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Mic 60-4060

BAER, Harry L ionet THE ASSOCIATION BETWEEN CERTAIN EXTRACELLULAR FACTORS OF ERYTHROCYTES AND SEVERAL MEASURABLE PERFORMANCE TRAITS IN DAIRY CATTLE.

The Ohio State University, Ph. D ., 1960 Biology - Genetics

University Microfilms, Inc., Ann Arbor, Michigan TIE ASSOCIATION BETWEEN CERTAIN EXTRACELLULAR FACTORS OF ERYTHROCYTES AND SEVERAL MEASURABLE PERFORMANCE TRAITS IN DAIRY CATTLE

DISSERTATION

Presented. In Partial Fulfillment of the Requirements for the Degree Dootor of Philosophy in the Graduate Sohool of The Ohio State University

By

HARRY LIONEL BARR, B. S. in A gr., M. Sc.

######

The Ohio S ta te U n iv ersity 1960

Approved by

Soienoe ACKN0W1EDC3£BNT

It would bo very difficult to mention each person who has con­ tributed in some way to the completion of this study. Therefore. I w ill mention the few without whose help the work would have been severely handioapped.

I wish to express my appreciation to my adviser. Dr. Thomas

Ludwiok for first stimulating my interest in graduate study, and then for supplying me with the aoademio and personal guidance with whioh to carry it through.

I would likB to extend my thanks to Dr. Fordyce Ely, Chairman of the Department of Dairy Science, for permitting me the opportunity of pursuing graduate study while serving as a member of his Btaff.

My thanks also to the personnel of the NC-2 Breeding Project for making available their store of data, and speoial thanks to Don

Richardson and Dr. Herman Riokard who were instrumental in the planning of this study*

To Dr. Edwin Hess and Dr. Donald Weseli. I wish to express my

appreciation for their counsel and advice not only in this study but through my entire graduate career.

i TABLE OF CONTENTS

Page

INTRODUCTION ...... 1

REVIEW OF LITERATURE...... 4 Blood Groups of Cattle ...... 4 Blood Groups and Selection in Man ...... 9 Blood Groups and Selection in Chickens...... 12 Application of Cellular Antigen Knowledge to Selection in Other Types of Lrvestook ...... 15 Selection for Heterozygosity ...... 20

MATERIALS AND PROCEDURE...... 23 Animals ••»•••••• ...... 23 Antigens ..... 23 Variables ...... 26 A nalysis ...... 28 Analysis of Combinations of Heterozygous L o c i ...... 30

RESULTS AND DISCUSSION...... 31 A Loous ...... 31 F/V L o c u s ...... 54 J Loous ...... 56 L Loous ...... ••••»• ...... 58 SU Loous ...... 63 Z Loous ...... 63 Combined Looi ...... 66

SUMMARY AND CONCLUSIONS...... 69

BIBLIOGRAPHY...... 72

APPENDIX...... 82

AUTOBIOGRAPHY ...... 137

i i TEXT AND APPENDIX TABIES

Table Page

1 Form for Calculating Unbiased. Estimate of Difference • • 29

2 Preliminary Analysis of Varianoe Form ...... 29

3 Comparison of the Milk Production of Animals Hetero­ zygous and Homozygous a t the A Loous ••«•••••>• 32

4 Preliminary Analysis of Variance for Milk Production as Affected by the A Loous ....•••• ...... 34

5 Analysis of Varianoe Corrected for Disproportionality fo r M ilk Production as A ffeoted by the A L o o u s...... 36

6 Comparison of the Butterfat Production of Animals Heterozygous and Homozygous at the A Loous ••••••• 37

7 Comparison of the Maturity Indexss of Animals Hetero­ zygous and Homozygous a t the A Loous ...... 38

8 Comparison of the Heart Girth Measurements at Birth of Animals Heterozygous and Homozygous at the A Locus . . . 40

9 Comparison of the Body Condition Evaluations at Three Months of Age for Animals Heterozygous and Homozygous at the A Loous ...... 41

10 Comparison of the Increase in Heart Girths From Three to Six Months of Age in Animals Heterozygous and Homozygous at the A Loous ...... 43

11 Comparison of the Increase in Heart Girths From Three Months of Age to Three Months After First Calving in Animals Heterozygous and Homozygous at the A Loous • • • 44

