Relationship Between Blood Type and Predicted Differences in Production of Holstein Sires in Artificial Insemination1, 2

Relationship Between Blood Type and Predicted Differences in Production of Holstein Sires in Artificial Insemination W. H. RAUSCH, E. W. BRUM, and T. M. LUDWICK

Department of Dairy Science, The Ohio State University; The Ohio Agricultural Research and Development Center, Columbus; DCRB, USDA, Beltsville, Maryland Abstract

I n an investigation which compared the blood types of 324 Jersey bulls with their USDA Predicted Differences (PD), Rausch et al. (16) found that bulls with A AI-I had progeny groups which were 0.06 unit lower in f a t percentage than their half-brothers that did not have the allele. Using Ohio NC-2 data, B a r r (2) found that cows heterozygous at the A locus ( A / D ) produced 210 kg more milk and 5 kg more f a t per lactation than cows which were homozygous (D/D). B-system. Because of the numerous alleles in this system, it is the most important locus for most cellular antigen studies, including parentage verification. I n a recent investigation which involved a large population of Ohio NC-2 Holstein cows, Brnm et al. (5) reported that the B alleles were associated with differences in fat percentage. F o u r different scientists have found an association between B BOxY2D' and superior f a t percentage. Andresen et al. (1) found a 0.07 unit superiority in f a t percentage with this allele. Rendel (17) found that animals with B BO~Y2D' tested higher (0.16 unit) in fat percentage than animals lacking this allele, l~eimann-Sorensen and Robertson (15) reported that cows with B BO~Y:D' were 0.06 unit above Although many investigations have shown contemporaries in f a t percentage. Conneally insignificant associations between blood poly- and Stone (6) found a superiority in fat permorphisms and production traits, this review centage (0.33 unit) with this allele. Two investigations, Andresen et al. (1) and is limited largely to those studies in which significant (P < .05) associations were found. Neimann-Sorensen and Robertson (15), found The systems included in this study are re- that allele B GD' was associated with high fat percentage and that B O~T~E~'K' was associated viewed by genetic system. A-system. Tolle (19) reported that cows with low fat percentage. Maijala (12) reported with A A~ produced more milk which had a lower that progeny of Finnish Ayrshire bulls with per cent f a t than cows lacking A A~. His later the allele B olA' exhibited a 0.02 unit increase (20) work verified the association between A A~ in fat percentage over progeny of their halfbrothers lacking the allele. Nair (14) reported and superior milk yield. an association between B BGIOxT-'A~'and low fat :Received for publication June 23, ]967. percentage in the Ohio NC-2 herds. This is ~A contribution from the 1~C-2 Dairy Cattle one of the few Holsteins in the United States Breeding Project in cooperation with Dairy Ge- where this allele is found. netics and Breeding Section, DCI~B, I~SDA, BeltsHogreve (11) suggested that the B System vil]e, Maryland. 2A project conducted in cooperation with the factor E ..," a factor present in many alleles, was Holstein-Friesian Association of America, Braft]e- linked with inferior lifetime performance. boro, Vermont. In the Jersey bull investigation, Rausch et al. The blood antigen types and estimates of transmitting ability for production (USDA Predicted Difference) were compared for 1,582 Holstein bulls. All blood-typed bulls which had Predicted Differences ( P D ) for 50 or more daughters listed in the U S D A Sire Summaries were included. These bulls had 804,195 daughters, averaging 508 daughters p e r bull. Analyses between eight blood polymorphic systems (A, B, F, J, L, M, S, and Z) and the PD estimates of breeding merit were conducted. In analyses within families of paternal half-brothers, both the B and L systems were related (P < .01) to PD for milk. The probability increased but was still less than .05 after adjustments were made for sire families and other blood systems. I n the B system, the constant for the BO1Y_~D' phenogroup was 305 ± 118 kg milk higher than the constant for the P 2 O~E~G 0 p phenogronp when the 20 most frequent phenogroups were considered. Milk production of females by bulls which did not have L exceeded the production of females which were sired by bulls having L by 55 ± 27 kg.

