R esearch Bulletin No.

105

October, 1927

The Genetics of Plumage Color

in Poultry
By O. W.

KNOX

AGRICULTURAL EXPERIMENT STATION

IOWA STATE OOLLEGE OF AGRICULTURE
AND MEOHANIC ARTS

POULTRY SECTION

AMES, IOWA

SUMMARY

1.
2.
3.

4.
5.
6.

7.
8.
9.

The chromogen gene for color, CC, acted as a simple

dominant.
The extension of black pigment, EE, was inherited on
a simple monohybrid basis.
The heterozygous extension and chromogen genes
(Cc Ee) had a balanced relationship. This combination of heterozygous factors weakened the expression
of the black melanic pigment in the presence of two
or more doses of buff.
Buff color was controlled on a dihybrid basis, the
following symbols being used, Bu Bu Bu' Bu'.
Three or more doses of the buff genes in the presence of homozygous color factors, CC, were epistatic
to black, and gave a buff color.
Two or more doses of the buff color determiners
were epistatic to black when they were in the presence of both the extension of black and chromogen
factors in a heterozygous state.
When the extension factor was homozygous, EE, the
buff color was hypostatic to black if not present in
three or more doses.
All the factors included thus far in the summary
were autosomal.
The sex-linked factor for barring (BB and B-) appeared to act as a simple dominant to black and buff
colors.

The Genetics of Plumage Color in Poultry

By C. W.

KNOX*

Much information on the inheritance of the different color

factors in poultry has accumulated since Bateson (2) first took
up the study shortly after the rediscovery of Mendelian inheritance in 1900. Many investigations of the various plumage
colors conducted in England, Europe, and the United States
have done much to place the colors and patterns on a factorial
basis, with the exception of buff and red. The main object of
the writer's investigations was to place buff color on a factorial
basis and to verify, in some measure, the previous experiments
of other investigators in respect to the inheritance of white,
black and barring.
Th e author's experiments with white, black, buff and barred
colors were so devised as to enable him to build up by inductive and deductive methods the inheritance of the plumage color
genes in White Plymouth Rocks, Black Langshans and Buff
Orpingtons.
For the purpose of simplifying the interpretation of the results obtained, the investigation of color is discussed in five distinct divisions, in the following order:
1.
2.
3.
4.
5.

Inheritance of color expression (white).

The heredity of the extension genes for black pigment.
The inheritance or the genes for the production of buff pig'
ment.
The heredity of the factor for barring in domestic fowls.
The genetic plumage color of White Plymouth Rocks, Black
Langshans and Buff Orpingtons.

Each division is treated separately in respect to the review

of previous literature, crosses used, data accumulated, discussion and results obtained.
The results from the different sections are summarized and
placed at the conclusion of this publication.
MATERIAL AND METHODS

The birds used in this study were purebred. The original

parental generations had been bred for several years at this
station. The following three crosses, involving three plumage
'Paper No. 12 from Iowa State College Poultry Husbandry Department, submitted
in part'al fulfillment of the requirements for the degree, Doctor of Philosophy.
"Acknowledgement is hereby made to H. A . Bittenbender of the Poultry Depart
ment and E. W. Lindstrom of the Genetics Department. who directed this investigation. for their valuable advice and criticism. The writer is also indebted to W. V.
Lambert for his cordial cooperation and able aS$,iijtance . .

108
colors, were made reciprocally, and the hybrid and back cross
generations from them studied.
1.
2.
3.

Black Langshan x White Plymouth Rock.

Black Langshan x Buff Orpingtons.
Buff Orpington x White Plymouth Rock .

The parental matings were made by placing a male of each

breed in a 10 by 12 foot shed-roof colony house. Each of these
males were mated to several hens from each of the other two
breeds.
The hybrid F2 and back cross generations were secured in the
following manner. The F, males were mated with F, females
and in most instances the same males were back crossed to both
recessive and dominant purebred females. Back crosses to
pure-bred recessive males were made and in two of the matings,
back crosses to the Buff Orpington males w'ere used.
The same methods and materials were used in all of the different parts of the investigation, except that each phase involved color crosses in different ways. 'l'hese crosses are discussed under each of the divisions.
Where only two color classes were involved in the summaries, the closeness of fit of the actual data was calculated by determining the deviations and calculating the probable errors
by the use of the formula P. E. = .6745Yp X q X n. P and q
equal the elements of the theoretical ratio expressed in percentage and n is the number of individuals involved.
In this paper, if the deviation from the expected is three
times or less than the probable error, the hypothesis is considered possible; if the deviation is greater than three times the
probable error, the hypothesis is considered untenable. Where
three or more classes were involved, the fit was tested by thc
use of Chi 2 (X2) and from this the probability (P) of the closeness of fit of the data was obtained. Pearson's (28) tables were
used to obtain the probabilities from the various X2'S.

INHERITANCE OF' COLOR EXPRESSION

Before any adequate discussion can be entered into concerning the inheritance of the color genes, a brief summary of the
research in this field needs be made.
The heredity of white plumage color was first mentioned by
Darwin (6). He made a cross between a Black Spanish cock
and a white game hen in which he pointed to the possible existence of a dominant factor for white in the white games of
that period, but his results were inconclusive due to a very limited number of birds raised from this cross.
This work was followed by that of Bateson and Saunders (2) .
They used White Wyandottes and White Dorkings in their in-

109

vestigations, and crossed these breeds with the White Leghorn.

In this experiment, 549 F2 chicks with light down were obtained, and 176 with dark down. This approximated a 3:1
ratio, and they concluded that the white color of the Leghorn
was dominant over the other whites. They also stated that the
inheritance of white plumage color was complex.
Later, Bateson and Punnett (3) reported on some crosses they
made between 'White Leghorn hens and White Dorking males.
'l'he F, generations were mated "inter se" and they obtained
96 white and 6 colored chicks in the second hybrid generation.
They concluded that the six colored chicks appeared because
all of the White Dorkings used were heterozygous for the
dominant white color.
The same authors, Bateson and Punnett (4), demonstrated
that the white color factor of the Leghorn acts as a simple
dominant, and that there are two kinds of white, one dominant
and the other recessive. The recessive white was found to be
in White Wyandottes, White Dorkings and White Silkies.
'l'hese same investigators, Bateson and Punnett (5), went a
step further and found in their experiments that there were two
kinds of recessive whites. They made the following crosses in
their experiments.
1.
2.
3.

White Silkies x pure colored birds.

F, (White Silkie x pure colored birds) x F, male.
Silkie x Brown Leghorn).
White Silkies x Recessive Whites.

(White

From the first cross, they obtained 220 colored chicks. The
second mating yielded 156 dark colored chicks and 44 light
colored ones. 'l'he theoretical expectation on the basis of a
simple monohybrid recessive for color, i. e., a 3 :1 ratio, was 150
dark to 50 light chicks. The third cross produced all colored
chicks. They therefore concluded that these were produced because of the complementary action of two different types of
recessive whites. From the foregoing results and those of their
previous experiments, they grouped the whites into three
classes.
1.
2.
3.

