INHERITANCE OF BLACK AND RED COAT COLORS I N CATTLE*

M. H. CAMPBELL Agricultural College and Experiment Station, University of Illinois, Urbana, Illinois

Received April 18, 1924

TABLE OF CONTENTS PAGE

INTRODUCTION.. ................................................................. 419 Source of Material.. ...................... ....................... 420 Hypothesis. .................................................................... 421 Presentation and discussion of data ....................... 421 S"RY AND CONCLUSIONS.. ..................................................... 427 LITERATURE CITED.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427 APPENDur-Table 6. ............................................................ 428

INTRODUCTION

BARRINGTON and PEARSON (1906) found from a study of the inheritance of blue-gray coat color in cattle that some pure-bred Galloway cows dropped red or roan calves when mated with a white Shorthorn male. LLOYD-JONES and EWARD (1916) made a further study of inheritance of coat color in blue-gray cattle. The data for the latter study were obtained from experiments in which pure-bred and grade Galloways were mated with Shorthorns. From these matings there were obtained 78 Fl's of which 67 were black or blue-gray calves, while 11 were red or red-roan. Five of these blue-gray F1 females were mated with a blue-gray F1 male, producing 21 calves in the second generation. Of this number 15 showed the black pigment, while 6 showed the red. The authors explained these results by assuming that black, B , is dominant to red, b. The results obtained in the first generation were due to the fact that part of the females were homozygous for black, BB, while part were heterozygous, Bb. The Fl's mated together were Bb in composition and consequently were expected to produce 3 black-pigmented off spring to 1 red-pigmented. The actual result of 15 to 6 was close to this ratio. The authors conclude

* The GALTON AND MENDEL MEMORIAL FUND has met in part the cost of setting table 6.

GENETICS 9: 419 s 1924

420 M. H. CAMPBELL

that, “The nature of black and red pigment in cattle as an independent allelomorphic pair of characters is clearly indicated.’’

I n discussing the roan color of the Shorthorn cattle, DUCK (1923) writes: “The question logically arises, why will a cross between red and black produce no roans? The reason is apparent when it is considered that red and black are genetically, as well as practically, true allelomorphs, resulting in complete dominance of black in the heterozygote, which in turn will separate out in the second generation in the definite Mendelian ratio of three dominants to one recessive.”

KUIPER (1921) has found from a study of the breeding of Dutch Belted cattle that some of the calves are self-colored. He states that,

“In this case they are mostly coal black, but occasionally when both parents are heterozygous as regards black hair, plain red calves were produced.”

I n a study of the color of the animals, as recorded in the herd-book of the AMERICAN ABERDEEN-ANGUS BREEDER’S ASSOCIATION, COLE and JONES (1920) found that the number of red animals per each thousand recorded in a volume of the herd-register ranged from 8.14 in the third volume down to . l 6 in the later volumes. The decrease in percentage of red animals is explained partly by the fact that red males could not be registered after volume 3, and partly because the red animals were unpopu- lar. I n addition, they believe that red is gradually being eliminated from this breed. The authors state, “Both parents of a red calf are equally responsible for the departure from the desired color. Furthermore, from each parent there must be an unbroken line of individuals red or ‘carrying red,’ that is ‘masqueraders,’ back to the remote red animals included in the ancestry of the breed.”

TEMPLETON (1923) has recently called attention to a red calf which resulted from mating a pure-bred Angus male with a pure-bred Hereford cow. He explains this by assuming that the Angus was heterozygous for coat color.

It is the purpose of this paper to present additional experimental data concerning the inheritance of black and red coat colors in cattle.

SOURCE OF MATERIAL

A n experimental study of inheritance in dairy cattle was undertaken by T. J. BOWLKER of Framingham, Massachusetts, in 1911. For this purpose Mr. BOWLKER made reciprocal crosses of the Holstein-Friesian and Guernsey breeds. The work was started with 41 Holstein and 18 Guernsey females. The F1 females resulting from the cross were kept in

INHERITANCE OF BLACK AND RED IN CATTLE 42 1

the herd and were mated with F1 males that had been selected for breeding purposes.