445

446

RAUS(]II, BF~UM, AND LUD~VICK

(16) found a relationship for the B alleles and milk and fat. However, when these data were analyzed within families of paternal halfbrothers these relationships were no longer significant. F-system. I n the same population of Holstein cattle mentioned previously, Bruin et al. (5) indicated that the F V / F v animals were highest for fat and milk yield, followed by F F / F F homozygotes, with F F / F v animals least desirable. J-syste,~. Rendel (17, 18) indicated that animals with cellular J produced milk with a higher fat percentage than those animals with no J. L-system. Rendel (17) reported that cows with L L produced milk which had a lower per cent fat than cows lacking L L. Treece et al. (21) reported that the absence of antigen L resulted in higher protein and casein percentages in the milk produced. These differences were of the magnitude of .14 to .17 percentage units. M-system. Mitseherlieh et ah (13), Tolle (19, 20), and Hogreve (11), working independently, all reported an association between ~[5~ and inferior milk production. Possibly supporting these results was Rendel's (17) finding' that cows with M M produced milk with a higher fat percentage than cows lacking

S-system. Tolle (19, 20) reported an association between S U-" and inferior milk yield. Hogreve (11) indicated that this same allele was associated with below average lifetime perfornmnee. Bruin et al. (5) found that the S alleles were implicated with differences in fat yield in the Ohio NC-2 Holstein herds. Rauseh et al. (16) demonstrated an association for the S alleles and milk and fat. However, these relationships were no longer significant in a within-family analysis. Z-system. Tolle (19) reported that cows with Zz produced less milk than cows that did not have this allele. Rausch et al. (16) found an association in a within-family analysis of the Jersey bulls between the Z system and diferences in milk and fat yield. I n most studies the blood type of individual cows was compared with their production, which was adjusted as much as possible for environmental influences. This investigation was designed to determine the relationship between discrete blood system and continuous milk production variables. One or possibly a few genes determine a blood system phenogroup and these blood polymorphic J. DAIRY SCIEI'~'CE VOL. 51, NO. 3

types are useful genetic markers (4). Variation in milk production results from the action of many genes and is greatly influenced by environmental variations. A bull's PD more accurately portrays his merit than the records of a cow portray her merit. With records expressed as 305-day 2 × ME deviations from herdmate averages, Van Vleek and Bradford (22) found a .50 correlation between cow breeding value and one lactation; .58 with two lactations; and .61 with three lactations, assuming equal heritabilities for all lactations. Bereskin and Lush (3) indicated that the expected correlations for various numbers of daughters in predicting the breeding value of a sire were .56, .69, .80, .86, .91, .93, .95, and 1.00 for 5, 10, 20, 30, 50, 70, 100, and an infinitely large number of daughters, respectively, assmning zero correlation among paternal sisters contributed by factors other than their sire. One would conclude that blood types of bulls provide an excellent discrete variable for this type of study and that the PD values provide a reliable estimate of the bulls' breeding values.

Experimental Procedure All dairy bulls used artificially by commercial organizations must be blood typed by a serology laboratory specified by the Purebred Dairy Cattle Association. Before 1954, blood typing was done in any one of three laboratories in the United States. Since 1954, typing of dairy bulls for permanent record has been done by the Serology Laboratory, University of California. The blood types used in this investigation were obtained from the files of the Holstein-Friesian Association of America for the bulls typed at California and the Laboratory of Genetics, University of Wisconsin, and from the files of the Cattle Blood Typing Laboratory, The Ohio State University, for the bulls typed at Ohio. Of the 1,582 bulls in this investigation, 1,131 were typed by California, 680 by Ohio, and 28 by Wisconsin. Five bulls were typed by all three laboratories, 235 by Ohio and California, 11 by Ohio and Wisconsin, and one by California and Wisconsin. The multiple laboratory typing provided a comparison for standardization of blood type codes. The blood types of the individual bulls were numerically coded according to Weseli's (23) coding system. Frequently a bull could not be coded for a particular system, either because he was not typed for that system or because he was an admixed twin. The relatively simple blood type loci, A, F, J, L, M, S, and Z were