Dominant White DDYYXX.

Silkie White ddYYxx.
Recessive White ddyyXX.

LEtter, Punnett (29) used the dominant II in place of DD because II would signify an inhibiting action on color. The three
classes of Whites appeared as follows:
1.
2.
3.

Dominant White IlXXYY (IlxxYY. or IlXXyy. or Ilxxyy).

Silkie White iixxYY.
Recessive White iiXXyy.

Following closely upon the work of Bateson and Punnett in

1908 was the report of Davenport (7) in which he showed conclusively the manner of inheritance of the Silkie 'White. He

110
TABLE 1. SUMMARY OF F, COL.O",R::.:.:...,'_-,--,;c;-_;--,-(D=-A~V--'--=,ENc:-=-PO=-R:.:T=-),--_
Parental crosses
I
Colored
I Non-colored I
Silkie (cc) x Buff Orpington (CC) ... I
Silkie (cc) x Black Minorca (CC) ... . I
Total observed .......... ........ 1
Total expected (3:1 basis) ....... 1

41
267
308
315

I
I
I
I

17
95
112
105

made two crosses and secured data on the F2 progeny in the

down stage. He used cc as expressing the white color of the
Silkie and its allelomorph CC as the expression of color. Table
I gives a summary of his results.
His results fit closely to the theory of a single gene difference
between Silkie 'W hite and colored birds, the factor for the expression of color being dominant. This is adequately indicated
by the relation of the deviation to its probable error, 7 6.0,
which shows a close fit between the actual and theoretical results.
Goodale (13) mentioned some experiments he conducted in
which he used the recessive white of the White Plymouth
Rocks. He crossed Brown Leghorns and White Rocks reciprocally and obtained the F, generation from them. In back crossing the F 1 females to a White Plymouth Rock male, he obtained 27 white and 26 barred chicks. This closely approximates the ratio 1 :1 which was expected if the White Rock was
a true recessive white. From a reciprocal back cross, 14 white
and 9 colored chicks were procUl'ed.
In some experiments of Hadley .(17), a White Leghorn male
was crossed with two White Plymouth Rock females. From
this mating, 63 chicks were hatched and the majority of them
raised to the age of 5 or 6 months. Five of these F, birds
showed barring in a few visible colored feathers. 'l'he following year an F 1 male was mated to four of his sisters. This mating gave 103 chicks of which 21 were colored. The next year
64 chicks were hatched, 52 of these being white and 12 dark
colored. He concluded from this that the White Leghorn male
bears the .genes IICC and the White Plymouth Rock females,
iicc. He also found that both of these breeds carried the barring factor (male BB, female B-) as a cryptomere and that
the barring factor was sex linked.
A summary of Hadley 's results showed that 134 white and
33 colored chicks were obtained in the F 2 generation. The
theoretical expectancy was 135.7 white and 31.3 colored, this
giving a deviation of 1.7 3.4 on a 13:3 basis
The investigations up to the present time have shown that
there are at least three types of white, one of them being due
to a dominant gene, II, which inhibits the production of pigment. Both of the others are r ecessive and have a complemen-

111
tary action with each other, producing color in the Fl generation.
Punnett (29) expresses these recessive types as xx and yy.
He states that one of the recessive whites, the White Silkie, is
of the type iixxYY and the other as iiXXyy. The latter is more
common than the former class of whites.
The use of these terms, xx and yy, is unfortunate as they are
now applied in general to the sex chromosomes in other species.
It would probably be better with respect to the recessive whites,
to use, the genes cc, the lack of chromogen, as indicated by
Hadley (16). Furthermore, this deficiency of color may be due
to an actual lack of chromogen material and that of the other
recessive white, the Silkie, might be represented as 00, a deficiency of an oxidase making possible the production of color.
Some chemical foundation exists for these latter terms and
the action that they have in producing color. Such evidence
has been advanced by Gortner (15). He found that the inhibiting factor, II, of the dominant white was due to the presence
of an inhibitory enzyme in the epithelial cells, which prevents
the action of the oxidase, and that the recessive whites differ
from the dominant types by having neither the power to produce pigments 00, lacking the chromogen material cc, or both,
ccoo; nor do they possess the inhibitor or anti-enzyme, II.
Therefore, to avoid possible confusion and perhaps to correspond with the chemical analyses, the following classification
and symbols might be used, grouped in the same way as that
made by Punnett (29) .
1.
2.
3.

Dominant White IICCOO (or IICCoo, or IIccOO, or Uccoo).

Silkie White iiCCoo.
Recessive White iiccOO.
II == Presence of inhibiting enzyme or anti-oxidase.
CC
00

== Presense of chromogen
== Presence of oxidase.

material.

EXPERIMENTAL

In this part of the experiment White Plymouth Rocks were

used, and from previous investigations they would be regarded
as recessive Whites (iiccOO) . They were crossed reciprocally
with Black Langshans (iiCCOO) and Buff Orpingtons
(iiCCOO). 'l'he hybrid and back cross generations, were also
secured, 'l'he breed crosses made were as follows:
1.
2.

White Plymouth Rock (cc) x Black Langshan (CC).

White Plymouth Rock (cc) x Buff Orpington (CC).

As the genes ii and 00 are common to all three breeds of

fowls, they may be omitted and only the factors CC and cc considered.
From the first cross, 69 colored F 1 chicks were hatched. In
the second mating, 167 colored progeny were obtained in the
first generation. In both of these crosses, all the observations

112
TABLE II.

SUMMARY OF THE DOWN OOLOR IN F2 AND BACK CROSSED

CHICKS, INVOLVLNG THE Cc FACTORS
Fz Down Observations
I

Parental crosses

1.
2.

3.
4.

Color

W. R. (cc) x B. o. (CC) ........... . ............ I

B. O. (CC) x W. R. (cc) .... ... . ... ....... .... 1
W. R. (cc) x B. L. (CC) . ................ ... .. . I
B. L. (CC) x W. R. (cc) ................ . .... . . 1
Total observed .......... . ..... ... ............ 1
Total expected (3:1 basis) ............. . ... ... 1

18
87
19
80
204
197.2

Noncolor
10
17
8
24
59
65.8

Back Crosses
5.
6.
7.
8.

W. R. (cc) X Fl (W. R. x B. 0.) .............. .

W. R. (cc) X Fl (W. R. x B. L.) ..... .. ....... .
Total observed .. .. .. . ............. ... .. ..... .
Total expected (1:1 basis) . ... .............. .
B . o. (CC) X Fl (W. R. x B. 0.)."' .......... ..
B. L . (CC) X Fl (W. R. x B. L.) ..... . ....... . .
Total observed ......... . . ...... ... . .. ......
Total expected (1:1 basis) .................. .