12 Comparison of the Services Per Conception for Animals Heterozygous and Homozygous at the A Loous ...... 45

i i i iv

Table Page

IS Comparison of the Heart Girths of the First Offspring of Animals Heterozygous and Homozygous a t th e A Loous • * 46

14 Comparison o f th e M ilk Production of Animals Homozygous Dominant and Homozygous R ecessive a t th e A Loous • • . • 49

15 Comparison of the Milk Production of Animals Hetero­ zygous and Homozygous Dominant a t the A Loous •••••• 51

16 Summary of Unbiased Differences, for the Nine Variables, Between Animals Heterozygous and Homozygous a t the A Loous ...... * ...... 55

17 Summary of the Unbiased Differences, for the Nine V a ria b le s, Between Animals Heterozygous and Homozygous at the F/V Loous ••••••••»••••••••••• 57

18 Summary of th e Unbiased D iffe re n c e s, fo r th e Nine V a ria b le s, Between Animals Heterozygous and Homozygous at the J Loous • ••••••••••.•.*•• •••• 59

19 Summary of the Unbiased D iffe re n c e s, fo r th e Nine V a ria b le s, Between Animals Heterozygous and Homozygous at the L Loous •..••••••••• ...... • 61

20 Summary of the Unbiased Differences, for the Nine V a ria b le s, Between Animals Heterozygous and Homozygous at the SU Locus ...... 64

21 Summary of th e Unbiased D iffe re n c e s, fo r the Nine V a ria b les, Between Animals Heterozygous and Homozygous at the Z Locus 65

22 Summary of the Unbiased Differences, for the Nine Variables, Between Animals Heterozygous at Two or More Loci and Animals Heterozygous at Not More Than One Locus 67

23 Summary of A ll Comparisons Between Animals Heterozygous and Homozygous a t the S ix Loci • • • ...... 68

24 Comparison of the Milk Production of Animals Heterozygous and Homozygous a t th e F/V Loous • 83

25 Comparison of the Butterfat Production of Animals Heterozygous and Homozygous a t the F/V Locus 84 V

Table Pag©

26 Comparison of the Maturity Indexes of Animals Hetero­ zygous and Homozygous a t th e F/V Loous •*••••••• 85

27 Comparison of the He curt Girth Measurements at Birth of Animals Heterozygous and Homozygous at the F/V Loous • • 86

28 Comparison of the Body Condition Evaluations at Three Months of Age fo r Animals Heterozygous and Homozygous at the F/V Loous . • ...... 87

29 Comparison of the Increase in Heart Girths From Tnree to Six Months of Age in Animals Heterozygous and Homo­ zygous at the F/V L ocus ...... 88

30 Comparison of the Increase in Heart Girths From Three Months of Age to Three Months After F irst Calving in Animals Heterozygous and Homozygous a t the F/V Locus . • 89

31 Comparison of the Services Per Conception for Animals Heterozygous and Homozygous a t th e F/V L o o u s ...... 90

32 Comparison of the Heart Girths of the First Offspring of Animals Heterozygous and Homozygous a t the F/V Locus 91

33 Comparison of the Milk Production of Animals Hetero­ zygous and Homozygous a t the J L o c u s ...... 92

34 Comparison of the Butterfat Production of Animals Heterozygous and Homozygous a t th e J Loous •■«•••• 93

35 Comparison of the Maturity Indexes of Animals Hetero­ zygous and Homozygous a t the J Locus ...... 94

36 Comparison of the Heart Girth Measurements at Birth of Animals Heterozygous and Homozygous a t the J Locus . . . 95

37 Comparison of the Body Condition Evaluations at Three Months of Age of Animals Heterozygous and Homozygous a t the J Locus ...... • •••••.«•• 96

38 Comparison of the Increase in Heart Girths From Three to Six Months of Age in Animals Heterozygous and Homozygous at the J Loous 9 7 Vi

Table Page

39 Comparison of the Increase in Heart Girths From Three Months of Age to Three Months After First Calving in Animals Heterozygous and Homozygous a t the J Loous • . • 98

40 Comparison of the Services Bar Conception for Animals Heterozygous and Homozygous a t the J Loous .•••••• 99

41 Comparison of the Heart Girths of the First Offspring of Animals Heterozygous and Homozygous a t the J Loous • • • 100