447

BLOOD TYPE AND P R O D U C T I O N

coded directly from the results of the serological test. Four alleles were coded in the A system and the bulls' genotypes were coded whenever possible. All except nine bulls were coded in this system. A total of 1,561 bulls were coded for the codominant F system. This was the only system, of the eight investigated, where the bulls' genotypes were always known if the bulls were typed. I n the J system, 1,579 bulls were coded either for the presence or absence of the cellular J factor. At the L locus, 1,277 bulls were coded for the presence or absence of the L allele. Fewer bulls were typed for the M system than for any other, with 712 bulls coded either for the presence or absence of M. At the multiple allelic S locus, 1,526 bulls were coded for five phenogroups and genotyped when possible. The use of quantitative antisera made it possible to code the genotype of nmst bulls for the Z system of two alleles. Eighty-nine phenogroups were present in the B system. The individual laboratories phenogrouped this system from the serological reactions, generally with the aid of phenogroup frequency tables. The bull's blood type was compared with that of his sire when available to determine heterozygous animals in the simple systems and hidden phenogroups i n the nmltiple allelie systems. With this and the methods mentioned previously, 1,038 bulls were genotyped for the B system and 1,366 for the Z system. The hulls' most recent PD for pounds of milk and fat were obtained from four Dairy Herd Improvement Association Sire Summary Lists (7-10), including smnmaries from August 1963, through August, 1966. The PD values of bulls are the estimates of transmitting abili t y of artificial insemination bulls currently published by the USDA. The PD values for pounds of milk and fat were taken directly from the lists. The per cent fat values were calculated as follows, using 5,991.1 kg milk, 3.6455% fat, and 218.4 kg fat as the breed average : fat PD + fat breed avg % fat = -- breed avg milk PD + milk breed avg for % fat. All analyses, unless higher minimum numbers are indicated, included bulls with a minimum of 50 daughters in the PD. The data for analyses within fanfilies of paternal halfbrothers included 86 sire families with four or more sons. The F, J, L, M, and Z systems were coded as mentioned previously. I n addition to the phenogroup coding of the S system, the bulls were coded for the presence or ab-

sence of S. The A system was coded two additional ways: 1) the homozygous D bulls were coded with a 3 and all other blood typed bulls with a 0; 2) each phenogroup of an animal was classified independently, thus yielding results representing the additive effects of the alleles. The B system phenogroups of an animal were classified independently, for two different numbers of phenogroups, 1) the 20 most frequent, and 2) the 40 most frequent. I n the 40 phenogroup classification the least frequent phenogroups occurred in four or more bulls with one of the other 39 phenogroups. The analyses of the A and B systems using the phenogroup classification included only bulls with identifiable genotypes. Results and Discussion

Tiffs study included 1,582 blood typed Holstein bulls with 50 or nmre daughters with herdmates in the PD. These bulls had 804,195 daughters with records (an average of 508 daughters per bull). The PD'S were not reluted to the year of birth of bulls. The means of the PD's were --2S.8 kg milk, +0.01% fat, and --0.3 kg fat. Standard deviations for the three PD's were 228.0 kg, 0.10 percentage unit, and 8.0 kg, respectively. The phenotypie correlations among PD's were +.73, --.42, and +.31 for milk and fat, milk and fat per cent, and fat per cent and fat, respectively. Table 1 gives the mean squares from leastsquares analyses for the relationship of blood system polymorphisms and PD. Most of these stone analyses, adjusted for differences between sets of paternal brothers (sire families), are found in Table 2. This adjustment reduced the number of bulls included in the analyses generally by 59%. The significant mean squares in Table 1 that are not significant in Table 2 indicate a relationship between sh'e families and PD. Considering the constant estimates from the Table 2 analyses, in the B system the BO1Y~D' phenogroup was 304 ±_ 171 kg milk higher than the O~E/G'O' phenogroup, and B~GI was 10.7 ± 5.1 kg fat higher than OxE/G'O', when the largest difference among the 20 most frequent in the 40 phenogroup analysis was determined. Absence of J exceeded the presence of J by 38 ± 27 kg milk. Absence of L exceeded the presence of L by 59 --+ 32 kg milk and 1.7 ± 1.1 kg fat. Differences that approached significance favored the AH phenogroup for milk and fat in the A system, the absence of J for fat in the J system, and the absence of S for fat in the S system. Table 3 gives the mean squares from leastJ. DAIRY" SCI~+CE VOL. 51, NO. 3

448

RAUSCtt, BRUM, AND LUDWICK

TABLE 1 Mean squares from least-squares analyses for relationship of blood system and Predicted Difference Blood system

D.F.