After the death of Mr. BOWLKER in 1917, his family undertook the completion of the work under the supervision of Doctor W. E. CASTLE. In 1919 Mrs. BOWLKER decided to discontinue the work and the herd was purchased by the UNIVERSITY OF ILLINOIS. Since that time the work has been carried on under the direction of the Department of Dairy Husbandry. A preliminary report of the experiment from 1911 to 1919 has been made by CASTLE (1919).

HYPOTHESIS

The theory that black and red in cattle are an independent allelomorphic pair of characters and that black is dominant to red will be assumed as a basis for the expianation of the results obtained in this experiment. Upon this basis the black-and-white animals are homozygous for black, BB, or carry the factors for both black and red, making them heteroqgous, Bb. The red-and-white animals are homozygous for red, bb, regardless of the generation to which they belong. Therefore, it will not be necessary to indicate whether a red-and-white animal is pure-bred, F1 or FP.

PmSENTATION AND DISCUSSION O F DATA

Table 6 (appendix) shows all the matings made and animals born in the BOWLKER hybrid herd previous to November 8, 1922. It also gives the name or number, date of bifth, color and sex of each animal born, together with the name, number and color of the parents. The table is so arranged that the pedigree of any individual can be traced back to the registered ancestors.

TABLE 1

Summary of the reciprocal matings of pure-bred animals showing color of ojspring.

PARENTS I l OFFSPRING

Male 1 Female Black-and-white Red-and-white ""

Guernsey.. . . . . Holstein 88 7 Holstein. ..... .I Guernsey I 31 1 0 - -I l I Total. . . . . . . . . I I 119 1 7

A summary of the matings of purebred animals, together with the number and color of their offspring, is given in table l.' These data show that when the Holstein males were mated with the Guernsey females

There have been a few animals born for which the color hacnot been recorded. These =re mostly males, which were doubtless disposed of at birth.

GENETICS 9: S 1924

42 2 M. H. CAMPBELL

all of the offspring were black and white; that when Guernsey males were mated with Holstein females the ratio was 88 black-and-white and 7 red-and-white offspring.

Fifteen calves were produced by mating Holstein females with red-and- white F1 males, 4 by mating Guernsey females with red-and-white F1 males, 1 by mating a Holstein female with a black-and-white F1 male, and 1 by mating a Guernsey female with a black-and-white F1 male. The results of these matings are shown in table 2. It will be seen from this

TABLE 2

Summary of matings of pure-bred females with F1 males.

PARENT OFFSPRING

Male Red-and-white Black-and-white Female - - Red-and-white R.. . . . . .

1 0 Guernsey Black-and-white F1. . . . . 0 1 Holstein Black-and-white F1. . . . . 4 0 Guernsey Red-and-white FI.. . . . . . 1 14 Holstein

table that the Holstein females mated with red-and-white F1 males pro- duced 14 black-and-white calves to 1 red-and-white. When Guernsey females were mated with the same F1 male, 4 red-and-white calves were produced but no black-and-white ones. The Holstein cow produced a black-and-white calf when mated with a black-and-white F1 male, but the Guernsey cow produced a red-and-white calf when mated with the same male.

It is interesting to know that the 8 red-and-white calves produced by the Holsteins, as shown by tables 1 and 2, came from 3 cows. These three cows also produced 3 black-and-white calves. The matings of these 3 cows, together with the color of their offspring, are shown in table 3.

TABLE 3

Matings and offsw-ng of pure-bred Holsteins which produced red-and-white calves.

PED-AND-WHITE I PURE-BRED HOLSTEIN I OFFSPRING I

Male parent Female parent Black-and-

white

Langwater Moderator 14556. . . Hailey Segis Erie Colantha 172915.. . . . Hailey Dimple Boy 22309. . . . . Hailey Segis Erie Colantha 172915.. . . .