239
217
456
466
355
15
370
381

255
221
476
466
11
0
11
0

==
=

W. R.
White Plymouth Rock.
B. O.
Buff Orpington.
B. L.
Black Langshan.
These symbols will be used in all of the tables which follow.

of the down on the chicks showed them to be fully colored.

Table II gives the results of the observations on the down color
of the F z and back cross generations.
These data show that the down color is inherited in a simple
monohybrid fashion, the chromogen color, CC, being- dominant.
Both the Black Langshan and Buff Orpington breeds have the
CC factor, while the White Plymouth Rocks behave as recessive
whites, cc.
Most investigations on the inheritance of this gene have been
based on the color of down in the chick stage. Very few observations have been made as to the inheritance of this factor
(CC) in the adult plumage. From the crosses used, 70 adult
TABLE III.

SUMMARY OF COLOR INHERITANCE (Cc) IN THE MATURE

BIRDS
F 2 Generations of breed crosses

Parental crosses

B. L . (CC) x W. R. (cc) .... ........ ........ ..... . 1

B. o. (CC) x W. R. (cc) ............... . .......... 1
Total observed ................................ 1
Total expected (3:1 basis) .................... 1

Color
45
48
93
85 .5

Non-color
14
7
21
28.5

Back Crosses

(B.L. x W. R.) Fl '(Ce) x W. R. (cc) ........ . ....... 1

(B. o. x W. R.) F l (Cc) X W. R. (cc) .............. 1
Total observed .... .. ............ , ............ .. 1
Total expected (1:1 basis) ....... ..... ....... . 1
(B. L. x w. R.) Fl (Cc) x B. L. (CC) ..... . ........ I
(B. o. x W. R.) Fl (Cc) x B. O. (CC) . ... ....... ... I
Total observed .. ....... .................... .. 1
Total expected ( 1: 0 basis) ..................... 1
1

85
138
223
205
14
146
160
169

82
105
187
205

9
9

113
birds were secured in the Fl generation, all of which were colored. Table III gives the summary for the F2 and back cross
generations.
'l'he results of these crosses agree closely with those of the
expected or theoretical. The only departures that need explaining are the appearance of whites in the back cross to the dominant types and the deficiency of whites in the F2 and back cross
(recessive) generation. The former is probably due to errors
in marking the eggs at the time they were laid, as all three
crosses were in the same house, and it would be very easy to
make a slight mistake in reading the number of the hen's leg
band. The deficiency of whites, however, cannot be explained
except that it may be due to differential mortality. This is noticeable in all of the data given in the following pages. In spite
of these errors the data show a close fit between the actual and
the theoretical calculations.
Therefore, these results verify those of the many previous
investigators. The white of the ,Vhite Plymouth Rock is due
to a lack of chr9mogen material, cc, which acts as a simple recessive to its allelomorph, CC, the presence of chromogen material. Both the down and plumage color data appear to support this theory. The Cc genes are not sex linked and accordingly must reside on a pair of autosomes.

THE HEREDITY OF THE EXTENSION GENE FOR

BLAOK PIGMENT
The genetic analysis of the inheritance of white plumage
color is relatively a very simple matter in comparison with
black and buff colors, especially the latter. Very little definite
work has been done with the inheritance of black in poultry
until recently.
Hurst (20) crossed Black Hamburg females with a Buff
Co chin male. The first hybrid progeny were black with various
mixtures of brown. This showed that the black color is dominant altho not completely so. The results of the observations
on down color in the second hybrid generation were 88 black
(71 of which were variously marked with buff) and 31 buff
pigmented. 'l'his appears to be a case of a simple monohybrid
inheritance of black. But it is very difficult to base the inheritance of adult plumage pigments upon the down color of
the chicks. The writer has found, in his investigations, that
many of the black chicks develop into birds whose adult plumage would be classed as buffs, while chicks classed as buffs very
rarely develop adult plumage which would place them in the
black group.
In making studies of the Black Hamburg and White Faced

]14

Black Spanish fowls, Davenport (7) found that the blacks were
of different sorts, but the numbers used were small which made
the hypothesis inconclusive. Hadley (16) reported in his
studies with the ,Vhite Leghorn breed, the zygotic constitution
of the different black breeds of chickens used in some of his
experimental matings. In these investigations, he crossed several black breeds of poultry with the White Leghorn and found
that the blacks had a formula of bbCCii. The dominant CC
produces chromogen coloring ii indicates that there is no antioxidase present, and bb is used as representing nonbarring.
The first application of the gene EE for the extension of
black pigment was mentioned by Lippincott (21). In the case
of the Blue Andalusian, he used the zygotic formula for black
Andalusion as EErr, rr representing the non clumping (restriction) of the black pigment granules.
J..Jater I..Jippincott (22 ) went a little furth er in his investigations of the blue color in the Andalusians. In this publication he
stated thai the Blue Andalusians carried pigment and restriction factors but lacked the extension factor, EE. In one of
his crosses, h e used "White Plymouth Rocks and found that they
carried the gene EE for the extension of black pigment.
In his investigations of the heredity of black pigment in
domestic fowls, Dunn (8) reported that a gene Em was present
in black breeds of fowls. 'rhis factor acted practically in the
same manner as the gene E found by Lippincott, which was reported by him in 1918 and 1921. The factor Em causes the
extension of black pigment, and was found by Dunn (8) to be
present in Barred and White Plymouth Rocks, and absent in
Buff Orpington, Rhode Island Reds, and Columbian fowls.
The following year, 1923, Lippincott (23 ) confirmed Dunn's
findings of the extension factor EIIl. Lippincott used a cross
between a W hite Rock male and Light Brahma (columbian pattern ) females procuring the extended black pigment in both
the males and females of the first hybrid generation. He suggested that Buff Orpingtons, Rhode Island Reds and Lakenfelders carry the autosomal gene E and not Em while the
splashed blue Andalusian carried Em and not E. He further
stated that the factor E is carried by Black Langshans, White
Rocks, Whiie Wyandottes, and White Leghorns. 'l'his latter
statement agrees with that of Dunn (9), but they take directly opposite views of the gene carried by buff and r ed varieties. Dunn states that these varieties are of the recessive, nonextension type (ee) while Lippincott suggests that they are of
the dominant type (EE) .
Almost simultaneously with Lippincott 's r eport, Dunn (10 )
published his results on the genetic relationship of the black,

115
buff and columbian coloration in domestic fowls. He concluded
that black fowls are genetically buffs with the extension factor
superimposed. He used matings of columbian with black and
buff birds. He obtained 6 black females (Ees-) and 3 white duckwing males (EeSs) in the first generation when he crossed the
columbian females with black males. As a result of a back
cross to the columbian pattern fowls with an Fl male (black
female by a columbian male), he obtained silver duckwing
males (EeSS & EeSs ), columbian males, (eeSS & eeSs), and
females (eeS -), silver EeS - ) and buff birchen females
(Ees - ) and buff females (ees - ) . It will be noted that the
blacks of the first mentioned cross are of the formula Eesand that the same formula applies to buff birchens.
Ag'ar (1) verified the Elllfactor for the extension of black
pigment. He found that this gene was present in Barred Plymouth Rocks and absent in Rhode Island Reds, and Golden
Laced Wyandottes.
EXPERIMENT AL

In investigating this phase of the experiment, two crosses involving the genes EE and its recessive, ee, were used. The matings were made reciprocally and the proper hybrid and back cross
generations were secured. The following crosses were used.
1.