Milk (kg)

A (D/D & - / - ) Residual A (genotype) Residual B (40 phenogroups) Residual B (20 phenogroups) Residual F Residual ff Residual L Residual M Residual S Residual Z Residual

] 1,571 3 ] ,095 39 931 19 794 2 1,558 1 1,577 1 1,275 1 710 1 1,524 2 1,363

62,748 51,971 38,250 51,847 99.765 ~ 51,807 185,547 ~ 49,261 108,914 52,182 183,340 51,957 339,642 ~ 53,822 5,753 59,712 2,022 52,425 32,051 53,628

Fat (%) 0.01 100.43 194.18 ] 00.17 214.45 ~" 104.34 273.97 ~ 103.12 234.43 100.44 1,862.07 ~ 99.31 57.83 106.03 475.27 ~ l J 2.70 93.79 101.25 530.04 ~ 103.90

F a t (leg) 87.76 63.30 3.87 64.84 ] J 8.] 8 ~ 63.75 180.06 ~ 62.34 33.65 63.94 98.04 63.39 289.85 ~ 66.65 112.22 72.45 16.34 64.02 49.48 66.38

Significant at the 0.05 probability level. ~ Significant at the 0.0l probability level. squares analyses of the r e l a t i o n s h i p s between blood system and P D a d j u s t e d f o r sire f a m i l y and other blood system effects. These analyses should eliminate the possible related effects of blood t y p e at other systems on the system u n d e r consideration, in addition to the possible sire f a m i l y effects. The mean squares r e m a i n e d significant a n d were very close to 0.01 level of significance f o r the B system a n d milk, when the effects o f the L system were removed. The mean squares f o r the J system were not significant when a d j u s t e d f o r B system effects.

This would suggest t h a t the significance of the difference in the J system, w h e n this system was analyzed individually or w i t h systems other t h a n B, was due to related B system effects. I n an analysis t h a t involved the B, J , and L systems, the mean squares were essentially the same as in the B and J, and B and L analyses given in Table 3. The B a n d L systems were significantly related with milk P D a f t e r adj u s t m e n t s were made f o r most e n v i r o n m e n t a l influences. The constants and s t a n d a r d errors f o r the

TABLE 2 Mean squares from least-squares analyses for relationship of blood system a~d Predicted Difference adjusted for sire effects Blood system

D.F.

Milk (kg)

Fat (%)

F a t (kg)

Residual B (40 phenogroups) Residual B (20 phenogroups) Residual F Residual J Residual L Residual M Residual S Residual Z Residual

3 456 39 357 19 307 2 632 1 641 1 521 1 287 1 622 2 558

89ff93 40,565 76,655 ~ 42,162 95,190 ~ 40_,580 19,045 41,229 174,332 ~ 4(),742 313,596 ~ 42,972 43,189 47,540 65,536 41,726 2,244 43,092

99.27 82.83 92.59 82.34 102.91 82.24 80.30 80.83 66.93 81.69 61.11 85.11 118.99 91.67 43.43 82.24 26.29 86.72

103.66 45.92 68.17 ~ 46.76 75.83 47.28 6.50 46.97 115.88 46.60 258.87 ~ 49.15 0.47 50.29 162.02 47.03 11.70 49.34

A

Significant at the 0.05 probability level. ~ Significant at the 0.01 probability level. J. DAIRY SCIENCE VOL. 51, NO. 3

BLOOD TYPE AND P~ODUCTION

449

TABLE 3 Mean squares fromleast-squares analyses for relationship of blood system and Predicted Difference adjusted for effects of sire families and some o~er blood systems Blood system

D.F.

Milk (~g)

Fat (%)

Fat (kg)

B (40 phenogroups) J Residual B (40 phenogroups) L Residual J L Residual L S Residual J L S Residual

39 1 356 39 1 324 1 1 520 1 1 515 1 1 1 514

76,762 +* 41,444 42,164 70:649" 177,496 + 42,600 200,113" 256,985 + 42,670 249±011" 56,962 43,170 199,969 + 197,542 + 58,642 42,865

91.28 0.26 82.57 93.52 146.84 84.45 103.48 44.10 85.07 65.16 44.55 85.20 70.02 49.84 43.68 17.54

69.39 + 58.90 46.73 66.16 69.09 47.89 114.72 219.12" 49.03 188.54 146.31 49.12 138.54 151.07 148.54 48.95