Hailey Segis Perfecta 188896.. . . . . . . . . Guernstein Sultan FI. . . . . . . . . Hailey Segis Perfecta 188896.. . . . . . . . . Hailey Dimple Boy 22309. . . . . Hailey Perfecta Clothilde 138099. . . . . . Guernstein Prince FI.. . . . . . . . . Hailey Perfecta Clothilde 138099. . . . . . Silverwood Brilliant 35980.. . . . HaiIey Perfecta Clothilde 138099. . . . . . Hailey Dimple Boy 22309. . . . .

Red-and- white

1 3 1 1 1 1 0

INHERITANCE OF BLACK AND RED IN CATTLE 423

If it is assumed that red is recessive to black, as has been previously pointed out, it would be necessary for the three Holstein cows, given in table 3, to be heterozygous for black. Accordingly, when mated with red-and-white bulls they would be expected to produce black-and-white and red-and-white calves in equal numbers. The actual result obtained was 3 : 8, respectively. The probable error for this result has been deter- mined according to the method suggested by WELDON (1901). This shows the deviation to be 2.5, the probable error & 1 .l 185, and deviation divided by probable error, 2.235. This shows that the result obtained is not very close to the expected numbers. However, because of the small numbers, this result should not be given much consideration.

The pedigrees of the three Holstein cows in question are presented on the following page in order that the relationship between them may be studied. Two of the cows, Hailey Segis Perfecta and Hailey Segis Erie Colantha, have the same sire, King Vernon Johanna Segis. The question arises as to whether these two cows inherited the factor for red from their sire or from their dams. Their sire, King Vernon Johanna Segis, was in the experimental herd and was used in 10 of the 31 matings with Guernsey cows which produced all black-and-white calves. Had he carried the factor for red, it would have been expected that half his calves would be black-and-white and half red-and-white, when he was mated with Guern- sey cows. Instead, all ten of his calves from such matings were black-and- white. According to the theory of probabilities, this result would not have been expected more than once in 1024 times had he carried the factor for red. On this basis, it seems safe to conclude that the bull was homozygous for black, and therefore could not have transmitted the factor for red to his daughters.

Since the two cows, Hailey Segis Perfecta and Hailey Segis Erie Colan- tha, did not inherit the factor for red from their sire they must have inherited it through their dams. This is quite evident in the case of the former, since it is known that her dam, Hailey Perfecta Clothilde, carried the factor for red because she produced three red-and-white calves out of four matings to red-and-white males. There is no direct evidence that Hailey Erie Colantha, the dam of Hailey Segis Erie Colantha, carried the factor for red; but from the data presented it seems safe to conclude that she was heterozygous.

It is impossible to determine which animals, in the pedigrees of Hailey Erie Colantha and Hailey Perfecta Clothilde, carried the factor for red. All of the registered animals in their pedigrees were black-and-white and did not produce, so far as is known, any red-and-white offspring. The GENETICS 9 : S 1924

M. H. CAMPBELL 424

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INHERITANCE OF BLACK AND RED IN CATTLE 425

factor for red can be accounted for only on the basis that it was carried by one animal in each generation of each pedigree back to the time when red- and-white animals appeared among the ancestors. This shows that the factor for red may be carried through many generations without the appearance of red hair. Likewise, this factor is doubtless carried by an unknown number of pure-bred Holsteins a t the present time.

TABLE 4

Number and color of animals obtained from F1 matings

COWR OE PARENTS ] OFFSPRING

Male Female 1 white 1 white Black-and- Red-and-

Red-and-white . . . . .

0 0 Red-and-white. . . . Black-and-white. . . . 8 21 Black-and-white.. . Black-and-white.. . . 3 0 Red-and-white. . . . Red-and-white . . . . .

31 33 Black-and-white.. .

DEV.