2.

Buff Orpington x Black Langshan.

White Plymouth Rock x Buff Orpington.

The first cross gave only extended, black, F 1 progeny, altho

they carried a slight amount of buff across the wings, back and
some in the neck. The results of the second hybrid and back
cross generations of this cross are placed in table IV.
The first hybrid generation males obtained from the second
cross were buff barred. Some of these carried a large amount
of black in the bars and had a slaty under color. The females
were predominantly buff, varying from a good buff color carrying a slight amount of black in the neck, primaries and tail, to
very dark birds which had black crescentic penciling across the
back and cushion.
The F 1 progeny of th e first cross showed buff to be recessive
which verifies Dunn's (8) hypothesis, but the hybrids from the
second cross showed buff to be dominant. This
be referred
to again in discussing buff plumage color.
The author believes it advisable to use the factor EE for the
extension gene. This is the same factor that was reported by
Lippincott (21). '{he writ.er can see no adequate reason for t.he
use of Em which was reported later and has t.he same meaning
and action. The use of this biliteral term with that of E makes
it difficult to correlate the literature in this field.

,,,ill

llG
TABLE IV.

ADULT PLUMAGE DATA ON THE INHERITANCE OF THE GENE

FOR THE EXTENSION OF MELANIC PIGMENT
F2 generations of breed crosses

Parental breed crosses

1.

2.

B. O. x B. L ......................... 1
"Total expected on 3 : 1 basis ....... . 1
Total expected on 33 :31 basis ..... . 1
W. R. x B. 0 ....................... . 1
"Total expected on 3:1 basis ........ 1
Total expected on 20 :44 bas es ...... 1

Extended
24
37.5
25.8
10
36
15

Non-extended

I
1
1

1
1
1

26
12.5
24.2
38
12
33

Difference

1
1
1

13.52.1
1.82.4

1
1

16+2.0
52.2

Back cross generations

3.

4.
5.

6.

(B. O.
"Total
Total
(B. O.
"Total
Total
(B. O.
*Total
Total
(B. O.
"Total
Total

x B. L.) Fl X B. 0 ... ........ 1

expected on 1: 1 basis..... . . 1
expected on 1:7 basis ..... . . I
x B. L.) Fl X B. L ............ !
expected ................. . . 1
expected .................... 1
x W. R.) Fl X B. 0 ....... .. .. 1
expected on 1: 1 basis ....... I
expected on 1: 15 bas is ....... 1
x W. R.) Fl X W. R. ........ . . I
expected on 1:0 basis ...... . . 1
expected on 7: 1 basis ....... .

12
47
11. 75
30
30
30
5
73
9.1
122
138
120.7

82
47
82.25

o
o
o

352.3
.25 2.2

141
73
136.9
16

68+3.9
4.12.0

17.2

1.25 1.9

Calculations made according to th e single gene hypothes is.

The results of the observation on the adult plumage from

the second hybrid and back cross generations, are placed in
table IV.
The first theoretical calculations are based on a simple 3:1
ratio as found by Dunn and Lippincott, and this gives a very
poor fit for the data at hand. The writer has found that there
is possibly a more adequate explanation for these poor fits which
are due to the interaction of other factors. 'l'he ratios given in
the second theoretical calculations in table IV give a good fit
for the present data and are based on the author's hypothesis
of buff color genes and the interaction between Ee and Cc. These
ratios cannot be explained adequately until the interaction of
these heterozygous genes and the inheritance of buff color are
discussed.
Altho more than 1,000 observations of down color were made,
it was impossible to obtain a simple classification of the various
grades. Many of the black graded chicks developed buff plumage. The percentage developing into buffs in each of these particular groups varied widely. Pearl (26) states on this same
subject, that in general the use of chicks in the down plumage
for the study of the inheritance of pattern and color characters
in poultry is open to very serious criticism. The reason for
this is that in many cases there is no definite or fixed relation
between color of the chick in the down and in the adult plumage.

117

THE INHERITANCE OF THE GENE FOR THE PRODUCTION OF BUFF PIGMENT

Altho breeds possessing buff pigment in the adult plumage
have been used by various investig'ators since 1905, very little
has been accomplished in explaining the heredity of the genes
responsible for this color, Some factors have been reported
which fitted the buff color for the particular crosses used, but
evidently do not fit all crosses, If the hypothesis is correct,
it should fit closely the data of all experiments,
Hurst (20) gives a brief account of a cross between Buff Cochin
females and a Mottled Houdan male and also of a cross between
Black Hamburg' females and a Buff Cochin male, He made
observations of the down in the second generation of the latter
cross only. Of 119 chicks obtained in the second hybrid generation, 88 were predominantly black while 31 were of various
kinds of buff. This close approximation of a 3:1 ratio suggests
a single factor difference with the factor for black being dominant. Hurst noted also that 17 of the F 2 birds developed into
full blacks while the others developed into a medley of various
mixtures of brown and black.
Davenport (7) records experiments conducted with Black
and Buff Cochin crosses. His observations were not made
beyond the chick stage in the F 2 generation and his results corresponded with those of Hurst. Neither of the above authors
attempted to place the inheritance of buff 'color on a factorial
basis. Altho several investigators used buff birds in back
crosses, no endeavor was made to analyze buff pigments, until
Dunn (8) published a series of investigations with the extension of melanic pigment and buff color.
Dunn (8) used crosses of columbian and buff fowls and he
concluded that only a single factor difference exists between these
birds. 'rhe columbian colored birds carry a sexlinked gene
tor silver, S, and the buff its allelomorph, s.
The same author (9) published a short article on his experiments with a gene for the extension of black pigment in domestic
fowls . He concluded that the black birds used had a zygotic
formula of E'm Em ss, the columbian em em SS. The extension
of black melanin is reported as due to an autosomal gene and
that of silver to a sex-linked one.
Later Dunn (10) r'e ported on this experiment in greater
detail. In his investigations, he used buff, columbian and blacl):
colored fowls, and concluded from back cross data that buff
and columbian colorations differ only in one principal gene de.t ermining the presence or absence of buff in the plumage, and

118

probably in multiple factors determining the amount of black

developed in the wing, tail and hackle feathers.
Both buff and columbian fowls lack the autosomal gene for the
extension of black. Dunn also adds that black fowls are genetically buff. The following formulae give the genotypic constitution of the different colors involved:
1.

2.
3.