* Significant at the 0.05 probability level. +* Significant at the 0.01 probability level. B and L analysis in Table 3 are listed in Table 4. The phenogroups in the B system are listed by frequency of occurrence from most to least frequent. Table 5 gives the mean squares from leastsquares analyses of blood system and PD determined from a minimum of 100 or 200 daughters. These analyses were adjusted for sire family effects. These analyses compared the relationships between blood type and PD, using a minimum of 50 daughters in the PD with the relationships using a larger minimum number of daughters. The PD should more aecm'ately estimate breeding value as more daughters are included. However, in this study the number of bulls in the analyses decreased as more daughters were included in the PD. A direct comparison can be made with Table 2, which includes identical analyses using a minimum of 50 daughters. The mean square for fat became significant in the S system when 200 or nmre daughters were included in the PD. Associations between polymorphisms and PD may be due either to direct effects of po]ymorphisms or the chance occurrence of certain polymorphisms exhibited by bulls which for other reasons exhibit differences in PD. Certain analyses were completed within paternal half-sib families to remove from consideration differences in gene frequencies and PD's among these families. The extent of the association between polymorphic variations and differences in estimated breeding values among paternal half-sib groups was examined by comparing sire variance components adjusted for these polymo~hisms with the sire variance components from a one-way analysis. Polymor-

phisms of the B and L systems were utilized for these purposes, since these systems appeared most highly related with the PD values generally. Adjustment of the sire components for B and L system polymm~phisms did not alter the components for either milk or fat PD. l~owever, concerning fat percentage, the adjustment for polymorphisms resulted in a 27% reduction of the sire variance component. The most likely explanations for the apparent direct relationships between blood type and production as demonstrated in this study would be pleiotropie gene action or chromosomal linkage. I f the relationships are the result of pleitropy, they would probably occur whenever the gene is present in any breed of dairy cattle. I f the relationships are the result of linkage between blood type alleles and major genes that influence production, one would not necessarily expect these same relationships to be present in dairy cattle of different genetic origin and possibly not in subpopulations of a particular breed. In a closely linked situation, the relationships would probably behave sinfilarly to pleiotrooy and it would be difficult to distinguish one from the other. I t is possible that the relationships result from pleiotropic gene action at some loci and linkage at others.

Acknowledgments The authors are grateful to Dr. W. R. Harvey for his ~.dvice in statistical procedures and for the use of his least-squares program. We also express our appreciation to Drs. F. R. Allaire, H. C. Hines, and H. L. Barr for their assistance in the development of the data and we wish to J . ]DAII~Y SCIE~TCE VOL. 51, NO. 3

450

nAUSCH, BRUM, AND LUDWICK

TABLE 4 Constant estimates and s t a n d a r d errors f r o m least-squares analysis of the relationships between blood type and Predicted Difference a d j u s t e d f o r the effects of sire families and the other blood system Blood type class GY2Ed --

BO1 OxDtE~'F'G'O ' I2

08J'K'O' OxE:,' O~A_." BO_-Y~A~'EJG' B,_,GI I' E~' BO~¥oD' B_oGY~D' O~Y~E~'G' O~E,~'G' O~Y~Ao'Y' O~K' O~E~'G'O' EGKO~EdF'O' O~Y~A._.' BGKO~A'E:~'O' O~Y~EdG'Y' EdG'I' BOMB' O~Ad BI BGKOxY~A'O' O: O3Y2.:PK'O' P GO~Y,~ B~GO~Y2D'E~' PI' O~O' BO~E~' BO2AI'E3'G' B2G0~

O~¥2A~' O~QE~' L/ -/-

Milk ( k g ) 24.8 --120.4 --83.7 --21.9 --33.8 -91.9 --47.1 --32.6 32.1 --21.4 11.1 --75.8 114.8 --86.7 --39.2 59.2 5.2 86.0 --189.7 --39.3 24.7 --40.0 204.9 --159.1 --129.5 111.2 --88.4 114.8 213.2 --203.7 213.3 86.7 58.2 90.4 435.4 358.5 --85.5 --131.3 --197.5 83.7