42 Comparison of the Milk Produotion of Animals Hetero­ zygous and Homozygous a t the L Loous ...••••••• 101

43 Comparison of the Butterfat Produotion of Animals Heterozygous and Homozygous a t th e L Loous • •••••. 102

44 Comparison of the Maturity Indexes of Animals Hetero­ zygous and Homozygous a t th e L Locus 103

45 Comparison of the Heart Girth Measurements at Birth of Animals Heterozygous and Homozygous a t the L Loous • • • 104

46 Comparison of the Body Condition Evaluations at Three Months o f Age of Animals Heterozygous and Homozygous a t th e L Loous # v t «?« » v « o « • * . * * * 4 9 • • ^ « 105

47 Comparison of the Increase in Heart Girths From Three to Six Months of Age in Animals Heterozygous and Homo­ zygous at the L Loous • 106

48 Comparison of the Increase in Heart Girths From Three Months of Age to Three Months After Calving in Animals Heterozygous and Homozygous a t the L Loous • •••••• 107

49 Comparison of the Services Per Conception for Animals Heterozygous and Homozygous a t th e L Locus .••••.. 108

50 Comparison of the Heart Girths of the First Offspring of Animals Heterozygous and Homozygous a t th e L Locus . . . 109

51 Comparison of the Milk Produotion of Animals Hetero­ zygous and Homozygous a t the SU Loous 110 v i i

Table Page

52 Comparison of the Butterfat Produotion of Animals Hetero­ zygous and Homozygous a t the SU L o o u s ...... I l l

53 Comparison of the Maturity Indexes of Animals Hetero­ zygous and Homozygous a t th e SU Locus 112

54 Comparison of the Heart Girth Measurements at Birth of Animals Heterozygous and Homozygous a t th e SU Loous • • • 113

55 Comparison o f the Body C ondition E valuations a t Three Months of Age of Animals Heterozygous and Homozygous a t the SU Loous 114

56 Comparison of the Increase in Heart Girths Prom Three to S ix Months of Age in Animals Heterozygous and Homozygous at the SU Loous ...... 115

57 Comparison of the Increase in Heart Girths From Three Months of Ageto Three Months After Calving in Animals Heterozygous and Homozygous a t the SU Loous ••••••• 116

58 Comparison of the Services Per Conception for Animals Heterozygous and Homozygous a t the SU Loous •••••«• 117

59 Comparison of the Heart Girths of the First Offspring of Animals Heterozygous and Homozygous a t the SU Locus • • • 118

60 Comparison of the Milk Production of Animals Heterozygous and Homozygous a t the Z Locus •••••••••••••• 119

61 Comparison of the Butterfat Produotion of Animals Heterozygous and Homozygous a t the Z Locus •••••>• 120

62 Comparison of the Maturity Indexes of Animals Hetero­ zygous and Homozygous a t th e Z L o c u s ...... 121

63 Comparison of the Heart Girth Measurements at Birth of Animals Heterozygous and Homozygous a t th e Z Loous • • • 122

64 Comparison of the Body Condition Evaluations at Three Months of Age of Animals Heterozygous and Homozygous a t the Z Locus ...... 123 v i i i

Table Page

65 Comparison of the Increase in Heart Girths Prom Three to Six Months of Age in Animals Heterozygous and Homozygous at the Z Locus ...... 124

66 Comparison of the Inorease in Heart Girths Prom Three Months of Age to Three Months After Calving in Animals Heterozygous and Homozygous a t th e Z L o o u s ...... 125

67 Comparison of the Services fbr Conception for Animals Heterozygous and Homozygous a t the Z Locus • 126

68 Comparison of the Heart Girths of the First Offspring of Animals Heterozygous and Homozygous a t the Z Loous • . • 127

69 Comparison of the Milk Production of Animals Hetero­ zygous at ‘Two or More Loci ana Animals Heterozygous at Not More Than One Locus ...... 128

70 Comparison of the Butterfat Produotion of Animals Heterozygous at Two or More Loci and Animals Heterozygous at Not More Than One Locus ••••••.*•• ...... 129

71 Comparison of the Maturity Indexes of Animals Hetero- zygous at Two or More Looi and Animals Heterozygous at Not More Than One Loous ...... 130