Black Em Em ss, Em Em s-*

Columbian em em SS, em em S-*
Buff em em ss, em em s-*

This was the first attempt to place the buff color on a definite
factorial basis, but this color appears to be inherited in a more
complex manner than by a simple factor difference. Furthermore, there seems to be a definite interaction with the extension of black, the expression of color, and buff pigments, which
tends further to complicate investigational work.
EXPERIMENT AL

The author's experiments conducted at this station involved

two breed crosses. These were mated reciprocally and the
second hybrid and back cross generations secured. The following crosses were made:
1.

2.

Buff Orpington x Black Langshan.

Buff Orpington x White Plymouth Rock.

'fhe first cross gave only predominantly black individuals in

the first generation. The black males carried an intense buff
(red 1) in the wing bows, back, shoulders and a slight amount
in the hackle. This type of color and its pattern is analogous to
that of the Buff Duckwing males reported by Dunn in 1926.
The females were similar to his Buff Birchens. These were mostly
black with a goodly amount of buff mixed with the black, especially in the regions of the breast, back, cushions and wings.
The second cross, Buff Orpington with White Plymouth Rocks,
gave different results. All of the birds obtained in the F 1 generation were predominantly buff. The males were buff barred,
some had black undercolor and sprinkling of black on the buff
bars. The females were of two types, one carrying a small amount
of black in the neck, primaries and tail, while the other was crescentic penciled having a greater amount of black than the first
type. All of these birds, whether buff barred or buff with
crescentic penciling, were classed as buff.
In view of the diametrically opposite results of these two Fl
generations, some other explanation than a simple relationship
of the extension of black, EE, with buff, ee (1), must be made.
Previous investigators have shown that both the blacks and
recessive whites (White Plymouth Rocks) contain the factor for
The - indicates the non-carriage of S on the W (or Y) chromosome.

119
extension of melanic pigment EE (the author prefers the use
of EE rather than Em for reasons previously stated) the only
difference between the two colors, black and recessive white,
being that the White Plymouth Rock carries the barring factor
and lacks the chromogen genes, while the Black Langshan does
not have the sex-linked barring factor and carries the chromogen
determiner. The buff color, according to Dunn, is due merely to
a single gene difference and all buff birds are of the type ee. If
this hypothesis is correct, the breeds used in this study would
have the following formulae:
1. White Plymouth Rock EE BB cc 00 ii.
EE bb CC 00 ii.
2. Black Langshan
3. Buff Orpington
ee bb CC 00 ii.

As the barring factor will not influence the results of this particular phase of the work, it may be omitted and the buff or
black barred birds classed as buff or black, respectively. The
factors for oxidase, 00, and the non inhibitor, ii, are common
to all three breeds, and these may also be ignored. Thus it is
only necessary to study the effect of the two genes EE and CO
excepting', of course for any modifying or new factors.
In the first cross, CC is common to both breeds and the effect
produced on the first generation can be due only to the dominance
of E , which produced pr~dominantly black birds.
The second cross (Buff Orpington with White Plymouth Rock)
involved the same gene and also the one for the lack of chromogen
material. From this cross, only F 1 birds were secured that were
phenotypically buffs. The only conclusion that can possibly be
drawn is that a definite interaction exists between the heterozygous chromogen genes Cc and black E e, the heterozygous condition Cc weakening the effect of the black extension factor, and
thus permitting the buff color to appear.
The first hybrid generations would have the following colors
and genotypes in the different breed crosses.
1.
2.

Buff Orpington x Black Langshan = F , - Ee CC (Black).

Buff Orpington x White Plymouth Rocks = F , - Ee Cc
(Buff) .

A tabulated summary of the second hybrid and back cross

generations from these crosses is given in table V .
All the white birds are omitted from the classification because
the recessive cc factor covers up the genes carrying buff or black
color. Accordingly, the various theoretical ratios given in table
V will lack the white component of the phenotypic ratio.
It will be noticed that the closeness of fit is extremely poor
for the hypothesis that there is only a sing'le factor difference
between! black and buff (EE and ee respectively ) . This is
also true when the interaction between Ee and Cc is considered
altho to a lesser extent. Both of these hypotheses have been

120
""TABLE V.

SUMMARY OF ADULT BUFF PLUMAGE COLOR INHERITANCE

- F2 AND BACK CROSS GENERATIONS
F2 Generations
Parental breed crosses

1.
2.

I Black

Buff

Difference

------~I~c~~~~~~I~~~~

B. O. x B. L .............. . ... .. .... . . .. ..... . . 1

Total expected" (3:1 basis) . , .... . .. .. .... . 1
Total expected (33:31 basis) ......... . .... 1

B. O. x W. R. .. .. ....... ... ................... I

Total expected" (3:1 basis) ... ., ............ 1
Total expected' (including interaction of 1
Cc and Ee. 5:7 basis) .................. 1
Total expected (75:117 basis) ...... .... .. 1

24
37.5
25.8
12
36

26
12.5
24.2
36
12

20
18.75

28
29.25

8+2.3
1 6.752.3

12
47
11. 75
30
30
30
5
73

82
47
82 .25
0
0
0
141
73

I
1

13.52.1
1.82.4
242.0

Back Cross Generations

3.
4.

5.

6.

(B. O. x B. L.) F , x B. 0 ............. ... ..... 1

Total expected" (l:L basis) .. . ... .. . ... . .. . . 1
Total expected (1:7 basis) .............. . .. 1
(B. O. x B. L.) F, X B. L ........... ..... .. . .. I
Total expected' (1:0 basis) .... . .. ...... . . . . 1
Total expected (1:0 basis) .............. .. 1
(B. O. x w. R.) F, x B. 0 .. . ...... .... . .... . . 1
Total expected" (1:1 basis) ........ . ........ 1

Total expected. (including interaction of I

Cc and Ee. 1:3 basis) ...... . ........... 1
36.5
Total expected (1 :15 basis) ....... ........ ... 1
9.1
(B. O. x W . R.) F, x W. R. ........ .. .. .. .. ... \ 122
Total expected' (1:0 basis) .................. 1 138
Total expected, including interaction of I
Cc and Ee. 1:1 bas is) .. ................ 1 69
Total expected (7:1 basis) .................. 1 120.75

109.5
136 .9
16
0
69
17.25

35+3.1
. 252.2

683.9
31.53.5
4.12.0

53+4.0
1. 252. 6

** All white birds are omitted from classification.

*Calculations made according to single gene hypothesis.

tested by the data at hand and the results given in table V. In

addition to these, a third calculation of expected ratios derived
from a new hypothesis offered by the writer is given. In this
case, the calculated closeness of fit, for practically all of the F 2
and back cross data is good.
This new hypothesis suggests a certain interaction of factors Ee and Cc with two new pairs of genes for buff color. All
of the factors concerned are autosomal. 'l.'he writer has designated these new color factors for buff symbols Bu Bu Bu' Bu'.
The use of Bu (Bu Bu Bu' Bu' ) instead of B was adopted in order
to avoid confusion with the barring factor B which has the right
of priority. These buff genes are almost wholly recessive to
black in an invididual having only two doses of buff, and being
heterozygous for the extension of black pigment, and homozygous for the chromogen factor Cc (Bu bu Bu' bu' EeCC).
When the same combination occurs in an individual that is
heterozygous for Cc, buff color is developed (Bu bu Bu' bu' Ee
Cc). Either buff factor (Bu or Bu' ) gives the same effect as
the other. This situation is clearly portrayed in the F 1 genflrations of the following crosses used in this investigation.
1.

