, we find a return of the mode to high numbers, such as 8 to 10 (but also 5). There is no doubt of segregation.

Table 44.Brahma crosses. (All entries are percentages.)

If a heterozygous bird be mated to a recessive the variability of the offspring is much increased, owing to the occurrence in the progeny of both DR and RR individuals (table 40). The offspring do not, to be sure, fall into two distinct and well-defined types, as in typical Mendelian cases; but one part of the range of variation agrees fairly with that of pure RR's, i. e., Brahmas, and the remainder with that of heterozygotes. And if we make the division in the middle of the middle class, viz, 5, we shall find a close approximation to that equality of extracted recessives and heterozygotes that the segregation theory calls for (table 44).

If, again, two heterozygous birds be mated, the variability is still greater and the proportion of clean-footed offspring rises to 12 per cent. These, together with some of the extremely slightly booted offspring, represent the extracted dominants. The whole range now falls into three regions divided by the middle of grades 2 and 5. These regions correspond to the DD's, the DR's, and the RR's of typical cases of segregation, and their relative proportions are approximately as 25: 50: 25.

Finally, if a heterozygote be mated to an extracted dominant the proportion of clean-footed offspring rises to about 30 per cent and the whole range of variation falls readily into two parts, the one comprising grades 0 and 1, the other grades 2 and above. The first includes the DD offspring; the second, the DR's; and their frequency is equal. One will not fail to note that we are not here dealing with a case of blending simply, and the inheritance of the blend; such a view is negatived by the fact of the much greater variability of DR × DR cross over the simple D × R cross of the first generation. One may safely conclude, then, that, despite the apparent blending of booting characters in the first generation of hybrids, true segregation takes place. But this is always to be seen through the veil of imperfect dominance.

A casual examination of table 38 would seem to show a correlation between the grade of booting of the parents and that of the average of their progeny. Thus, on the whole, the parental grades run high in the upper part of the table and run low in the lower part. This relation would thus seem to confirm Castle's conclusion for polydactylism in guinea-pigs that there is an inheritance of the degree of a character. One consequence of such an inheritance would be that it would be possible in a few generations to increase or diminish the grade of a character and fix any required grade in the germ-plasm. A more careful consideration of the facts of the case shows that this relation has another interpretation. The grade of boot of the different parents varies largely because their gametic constitution is diverse. As table 39 shows, the parents of the upper part of table 38 are chiefly extracted recessives, and consequently their booting and that of their offspring are characterized by high grades. On the other hand, the parents of the lower part of the table are heterozygous or extracted dominants and, consequently, their grades and also those of their offspring average low. On account of the lack of homogeneity of the families in table 38, one can draw from it no proper conclusions as to relation between parental and filial grades. On the other hand, from a homogeneous table, like table 39, we can hope to reach a conclusion as to the existence of such a relation. I have calculated, in the usual biometric fashion, the coefficient of correlation between average parental and filial grades, and found it to be -0.17±0.13. This can only be interpreted to mean that in a homogeneous assemblage of families there is no correlation between the grade of booting of parents and offspring.

                                                                                                                                                                                                                                                                                                           

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