_+ 28.7 _+ 35.1 ___ 40.0 _+ 42.6 _ 45.4 +_ 41.0 -4- 45.4 _+ 53.6 ± 49.2 ___ 65.8 _+ 51.9 +_ 58.1 +_ 76.1 _+ 73.8 _+ 101.6 _+ 57.8 ~ 98.4 _+ 144.7 _+ 89.4 ~ 86.9 _+ 90.5 _+ 108.0 ___ 84.2 _+ 125.7 _+ 122.9 +_ 163.2 _+ 156.3 _+ 156.8 _+ 105.9 ~ 158.8 _+ 149.6 _+ 125.8 _+ 171.1 _+ 116.1 ___ 291.1 -4- 214.2 +_ 151.9 _+ 131.1 _+ 214.1 _+ 153.9

--27.4 +- 13.4 27.4 _+ 13.4

Fat (%) B System (40 p h e n o g r o u p s ) --.007 _+ .013 .045 +_ .016 .023 ___ .018 .011 _+ .019 --.003 ___ .020 .001 +_ .018 .027 ± .020 --.003 + .024 .019 -4- .022 ,091 -4- .029 .o08 ± .023 .013 + .026 --.038 _+ .~34 .016 _+ .033 .026 _+ .045 --,011 + .026 .012 _+ .044 --.033 +_ .064 --.010 _+ .040 .002 +_ .039 --.030 ± .040 .046 _+ .048 .O00 _+ .037 .039 -4- .056 --.005 _+ .055 --.077 -4- .073 .041 _+ .070 --.098 _+ .070 - . 0 6 9 _+ .047 .081 _+ .071 .045 -4- .067 - . 0 0 4 +_ .056 .~30 _+ .076 --.00O _+ .052 --.220 -4- .130 .050 _+ .095 --.0O0 _+ .068 .008 _+ .058 --.095 -4- .095 .067 +_ .069 L System .008 _+ .006 --.008 ~ .006

thank the University Research Computing Center for use of their computers.

References (1) Andresen, E., H o j g a a r d , N., Jylling, Birthe, Larsen, B., Moller, F., Moustgaard, J., and Neimann-Sorensen, A. 1959. Bloodand Serum-Group I n v e s t i g a t i o n s on Cattle, Pig, and Dog in Denmark. Rept. ¥ I t h I n tern. Bloodgroup Congr. (Munich), p. 24. (2) Barr, H. L. 1960. The Association Between Extracellular F a c t o r s of E r y t h r o c y t e s and Several Measurable P e r f o r m a n c e Traits ixl D a i r y Cattle. Ph.D. dissertation, Ohio State University, Columbus. J

DAIRY SCIENCE VOL. 51, NO. 3

F a t (]cg) 0.54 + --1.78 + --1.69 +-0.O0 ± --1.43 +__ - 3 . 4 3 +_ --0.19 _+ --1.26 _+ 2.41 _+ 4.55 ~ 0.96 _+ --1.77 _+ 1.76 _+ --2.06 +_ 0.26 +_ 1.73 _+ 0.93 _+ 1.16 -4--7.74 +_ --1.50 -4--0.89 ± 1.31 _+ --7.41 +_ --3.54 ± --4.87 ~ --0.29 _+ --1.16 +_ --1.48 ___ 3.33 _+ --2.90 ± 10.74 + 2.90 _+ 4.27 _+ 3.46 _+ 1.99 ~ 15.81 +_ --2.89 _ --4.48 ± --12.60 ~ 7.25 ±

0.96 1.18 1.34 1.43 1.52 1.37 1.52 1.80 1.65 2.20 1,74 1.95 2.55 2.47 3.41 1.94 3.30 4.85 3.00 2.91 3.04 3.62 2.82 4.22 4.12 5.47 5.24 5.26 3.55 5.32 5.02 4.22 5.74 3.89 9.76 7.18 5.09 4.40 7.18 5.16

--.54 _+ .54 _+

.45 .45

(3) Bereskin, Ben, and Lush, J. L. 1965. Genetic and E n v i r o n m e n t a l F a c t o r s in Dairy Sire Evaluations. I I I . Influence of Environmental and Other :Extraneous Correlations A m o n g the Daughters. J. D a i r y Sei., 48: 356. (4) Brewbaker, J. L. 1964. Agricultural Genetics. Prentice-ttall, Inc., Englewood Cliffs, New Jersey. (5) Bruin, E. W., Rausch, W. H., Hines, H. C., and Ludwick, T. M. 1967. Association Between Milk and Blood P o l y m o r p h i s m Types and Lactation T r a i t s of Holstein Cattle. ( A b s t r a c t . ) J. D a i r y Sei., 50: 987. (6) Conneally, P. M., and Stone, W. H. 1965.