2.

Buff Orpington x Black Langshans = F , Ee Bu bu Bu' bu' CC

(Black) .
Buff Orpington x White Rock = F, Ee Bu bu Bu' bu' Cc
(Buff) .

121
These two different F 1 generations, arISIng from crosses in
which the buff breed is the common parent, and in which only
the 0 factor is variable, suggested very strongly that the homozygous dominant condition of 0 permits a different expression
of the buff-black relation than does the heterozygous condition.
This, together with the fact that a single factor difference between black and buff is untenable, led to the deduction that buff
color is more complex in nature than was previously noted. Accordingly the two pairs of buff factors were introduced, and
these, c(1upled with the interaction of Ee and Oc, were found
to explain the experimental results surprisingly well.
In the particular F 1 generations noted above ,it may be seen
that two dosages of buff make buff color hypostatic to black when
o is homozygous (dominant) ; but when 0 is heterozygous these
same two genes are permitted to develop the buff color. F2
and back cross data have given verification of this interaction
and have afforded the further assumptions; namely, that three
or four doses of the buff genes are epistatic to black when acting
with 00. From this, it also follows logically that when 0 is
heterozygous two, three or four doses of the buff factors will
produce buff color.
The interaction of these various genes are portrayed by
table VI, showing gametes and their resultant phenotypic
colors.
If the hypothesis advanced is correct it should fit Dunn's (10)
data. 'rabIes VII and VIn give a summary of Dunn's results
and also the calculated ratios for his and the author's hypothesis.
TABLE VI. SUMMARY OF GAMETES AND ADULT PHENOTYPES OF THE
BACK CROSSES IN THE WHITE PLYMOUTH ROCK BY BUFF
ORPINGTON CROSS'
x
Fl Gametes

Back Cross to W. R.
E bu bu' c
Back cross phenotypes

---------------------x Back Cross to B. O.

e Bu Bu' C
Back cross phenotypes

Black
Buff
Bu Bu' C
Buff
Black
bu Bu' C
Buff
Buff
Bu Bu' C
Buff
Black
bu Bu' C
Buff
Black
Bu bu' C
Black
Black
bu bu' C
Black
Buff
Bu bu' C
Black
Buff
bu bu' C
White
Buff
Bu bu' c
Buff
White
bu Bu' c
Buff
White
Bu bu' c
Buff
White
bu bu' c
Buff
White
Bu bu' c
White
Buff
bu bu' c
Buff
White
Bu bU' c
Buff
White
bu bu' c
Theoretical Ratios
0:1 :15
8:7 :1
F2
64 :75 :117
.The phenotypes are expressed only as buffs and blacks but in the actual observations made, the predominantly buff birds were placed in 14 grades ranging from a
pure buff to a blackish huff color. The blacks were placed in seven different classes.
ranging from a self black to the almost blackish buff of the buff class.

E
E
e
e
E
E
e
e
E
E
e
e
E
E
e
e

122
TABLE VII.

SUMMARY OF DUNN'S (1923) ADULT PLUMMAGE DATA TESTED

BY NEW HYPOTHESIS
Crosses of Buff and Columbian Fowls
Parents
1
-=-----,----=-:---:-:--Buff
Columbian
1

1.
2.
3.
4.
5.

P , Buff x Columbian .......... . ...... . 1

Total expected ................ . 1
Total expected ........ .. ... . .... 1
B. C. Fl Columbian x Buff ..... . ... . .. 1
Total expected .... . .... . ....... 1
Total expected . ................. 1
B. C. F, B. C. Buff x B. C. Buff ...... 1
Total expected .................. 1
Total expected ....... . ......... . 1
P , Columbian x Buff ....... ........ 1
Total expected ......... . .. ... . . 1
Total expected . ................. 1
B. C. Buff X F, Columbian ............ 1
Total expected . ................. 1
Total expected .................. 1

17
17
21
17
17
15
16
16
8
10
10
6

17
16
16

3
10.5
10.5

13
17
17

12
10
10
6
10.5
10.5

*Calculation based on single gene hypothesis.

From the foregoing tables, it is readily seen that the writer's

hypothesis adequately fits the data of Dunn. In Dunn's table
VIII, there were tWg classes with identical zygotic formulae
Ees- for self black and for buff birchens, while the author's
assumption gives each class a different genetic constitution,
Ees-bu bu bu' bu' for black and Ees-Bu bu Bu' bu' for buff
birchen. In table VIII the data of the first cross do
not fit the writer's hypothesis unless the F, t carried buff. This
male could be a Silver duck-wing, yet carry buff (Ee Ss Bu bu
Bu' bu'). The author is inclined to believe that such must
have been the case.
Accordingly, the results indicate that the extension of black
pigment (Ee and EE) in the presence of CC is epistatic to the
buff color factors when these are present in less than three
doses. Buff is epistatic to black when it is in excess of two
doses and is hyperstatic also when present in more than one
dosage, the chromogen and black pigment being heterozygous
(Ee Cc). From Dunn's data, it is evident that silver is epistatic to the buff color factors.

THE HEREDITY OF THE -F ACTOR FOR BA_RRING

IN DOMESTIC FOWLS
The barring factor, which accounts for the characteristic appearance of such breeds as the Barred Plymouth Rocks and
Dominiques, is one of the few color genes the inheritance of
which has been satisfactorily determined.
This gene was first mentioned by Davenport in 1906, who
noticed that it was closely associated with maleness but he
regarded it erroneously as a "heterozygous condition found in
hybrids from a white and black parent."

TABLE VIII.

SUMMARY OF DUNN'S (1923) ADULT PLUMAGE DATA T

Crosses of black and Columb'an fowls

Color pattern
Dunn's hypothesis

Ftd x Columbian 'j"s

Total expected
'Total expected
F t d x Buff 'j"s
Total expected
'Total expected
Columbian'j"s x Black rJ
Total expected
'Total expected
F, Black 'j"s x Buff d
Total expected
'Total expected

New hypothesis

Duckwing d
I
White
Buff
I
EeSs or I
Eess
EeSS
I
I
I
4
I
6.5
I
6.5
I
2
2
I
2.3
2.3
I
2.3
2.3
I
3
I
4.5
I
4.5
I
4
I

Black 'j'
Ees-

I
I
I
I
I
I
I
I
I

6
4.5
4.5

I
I
I
I
I
I
I
I
I

White

Buff

EeS-

Ees-

3
3.3
3.3
2
2.3
2.3

2.5
I
2.5
I
I
I
I
I
!
I
I
I
I
EeSs
EeSI
I
I
I
bububu'bu' I
EesI bububu'bu'l
Eess
I
I
or EeSs I BubuBu'bu'1 bububu'bu' I or EeSBubuBu'bu' 1
I BubuBu'bu' 1
I
I
I
I
I

.Calculations according to single gene hypothesis.