72 Comparison of the Heart Girth Measurements at Birth of Animals Heterozygous at Two or More Looi and Animals Heterozygous a t Not More Than One Loous •••....* 131

73 Comparison of th e Body C ondition E valuations a t Throe Months of Age of Animals Heterozygous at Two or More Loci and Animals Heterozygous at Not More Than One Loous 132

74 Comparison of the Increase in Heart Girths From Three to S ix Months of Age in Animals H eterozygous a t Two or More Looi and Animal^ Heterozygous at Not More Than One Locus 133

75 Comparison of the Increase in Heart Girths From Three Months of Age to Three Months After Calving in Animals Heterozygous at Two or More Looi and Animals Hetero­ zygous at Not More Than One Locus 134 lx

Table Page

76 Comparison of the Services Per Conception for Animals Heterozygous at Two or More Looi and Animals Hetero­ zygous at Not More Than One Loous ••••••••••• 135

77 Comparison of the Heart Girths of the First Offspring of Animals Heterozygous a t Two or More Looi and Animals Heterozygous at More Than One Loous ...... 136 INTRODUCTION

The selection and breeding of cattle have occupied the minds of men for hundreds of years and the criteria by which animals were evaluated have been many. No doubt the original oriterion for selection was utility, but many things have orept into the selection procedures which have relegated u tility value to a position far from the top. The term 'purebred1 and the conformity to certain color patterns have come to command more respect than the usefulness of an animal in the minds of many of our influential oattlemen.

Although continuous efforts have been made by oattle breeders to select for characteristics which have a high u tility value, most of these traits have provided little information. The classic example of this was the belief in the significance of the 'escutcheon pattern' by an observer of livestock in France.

In this day of scientific approach to nearly every phase of agri­ c u ltu re , breeders of d a iry c a ttle s t i l l hold to many ideas of unknown origin in seleoting their animals, A few of these special points such as the 'mammary system' and 'd a ir y o h arao ter' have been shown to be correlated with the cow's main utility attribute, milk produotion.

However these indicator characteristics are expressed largely at an age when the animal is already in produotion, and any attempt to evaluate the produotion of a milking cow by such an indirect means seems about as ridiculous as guessing the weight of an object on the sc a le s by th e shadcrw i t oasts*

A more useful characteristic would be one that would help in the selection of dairy animals at an age long before their productive life had begun* Scientific exploration has been pursued in this direction and some modest adds have been developed, notably the measurement of the glandular development of immature females*

This present study involves investigation into another character­ istic of cattle which might give an early indication of the worthiness of an individual* The study of blood groups involves a characteristic in cattle which might give an early indication of an animal's eventual value* But even more important, an association between certain combi­ nations of blood antigens and usefulness would have an additional advantage. The inheritance of some various blood factors is simple enough and well enough known that we oould use them, not to evaluate the worth of an animal already in produotion nor to evaluate one that would be in produotion within a few years, but it oould be used to bring about the mating that would result in the proper antigenio pattern* Thus, not just superior animals would be seleoted from many, but rather only those of great promise would need be conceived*

The aforementioned ideas in conjunction with information concern­ ing association of blood groups with various traits, in species other 3 than cattle, has suggested the strong feasibility of suoh a study in dairy oat tie •

The high frequency that has apparently been established in man of

A and 0 of the ABO system with certain types of oanoer of the intestinal

tract, the reported association between ABO and other characters suoh

as elliptooytosis and nail pattela, and of course the well-known

relationship of erythroblastosis fetalis and the rhesus looi have all

added evidence to the selective advantage of certain blood factors.

In poultry the disoovery that the homozygosity at a given locus did

not fit those calculated by the coefficient of inbreeding and that at

least two alleles still segregated even after very intense inbreeding,

plus reports of greater growthjlivability and productive performance

among h e te ro zy g o tes, suggests th a t th e same m ight be tru e in o th er

types of livestock.

The objeotive of this study is to determine whether dairy cattle

heterozygous at any of several looi or at a combination of loci

perform differently in several measurable traits than do animals in

the same herd and by the same sire that are homozygous at these same

lo o i. HEYISW OF LITERATURE!