A summary of the F1 matings is given in table 4. This shows that when the red-and-white Fl males were mated with the black-and-white F1 females there were produced 64 offspring, of which 33 were black-and- white, and 31 were red-and-white. There were only 3 matings in which these same males were mated with red-and-white females, but all the offspring were red-and-white. When the black-and-white F1 male was mated with females of the same color the result was 21 black-and-white to 8 red-and-white offspring. There were no matings of the black-and-white male with red-and-white females.

In accordance with the postulated theory, the black-and-white F1 animals may be considered to be heterozygous for color, whereas the red-and-white animals may be considered to be homozygous for red. Consequently, when matings were made between the black-and-white and red-and-white an equal number of each color would be expected in the offspring. The result of 33 to 31 agrees very well with the expected numbers, as shown by the probable error in table 4.

The black-and-white Fl’s, when mated together, would be expected to produce black-and-white and red-and-white offspring in the ratio of 3 to 1, respectively. As shown by table 4 the’result 21 to 8, respectively, is very close to the expected ratio. Since the red-and-white animals are homozy- gous for the recessive red factor, they would produce only red-and-white offspring when mated together. This was the result obtained with three matings.

Only a few Fz females in the herd have freshened up to the present time. Some of these were mated with F1 males and some with pure-breds. A GPNERCS 9 : S 1924

426 M. H. CAMPBELL

few three-quarter-blood females2 also have been mated with F1 or pure- bred males. A summary of these matings, which a.re given in table 6 (appendix), is given in table 5. Since the animals represented by this table are not all of the same generation, the genetic composition of each group is indicated. The black-and-white three-quarter-bloods, Fl’s and Fz’s, with the possible exception of 724 (see note 3, table 6 , appendix) are

TABLE 5 Summary of malings of F2 and three-quarter-blaod females wilh F1 and pure-bred males.

MALE PARENTS I FEMALE PARENTS 1 OFFSPRING

Genetic Color

Genetic -white composition Color composition

Black-an

Red-and-white.. . . . . . . . . .

5 b b Red-and-white B B Black-and-white.. . . . . . . . . 6 B b Black-and-white B b Black-and-white. F1. . . . . . 5 b b Red-and-white B b Black-and-white F , . . . . . . . 3 B b Black-and-white* b b Red-and-white.. . . . . . . . . . 0 b b Red-and-white b b

Black-and-white. . . . . . . . . . B B Black-and-white B b 4 (Registered Holstein)

(Registered Holstein)

* It is possible that one of these Fa females, 724, is homozygous, BB.

Red-and- white

6 2 3 2 0

0

heterozygous, Bb. Although mated with red-and-white and heterozygous black-and-white females the two registered Holstein males have never sired a red-and-white calf, so they are considered homozygous, BB. The numbers obtained from these matings are small, but may well be considered.

When the red-and-white animals were mated together they produced only red-and-white offspring, as would be expected. The nekt two types of matings in the table may be considered together, for one parent is red- and-white, while the other is heterozygous. The results of these two types of matings are 8 black-and-white to 5 red-and-white, while an equality would be expected. As stated above, one of these F2 parents considered to be heterozygous may be homozygous for black. This may account for one of the black-and-white offspring which should not be included in the ratio. The mating together of black-and-white Fl’s, or heterozygous ani- mals, gave a result of 6: 2, respectively, which is a ratio of 3 to 1, as would be expected. The pure-bred Holsteins produced only black-and-white offspring when mated to both red-and-white and heterozygous black-and- white animals. This is to be expected with pure-bred Holsteins that are homozygous for black.

2 The animals which have one F1 parent and the other a pure-bred are referred to as three- quarter-bloods.

INHERITANCE OF BLACK AND RED IN CATTLE 42 7

The results obtained from this experiment bear out the theory that black is dominant to red and that the two pigments act as an independent allelomorphic pair of characters.