Birchen 'j'

3
3.3
3.3
2
2.3
2.3

I
I

I
I
I
I
I
I

I
I
I
I
I
I
2.5
I
2.5
I
I
I
EesI
BubuBu'bu'1
I

I B
I

124
Spillman (31) cleared up the inheritance of this factor, offering the explanation that the barring factor acted as a simple
dominant, and was sex-linked. Spillman's interpretation has
been amply verified by Goodale (12, 13, 14). In his investigations, Goodale used many different breed crosses and has
shown that the barring factor is dominant and sex-linked.
Numerous other investigators, Pearl and Surface (25), (26),
(27); Holde:fl.eisz, (19); Morgan and Goodale, (24); Hadley
(16), (17 ); Frateur (11) confirmed the observation of the previously mentioned authors. More recently attempts have been
made to determine the crossing-over percentage between another sex-linked gene, silver (S), and barring (B).
A few years ago Goodale (14 ) suggested linkage between
these factors. Haldane (18), Agar (1) and Serebrovsky (30)
calculated from their experiments the crossing-over percentage
between these genes and found values of 34.6 percent, 35.7 percent, and 435.6 percent, respectively.
From the accumulated data, it is well established that barring
is a simple, sex linked factor which is dominant over non-barring. 'l'his determiner may be regarded as a partial inhibitor
of pigment production, or better as a pattern gene. Normal
barred birds are genetically fully extended blacks or non-extended buffs which have added to them the pattern factor for
barring.
EXPERIMENT AL

In this experiment, no attempt was made to find the crossing

over percentages of the sex linked genes as the breeds used were
unsuitable for this purpose. The barring factor was included
as a minor investigation, and was used for the further confirmation of the previous investigations.
For this purpose, crosses between three different breeds were
used. The matings were made reciprocally and the second
hybrid and back cross generations secured. The following
parental matings were made reciprocally:
1.
2.

White Plymouth Rock x Black Langshan.

White Plymouth Rock x Buff Orpington.

The first hybrid generations obtained from the Black Langshan and Buff Orpington males mated to ,Vhite Plymouth Rock
females produced black or buff barred males and self black or
self buff females according to the particular breed used. The
reciprocal crosses gave black or buff barred males, black
barred females, and five non-barred buff females. The latter
were from the White Plymouth Rock male mated with Buff 01'pington females. The hatchability was good but due to heavy
mortality only five Fl birds were raised to 12 weeks of age, and
of these only one survived to be placed in an F2 mating, altho

125
adequate observations were made on two others before they
died. These females were non-barred and predominantly buff
with a few barred feathers in the region of the neck. The results of the second hybrid and back cross generations have been
summarized and are shown in table IX.
TABLE IX.

SUMMARY OF THE SEX-LINKED BARRING GENE"

F2 Generations
1

Parental breed crosses

1.
2.

3.
4.

Barred

Nonbarred

Deviation

----~I----~--'~~~--T_----------

w.

R. x B. L ........................ I
Total expected (1:1 bas is) ...... .. 1
B. L . x W . R ....................... 1
'rotal expected (3:1 basis) ........ 1
W. R. x B. O ....................... 1
Total expected (1: 1 bas is) ........ 1
B.O.xW.R. ....................... 1
Total expected (3:1 basis) ........ 1

2
4

25
27.2
20
20
5
6

6
4

2.0

.9

12
9.8
20
20
3
2

1.0

26
18

8.0

2.5

2.0

1.2

2.2 1.8
0.0 2.0
.8

Back Cross Generations

5.

W. R. X F, (W. R. x B. L . ) ... ..... /

Total expected (3:1 basis) ...... . .. 1
Barred F, x W . R. .................. I
Total expected (1:0 basis) ... ..... 1
7. B. L. X F, (W. R. x B. L .) ......... I
Total expected (1:1 basis) ....... . 1
8. (B. O. x w. R.) F, x B. 0 .......... 1
Total expected (0:1 basis) ........ 1
9. (B. O. x F, (W. R. x B. 0.) ......... 1
Total expected (1: 1 basis) ... ..... 1
10. W. R. X F, (W. R. x B. '0.) ........ 1
Total expected (3:1 basis) ........ 1
11. (W. R. x B. 0. ) x W. R. ............ I
Total expected (1:0 basis) ........ 1

6.

46
54
13
13
9
7

19
27
72
71.2
40

41

o
5

7
93
93
35
27
23
23.8
1

8 .0

2.4

0.8

2.8

White progeny are omitted from classification.

These data approximate closely those of previous investigators. The barring factor seems to affect the black and buff
birds in the same manner, being sex-linked and dominant, and
inherited as a simple sex-linked gene. Barring is designated
by practically all investigators by the letters BB in the male
and B-- in the female, while the non-barred birds are so distinguished by the allelomorphs bb and b--. In this experiment
the same symbols are used, and it was demonstrated that the
White Plymouth Rock carried the barring factor as a cryptomere (BBcc, B--cc), while the Black Langshan and Buff Orpingtons lacked this barring gene.

126

THE GENETIC PLU111AGE COLOR OPT WHITE PLYMOUTH ROCKS, BLACK LANGSHANS AND
BUFF ORPINGTONS
The color composition of the different breeds of poultry has
been neglected to a great extent. Many investigations have
been conducted on some of the factors concerned with the development of color in the different breeds, but few attempts
have been made to demonstrate in each breed the inheritance
of genetic determiners for color. Practically the only experiment to investigate the genetic color constitution of a particular breed, was that of Hadley (16 and 17).
From the foregoing analyses of the heredity (pages 107-126)
of the different factors used in this investigation, the genetic
constitution of White Plymouth Rocks, Black Langshans and
Buff Orpingtons can be described.
The autosomal genes 00 and ii are common to all three
breeds, and therefore are not used in this study altho they must
be included in the different breeds in order that the normal
color may develop.
From previous investigations, including the present experiment, it has been shown that CC probably represents a chromogen condition and acts as a simple mendelian dominant. This
factor is carried by the Black Langshan and Buff Orpington
but is lacking in the White Plymouth Rocks.
The extension of black pigment, EE, is a simple dominant
but has certain interaction, in a heterozygous state, with Cc
which destroys its balance, weakening its power of expression.
This determiner is found in the Black Langshan (probably all
black plumage birds bear this gene) and White Plymouth Rocks
evidently carry it as a cryptomere. Buff birds bear its allelomorph, non-extension of pigment, ee.
Buff plumage color seems to be inherited on a two factor
basis which the author designates as Bu Bu Bu' Bu'. The reason these symbols are used is to avoid confusion with the symbol B for barring. These buff factors are hypostatic to EE
when present in less than three of four dosages, and epistatic
when the individual bears more than two doses of the buff
genes. In addition to this, if an individual is heterozygous for
the chromogen determiner, Cc, then two or more doses of buff
are epistatic to black in the presence of this extension factor in
a heterozygous condition. The buff genes are carried by Buff
Orpingtons but are not present in the White Plymouth Rock or
the Black Langshan breeds. All of the color determiners mentioned thus far are autosomal in nature.
The next factor, barring, is sex-linked and this gene is present
in the White Plymouth Rocks as a cryptomere. The non-bar-

127
ring gene, b, is carried by the Black Langshans and Buff 01'pingtons.
From the inheritance of the genes already discussed it is
evident that the different breeds used must possess genetic
formulae as follows:
1.