Reviewers of blood group work, Ferguson (42), Rendel (90), and

Wiener (122), give oredit for the first discovery of the within species differences in blood which would bring about agglutination to Land- steiner whan, in 1900, he made his fundamental discovery of the AB and

0 groups of human blood*

Todd and Tflhite (2 0 ), 1910, appear to be th e f i r s t to have d e te c te d

•the presence of an isolysin in cattle blood, end at this same time they discovered the necessity of foreign complement in the lytic reaction*

In the same work these authors noted the great individuality of cell characteristics and reported the sim ilarity that existed between oells or related individuals* Thus we had the first suggestion that these factors might be used in the study of heredity*

By 1911 Ottenberg and Friedman (82) had demonstrated isoaggluti­ nation when steers* blood was mined, and Little (65), using fresh bovine serum, also dem 'trated the presence of normal antibodies*

This work caused the authors to make a futile attempt to fit cattle blood groups into the pattern of the ABO of human blood*

Blood Groups of Cattle

The beginning of intensive study of blood groups in oattle was reported by Ferguson (41) when he discovered seven antigens which, he

suggested were inherited as if each was controlled by a different gone. Two years later Ferguson et al. (43) reported the existence of

30 antigens as detected by Isoimmune sera. Stormont et al. (116) reported 40 oharaoters in cattle, but discovered that some of the characters occurred in oompiexes rather than singly. Evidence of overlapping serologioal relationships was published by Stormont (ill) with his discovery of subtypes of four previously known factors.

In the development of the blood group thinking* the term pheno- groups need be introduced. Stormont (113) defines phenogroups as moleoules affeoted by individual genes. He defines blood factors as serologioal reactions of the various blood typing antibodies (reagents) that characterize phenogroups.

B loous. At present there appear to be 11 non-lintoed loci controlling the antigenio pattern of cattle erythrooytes. The great number of blood factors at the B loous has led to two different hy­ potheses of the inheritance mechanism. This difference of opinion is similar to the controversy that Wiener (122) describes in the human Rh system. Rendel (90) merely suggests that these groups of factors* instead of being controlled at one loous* might be a chromosome segment with closely linked looi similar to the Fisher hypothesis of th e Rh system re p o rte d by Wiener (122) in human Rh f a c to r s . Storm ont et al. (117) prefer to think of the system as one loous with multiple

alleles giving expression to the many phenogroups. The early recog­ nition of these blooks of factors was reported by Stormont et al.

(117)^(116)^(114). More reoently Stormont et al. (116) assert the 6 presence of 164 alleles in the B system although some are as rare in frequenoy as *005. Stormont (116J gives the same figure for the number of phenogroups which can be detected by the use of 40 blood factor reagents*

C loous* The C system is the second highest in number of possible alleles* Stormont et al* (117J had discovered 18 possible combinations at this loous and by a later report, Stormont (113) enumerates 35 phenogroups and 11 blood factor reagents*

Z* lo c u s. The Z 1 loous according to Stormont (113) has one reagent (Z1) and the phenogroups consists of the presence or the absenoe of the Z* factor*

M and D loci* M and D loci both appear to fit the same pattern as Z’ in the summation by Stormont (113). Stormont and Suzuki (118) discovered that the frequenoy of D was so high as to be of little use in cattle. An additional disadvantage is the fact that the antisera must be made by immunization of rabbits.

A loous. An early disoovery of a factor designated as A has been reported by Ferguson (41) and was further elaborated by Stormont (113) in which he found that three reagents were available (A, Ag and H).

From these, six phenogroups are known, and they are as follows* absenoe of any factor, Ai» Ag# H, A]H, and AgH*

F/V locus. The F/V system as reported by Stormont (112) has two reagents, and one or the other, or both factors are present in all animals* This provides for three distinguishable genotypes, F/F, F/V or V/V. Z loous. In the Z system, also reported by Stormont (112), one reagent is available which, gives rise to phenogroups Z and no Z.

However, Stormont (108) related that the Z/Z animals can be distinguish­ ed from the Z/-, since those cells with two units of the Z factor react much more rapidly than do the ho to ro zygote bearing oells.

L loous. The L system is one of the simpler systems according to

Stormont (113). One faotor is present and the presence or absenoe of

it makes up the phenogroups.

