Author Institution: Department of Dairy Science, Ohio Agricultural Experiment Station, Wooste

Baldwin, C. S.

Fechheimer, N. S.

Gilmore, L. O.

The Ohio Journal of Science

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No. 5 SOUTHERN ILLINOIS AQUATIC PLANTS 273INHERITANCE OF BLACK HAIR PATTERNS IN CATTLELACKING THE EXTENSION FACTOR FOR BLACK (E).IV. PARTITIONING PHENOTYPES BY CASTRATION2L. O. G1LMORE, N. S. FECHHEIMER, AND C. S. BALDWIN1Department of Dairy Science, Ohio Agricultural Experiment Station, WoosterThe appearance of blackish hairs in genetically red cattle (Jerseys andAyrshires) was shown by Baldwin et al. (1954) to be affected by genotype, sex,and age. They pointed out that the areas on which these hairs are found under'Present address, Central Ohio Breeding Association, Columbus.2Journal Article No. 57-61, The Ohio Agricultural Experiment Station.THE OHIO JOURNAL OF SCIENCE 61 (5): 273, September, 1961.274 L. O. GILMORE, N. S. FECHHEIMER AND C. S. BALDWIN Vol. 61different conditions are eye, cheek, muzzle, ear, neck, shoulder, body, rump, legs,and switch. They differentiated between blackish and black hairs, microscopically,and showed that the frequency with which blackish hairs were found was differentfor various parts of the body. It was concluded that the presence of blackishhairs on the head depended on the presence of the Bs gene. Those on the bodyand rump were concluded to depend on a pattern modifier, Bpm, which exhibitedsexual dimorphism in heterozygotes (Baldwin et al., 1956). A trait characterizedas exhibiting sexually dimorphic reversal of dominance is also known as sex-influenced.Pattern baldness in man has been shown to be caused by a sex-influencedgene (Snyder and Yingling, 1935). When Hamilton (1942, 1948) studied thischaracter in eunuch, eunuchoid, and castrate males, he was unable to detect theappearance of baldness. When, however, testosterone therapy was applied tofour castrated men, patterned baldness began to appear; when the therapy washalted the progression of the baldness ceased. Other sex-influenced traits reviewedby Baldwin are scurs and a certain type of horns (African) in both sheep and cattle.That Bpm is a modifier of Bs has been questioned by Overton et al. (1958)who analyzed the data of Baldwin (1951) by a method of maximum likelihood totest the validity of assuming that the variations observed could be explained alsoby the action of three alleles, i.e., Bsm, Bs and bs. Although they showed thatan allelic hypothesis was also tenable for the Jersey data, it was rejected at the5 per cent level of significance in the Ayrshire data.TABLE 1The effect of castration on the persistence of black hairpatterns in Jersey malesTreatmentBeforeAfterNo.Head166with Black Hair on Specified AreaCrest155Body140Leg1212Switch1212sEar2*2*Both had black fringe hairs.Hancock (1952) concluded from the observations on nearly 300 sets of identicaltwins that Bs has the pleiotropic effect of producing a strip of brown on thegenerally black skin extending from the lower jaw along the ventral line to theanalvulvar region and along the inside of the tail. There may be a break in thebrown skin under the neck. The inner surface of the ear as well as the lowerportion of the outer surface is also brown skinned. Hancock found that therefrequently occurred a characteristically shaped black pigmented area on thebrown skin inside the ear (ear streak) and on the ano-vulvar region (a-v spot).The bilateral symmetry of the whole of this skin color pattern was so marked andthe concordance between identical twins so great as to warrant the conclusion thatall variations except those resulting from conditions of housing and lactation mustbe explained genetically. In a further effort to explain different genetic causesfor the black hairs on red cattle, Hancock proposed the existence of a third gene, asimple recessive, be, which, in homozygotes, possessing also the Bs gene differsfrom Be/—Bs/—cattle by having only the skin of the udder, belly, brisket andthe upper two-thirds of the inner surface of the ears brown while the rest is black.No intratwin pair differences were found with respect to the phenotypes of be/beanimals. Because of the presumed interaction of be/be and Bs, the hair of theinner fringe of the ears is generally jet black.No. SEX-INFLUENCED INHERITANCE 275After the observation by Baldwin (1956) that an Ayrshire freemartin, fromparents both of which were nonblackish patterned, exhibited the pattern, theneed to observe experimentally imposed castration effects was suggested. Thiswas done on three Ayrshire bull calves. This experiment was concluded pre-maturely but, nevertheless, the blackish pattern on one of the bulls disappeared.The purpose of this paper is to report further pertinent observations that weremade on 17 Jersey males, some of which were castrated for another experiment(Hibbset al., 1959)." . """:FIGURE 1. The blackish pattern disappears with castration. A. (top) Jersey bull, 191 2 mo.old. B. (bottom) Same male 1 1 ^ mo. after castration, at 4014 mo. of age. Theblackish pattern had completely disappeared.EXPERIMENTAL PROCEDUREThirteen of the 17 calves were castrated at approximately six months of ageby crushing the external pudic nerves and arteries with an emasculatome. Beforecastration and again at approximately 17 months, color slides were taken of eachside of each animal. The other four bulls were castrated at an older age bygonadectomy, three at 11 months, and one at 23 months. These four wereobserved carefully as well as