2.
3.

White Plymouth Rock male EE BB bu bu bu' bu' cc 00 ii.

female EE B- bu bu bu' bu' cc 00 ii.
Black Langshan male EE bb bu bu bu' bu' CC 00 ii.
. female EE b- bu bu bu' bu' CC 00 ii.
Buff Orpington male ee bb Bu Bu Bu' Bu' CC 00 ii.
female ee b- Bu Bu Bu' Bu' CC 00 ii.
E == Extension of black plumage color.
B
Barring of either black or buff pigments.
Bu Bu' == Buff pigment genes.
e == Chromogen material present.
o Oxidase present.
I
Color inhibition (anti-enzyme or anti -oxidase).

=
=
==

EXPERIMENT AL

With these three different breeds the following matings were

made reciprocally and the hybrid and back cross generations
secured.
1.

2.
3.

White Plymouth Rock x Black Langshan.

White Plymouth Rock x Buff Orpington.
Buff Orpington x Black Langshan.

The F, generation obtained from the White Plymouth Rock

male crossed with Black Langshan females were all black
barred males and females. The reciprocal cross produced males
which were black barred and females that were self black. The
results of the F2 and back cross generations are placed in
table X.
Further evidence as to the genetic color of White Plymouth
Rocks was given by crossing this breed with Buff Orpingtons.
'l'he F, progeny were buff barred males and buff females, some
of which were crescentic penciled with a few barred feathers.
The results of further generations are shown in table X.
In order to procure additional evidence as to the inheritance
of both black and buff, a cross was made between the Black
Langshan and Buff Orpington. From this mating black males
with buff (red?) in the wing bows, wing, back and shoulders
(buff duckwing color) were obtained while the females were
black with buff mixture in the breast, back and cushion (buff
birchens). The results of the F 2 and back cross generations
from reciprocal matings are given in table X.
The relation of the different combinations in the hypothesis
for color are clearly expressed in tables VI (page 121) and XI.
Table XI includes the gametes, phenotypes and theoretical ratios for White Plymouth Rock x Black Langshan and Buff
Orpington x Black Langshan crosses.

128
TABLE X. SUMMARY OF COLOR INHERITANCE IN WHITE PLYMOUTH
ROCKS. BLACK LANGSHANS . AND BUFF ORPINGTONS
F 2 Generations
_ _ _ _ _-:-1~W
.:..:.:
h.:.:
it=e..;.i_.::B=u::ff::.,.!....:==_+--C:===~
1
1
W . R. x B. L ......................... I 14
1
Total expected (3:1 basis) ............ 1 14. 8 1
W . R. x B. O . . . .......... ... ..... . .. .. 1
7
1
36
Total expected (64:117:75 bas is) ...... 1 13 . 8 1 25.2
B. O. x B. L .......................... I
1 26
Total expected (31 :33 basis) .......... 1
1 24.2
Parental crosses

1.
2.
3.

1
1
Back Cross Generations

4.
5.
6.
7.
8.
9.

W . R. X F, (W. R. x B. L .) . . .... ... ... I 82

Total expected (1 :1 bas is) ..... .. ..... 1 83.5
B . L. X F, (W. R. x B. L.) ............ . . 1
Total expected (0:1 basis) ..... ....... 1
W . R. X F , (W. R. x B . 0.) .. . ... 1 105
Total expected (8:1 :7 basis) ..... . .... 1 121. 5
B. O. X F, (W. R. x B. 0) 1 9"
Total expected (15 :1 basis) ... ..... .. 1 0
B. O. X F , (B. L . x B. 0 ) ...... .... 1
Total expected (7 :1 basis) .... . ....... 1
B. L. X F, (B. L. x B. 0) .. .. .... 1
Total expected (0 :1 bas is) ............ 1

1
1

85
83.5
14
1 14.0
16
1 122
15. 2 1 106 .3
141
1 5
136.9 1 9.1
82
1 12
82.2 1 11. 8
0
1 30
0
1 30

1.5 4.2

P =

.10

4.12.0
0.2 2.2

'Buff and black barred birds classified as buff or black. respectively.

*These nine whites are probably due to experimental error and are omitted from
the calculations. Furthermore, crosses number two and six were in the same pens
as cross seven, which has the discrepancy of too many whites and no doubt they
should be classified in cross two or s ix, both of which are deficient in the number of
whites.

'l'he summaries in table VI and XI give the theoretical ratios

of the inheritance of the adult plumage colors according to the
different hypotheses already mentioned. When these ratios
TABLE XI. SUMMARY OF GAMETES AND ADULT PHENOTYPES OF THE
BACK CROSSES INCLUDING THE THEORETICAL RATIOS
- PI White Plymouth Rock x Black Langshan
FI
F,

Gametes

Barred Barred Barred

Barred Barred Black
White
White White
White White
White

EBC
EBC
E-C
Ebc
EBc
Ebc
E-c
Ebc
Theoretical
Ratios F. = 4:9:3

2:2 :0
-

x
Fl Gametes
E Bu Bu'
E bu Bu'
e Bu Bu'
e bu Bu '
E Bu bu'
E bu bu'
e Bu bu'
e bu bu'
Theoretical
Ratios F. = 31 :34

back cross to W. R. x back cross to B. L .

EBC : EBc and E-c
EbC and E-C
Barred
Black
Barred
Black

4:3 :1

0:4 :4

PI. Buff Orpington x Black Langshan Back cross to B. O.

e Bu Bu'
Buff
Buff
Buff
Buff
Buff
Black
Buff
Buff

7:1

Barred
Black
Barred
Black

Back cross to B. L.
E bu bu'
Black
Black
Black
Black
Black
Black
Black
0:8

129
are calculated for the summarized data in table X, it will be noticed that the assumptions made give a close fit to the actual results. Furthermore, this close approximation of the theoretical
with the actual results d emonstrates that the breeds are of a
genetic constitution as given at the beginning of this discussion.

130
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(1)

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(8)

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----

1909.

1922.

A Gene for the Extension of Black Pigment in Domestic

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1909.

Sex and Its Relation to the Barring Factor in Poultry.

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1910.

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1917.
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1913.

B.

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