There is some variation in the intensity of black and red colors of the animals in the BOWLKER hybrid herd. Some are a jet black with white, while others are a brownish black or chocolate and white. The original records kept by Mr. BOWLKER report some of these animals as “seal color and white.’’ The red color varies from a very dark to a light red. The intensity of the black may be explained on the same basis as LLOYD- JONES and EVVARD (19 16) explain the variation in the black of the Galloways. In regard to this condition they state,

“This is produced by weakening or failure of the black in the tips of the hairs and its replacement by a rusty red pigment. . . : The rusty appear- ance so often observed in Galloways is not a criterion as to the purity of these animals for the black factor and is in fact quite unrelated geneti- cally to the condition known as red.”

It is hoped that a further study of this color may be made on the animals in the BOWLKER hybrid herd.

SUMMARY AND CONCLUSIONS

In 1911 T. J.BOWLKER started reciprocal crosses between Holsteins and Guernseys in order to study the inherited characteristics of dairy cattle. The experiment was later taken over by the UNIVERSITY OF ILLINOIS. The data obtained by this experiment furnish material for the study of the inheritance of black and red coat colors in cattle.

The matings which have been made in the experimental herd number

Three Holstein cows dropped red-and-white calves when mated with red-and-white males. This is evidence that some pure-bred Holstein cattle carry the factor for red.

The data presented conform to the proposition that black and red coat colors in cattle are an allelomorphic pair of characters, and that black is dominant to red.

The intensity of the color of the pigmented areas varies from a jet black to a brownish black or chocolate in the case of the black-and-white animals, and from a dark to a light red in the red-and-white animals.

291.

LITERATURE CITED

B ~ I N G T O N , A., and PEWON, K., 1906 On the inheritance of coat color in cattle. Biometrika

CASTLE, W. E., 1919 Inheritance of quantity and quality of milk production in dairy cattle. 4: 427464.

Proc. Nation. Acad. Sci. 5: 428434. GENETICS 9 : S 1924

428 M. H. CAMPBELL

COLE, L. J., and JONES, Sarah V. H., 1920 The occurrence of red calves in black breeds of

DUCE, R. W., 1923 Colors of Shorthorn cattle. Jour. Heredity 14: 65-74. KUIPER, K., 1921 Color inheritance in cattle. Jour. Heredity 12: 102-109. LLOYD-JONES, O., and EVVARJI, J. M,, 1916 Inheritance of color and horns in blue-gray cattle.

TEWLETON, G. S., 1923 Coat color of cross-bred steer, Quoman’s Perfection, not that ordinarily

WELDON, W. F. R., 1901 Mendel’s laws of alternative inheritance in peas. Biometrika 1:

cattle. Wisconsin Agric. Exp. Sta. Bull. 313. 36 pp.

Iowa Agric. Exp. Sta. Research Bull. 30. Pp. 67a-106a.

encountered in the Angus-Hereford cross. Jour. Heredity 14: 39-40.

228-254.

APPENDIX

I n order to save space in the following table the male is referred to by the initials. The names and numbers (when registered) which correspond to the initials given in the table are as follows:

Initial Name

L. M. = Langwater Moderator H. D. B. = Hailey Dimple Boy S. B. = Silverwood Brilliant A. V. F. P. = Archer’s Victor of Four Pine I. V. K. = Illini Victoria King I. M. K. Y. = Illini May King Yeksa K. V. J. S. = King Vernon Johanna Segis H. S. E. = Hailey Segis Elliston 0. C. H. = Ona Clothilde Hartog I. J. 0. = Illini Johanna Ona G. P. = Guernstein Prince G. S. = Guernstein Sultan G. D. = Guernstein Duke No. 3 = aNo. 3

Number Breed

14556 Guernsey 22309 Guernsey 35980 Guernsey 67661 Guernsey 68560 Guernsey 59565 Guernsey 59332 Holstein

170069 Holstein 226637 Holstein 349919 Holstein . . . . . . F1 ...... F1 ...... F1 ...... F1

INHERITANCE OF BLACK AND RED IN CATTLE 429

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