photographed at various times before and aftercastration.From each photograph, or animal, information regarding the presence of blackhairs on the head, ears, crest, body, legs, and switch was recorded.276 L. O. GILMORE, N. S. FECHHEIMER AND C. S. BALDWIN Vol. 61RESULTS AND DISCUSSIONFrom the observations summarized in table 1, it is seen that the six areasstudied can be divided into two separate categories, i.e., those areas which wereuniformly unchanged by castration and those areas affected at least in some ofthe cases. In no case was there a change in the gross nature or the extent of theblack hairs appearing on the tail and legs. From this it is concluded that thecolor expression of the black hairs in these areas are influenced by genes, theexpression of which is not affected by sex hormones. While the same resultwas observed on the black hairs of the ears as on the legs and switch, there wasan insufficient number of cases to justify a firm conclusion. Only two possessedthe black hair fringe on the ear described by Hancock (1952) as caused by theinteraction of Bs with the be/be genotype. In neither case, however, was a changerioted in the appearance of the black hair fringe following castration.From the observations made in this study, there is corroboration of the con-clusion by Baldwin (1951) that the black hairs in the switch and on the legs arepresent at birth. Furthermore, it has been shown that the pattern of black hairson the switch and distal part of the leg is not modified by brindling as are otherareas of the body on which blackish hairs are found (Baldwin et al., 1954).It has been observed that blackish hairs appear on the areas of the body otherthan legs and switch after approximately one month of age. The color expressionmay not be fully developed until 6 months or more in males and 2 to 3 years infemales. The blackish hairs on the body (including rump and thigh) disappearedin all 14 cases in which it was present before castration (table 1, fig. 1). Thus, itappears that blackish pattern in cattle responds to male hormone withdrawal inthe same manner as pattern baldness in man. For both head and crest, approxi-mately one-third of the experimental subjects failed to lose the blackish hairsin each area; three failed in both areas. So far the relation between Bs, Bpm,and be with different hormone levels has not been investigated with respect toeither of these areas; neither has the possibility of a differential response to castra-tion between the homozygote and heterozygote been excluded. It was impossiblein the present material to make a satisfactory diagnosis of genotype from thepedigrees and no definitive mat ings were made.SUMMARY AND CONCLUSIONSSeventeen Jersey bulls were castrated after they became old enough to showblackish hairs on the head, ears, crest, body, legs, and switch. Within one yearfrom the period of castration blackish hairs disappeared from the body, rumpand thigh areas in all 14 animals showing pre-castration blackish in these areas.This is interpreted to indicate absolute dependence of Bpm on the male hormonefor its expression. In 10 out of 16 cases the blackish hairs of the head disappeared,indicating either a partial reliance of Bs on the male sex hormone, incompletedominance or an interaction between hormonal function and the Bs alleles. Asimilar result for blackish or black hairs on the crest suggests the need for inves-tigating the cause of their appearance in this area. In no case did black hairsdisappear from the leg or switch. This is interpreted to indicate that these maybe black instead of blackish hairs and that a separate genetic cause is responsiblefor their appearance.REFERENCESBaldwin, C. S. 1951. Sex-Influenced Color Inheritance in Cattle. Master's Thesis. TheOhio State University., L. O. Gilmore, and N. S. Fechheimer. 1954. Inheritance of black hair patterns incattle lacking the extension factor for black (E). I. Blackish. J. Dairy Sci 37: 1368-1375., and D. C. Rife. 1956. Inheritance of black hair patterns in cattlelacking the extension factor for black (E). II. Blackish pattern. J. Dairy Sci. 39: 435-439.Hamilton, J. B. 1942. Male hormone stimulation is prerequisite and incitant in commonbaldness. Amer. J. Anat. 71: 451-480.No. 5 SEX-INFLUENCED INHERITANCE 277. 1948. The role of testicular secretions as indicated by the effects of castration inman and by pathological conditions and the short life span associated with maleness.Recent Ad. Hormone Res. 3: 252-322.Hancock, J. 1952. Identical cattle twins and colour inheritance. Endeavour 11: 78-86.Hibbs, J. W., E. W. Klosterman, and H. R. Conrad. 1959. Dairy beef production. Part 1.Performance of dairy steers from birth to slaughter. Ohio Agr. Expt. Sta. Bull. 833.Overton, Louise F., R. Laird, and N. S. Fechheimer. 1958. Inheritance of black hair patternsin cattle lacking the extension factor for black (E). III. A multiple allelic hypothesis toexplain the inheritance of blackish and blackish pattern. Ohio J. Sci. 58: 133-139.Snyder, L. H. and H. C. Yingling. 1935. The application of the gene-frequency method ofanalysis to sex-influenced factors with special reference to baldness. Human Biol. 7:608-615.

Inheritance of Black Hair Patterns in Cattle Lacking the Extension Factor for Black (E.). IV, Partitioning Phenotypes by Castration

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Inheritance of Black Hair Patterns in Cattle Lacking the Extension Factor for Black (E.). IV, Partitioning Phenotypes by Castration

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