451

BLOOD TYPE AND PI%ODUCTION

TABLE 5 Mean squares from least-squares azmtyses for relationship of blood system and Predicted Difference with 100 or more daughters adjusted for sire effects Blood system

D.F.

Milk (kg)

J

1 522 1 415 1 511 2 446

100 or More daughters 220,698 ~ 92.43 40,083 84.44 320,199 ~ 25.14 41,668 87.30 18,710 109.04 41,095 84.74 12,547 39.85 42,764 90.71

137.23 45.67 324.69 ~ 47.24 119.19 45.90 37.44 48.74

1 378 1 296 1 370

200 or More daughters 276,898 ~ 197.52 39,330 87.01 306,972 ~ 1.22 41,532 91.58 111,946 80.33 40,381 87.66

124.90 46.73 389.49 ~ 46.67 298.59 ~ 46.80

Residual L Residual S Residual Z Residual J Residual L Residual S Residual

Fat (%)

F a t (kg)

Significant at the 0.05 probability level. ~ Significant at the 0.01 probability level.

(7)

(8) (9) (10) (11)

(12)

(13)

(14)

(15)

Association Between a Blood Group and Butterfat Production in Dairy Cattle. Nature (Lond.), 206: 115. D I I I A Sire Smmnary List, August, 1963November, 1965, ARS 44-172, USDA, March, 1966. D t I I A Sire Summary List, February, 1966, ARS 44-174, USDA, May, 1966. DI{IA Sire Summary List, May, 1966, ARS 44-179, USDA, July, 1966. D t I I A Sire Summary List, August, 1966, ARS 44-184, USDA, November, 1966. tIogreve, F. 1965. Lifetime Performance and Blood Group Factors in Cattle. Investigations on Black Pied Lowland Cows. Tier~rztl. Umsch., 20: 17. Ma~jala, Kalle. 1966. On the Possibility of Predicting the Success of a Bull's Daughters from His Blood Type. Ann. agr. fenn., 5: 65. Mitscherlich, E., Tolle, A., and Walter, E. 1959. Untersuchungen fiber das Bestehen yon Beziehungen zwischen Blutgruppenfaktorch und Milehleistung des Rindes. Tierz. Zficht. biol., 72: 289. Nair, P. G. 1957. Studies on Associations Between Cellular Antigens and Butterfat Percentage in Dairy Cattle. Dairy Sci. Abstr., 19: 369. Neimann-Sorensen, A., mud Robertson, A. 1961. The Association Between Blood Groups and Several Production Characteristics in Three Danish Cattle Breeds. Aeta Agr. Scand., 11: 163.

(16) Rausch, W. H., Brmn, E. W., and Ludwick, T . M . 1967. Unpublished data. Ohio State University, Columbus. (17) Rondel, J. 1959. A Study oll Rela,tionships Between Blood Group and Production Characters in Cattle. Rept. V I t h Intern. Bloodgroup Congr. (Munich), p. 8. (18) Rendel, J. 1961. Relationship Between Blood Groups and the F a t Percentage of the Milk in Cattle. Nature (Lond.), 189: 408. (19) Tolle, A. 1959. Principles and Experimental Results Pertaining to Relationships Between Blood Group Factors and Heifer Lactation. Rept. VIth Intern. Bloodgroup Congr. (Munich), p. 40. (20) Totle, A. 1963. Die Blutgruppen des Rindes. M. and H. Schaper, Hanover, Germany. (21) Treecc, J. M., Gilmore, L. O., Washburn, R. G., and Fechheimer, N. S. 1959. Evidence of Inherited Influences Affecting Casein Yield and Percentage as Measured by Relationships to Sire Groups and Cellular Antigens. J. Dairy Sci., 42: 922. (22) Van Vleck, L. D., and Bradford, G. :E. 1966. Genetic and Maternal Influence on the First Three Lactations of Holstein Cows. J. Dairy Sci., 49: 45. (23) Weseli, D. F. 1963. Blood Group System Phenogroups in Cattle. Mimeograph by the Department of Dairy Science, The Ohio State University, Co]umbus.

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