S/AJ loous. The SU system has four reagents, which gives rise to

si* different phenogroups. The reagents are S, H', Uj, with the

six phenogroups being SH1, U]H', SUgH*, U£» H*, and absenoe of any

faotor. Stone and Miller (105) discovered that U2 reagent was a normal occurring antibody and oould be used to deteot U 1U2 individuals

from previously undeterminable UjUi or UgUg individuals.

J loous. The development of this system has been quite interest­

ing. Stormont (109) reported the presence of normal antibody in the

plasma of cattle, and he also found this factor to be similar to A in

humans. Stormont ( 110) presented evidence that the J substance was a

soluble seriologioally reaotive faotor and that the red blood oells

acquired it from contact with the soluble substance in the plasma.

Neimann-Sorensen et a l. (81) showed that R faotor in sheep which is

a lso solu ble was very much lik e J in o a ttle and A fa o to r in human

blood. Other peouiarities of the J faotor have been noted by several

workers. Stone and Irwin (102) postulated the following three alleles

at this loous 1 Jcs existed where the faotor was on the oells and in the serum, Js where the faotor was in the serum only, and Ja where the faotor did not exist hut had potential for the normal antibody*

E lloit and Ferguson ( 40) gave the frequenoy of these three types, and showed that anti J oould also be stimulated by isoimmunization*

Stone and Irwin (103) reported that J is present in newborn oalves but not on their red blood oells* Hayaski et al* (49) designated J substance as a polysaooharide and confirmed that if it were expressed in serum it might or might not be on cells* Jamieson

(51) reported that the J antigen and J antibody were independent of the erythrooyte mosaicism that ooours often in other faotors in bovine twinning* l'hat the season of year affected anti J concentration was shown by Stone (101)* Bednekoff et al. (9) reported that a J substance was a glycoprotein with eleotrophoretio m otility.

Recently Sprague and Stormont (100) reported the presence in cattle of an additional soluble faotor which oould be detected by inhibition test employing so called cattle antisheep 0 antibodies*

This substance is called Oo and is sometimes absorbed by erythrooytes, but the cells are not agglutinated or hemolyzed by anti Oo* These

authors believe Oo to be a part of the J system, which, if it is,

gives phenogroups J, JOo, Oo and no faotor at this locus* Sprague

(99) indicated that anti Oo was not deteotahle when either or both J

and Oo substances were present} therefore, the anti Oc gene was regard­

ed as hypostatio* 9

Blood Groups and Selection in Man

Evidence of the selective advantage of certain blood groups has been the basis of studies in human populations for a number of years.

As early as 1921 Alexander (4) arrived at the conclusion, from his work, that all blood groups were subject to malignant disease, but two groups were more susceptible and that the clinical type of the disease was more malignant in these two groups. The large number of reports associating the ABO system with oarcinoraa of the digestive tract has led most reviewers to conclude that this correlation is real. The frequenoy of carcinoma of stomach has been shown to have a high degree of association with the occurrence of A, Buokwalter et a l.

( 3 0), Buokwalter ab al. (31), Aird (l), Aird et ajL. (3). Mayr et al.

(71) and Woolf (125) analyzed all available reports and concluded that

the A blood group was able to increase the risk of carcinoma by as muoh as 39 per c e n t. The work re p o rte d above was a l l accomplished using white populations, however Buokwalter et^ al. (29) found evidence that in Negro populations the A group also had a significantly higher

frequency of gastric cancer. All reports, however, have not been in

agreement with these associations. Clarke (36), Clarke et al. (37),

Mayo and Fergeson (70), Brown et al. (27) and Buchanan et al. (28) all

found little or no association between ABO systems and gastric

caroinoma. Allan (5) questioned whether the donors of blood which

were used as oontrols in these studies were reliable samples and also

pointed out the possibility of stratification and occupational effects

upon the study. 10

Associations between blood group and duodenal uloera have been reported frequently in the literature. Workers who have found suoh association are Clarke (36), Brown et al. (27), Buokwalter fi* (30),

Buokwalter et a l. (32), Buokwalter et al. (29), Aird efc a l. (2) and

A ird ( 1 ) .

Other association have also been reported between blood groups and oanoer type diseases. McConnell and Clarke (74) reported a high frequenoy of Rh individuals in another oanoer type and Roberts (92) found that A existed frequently in patients with oanoer of the panoreas and the esophagus. In a study of patients of five different hospitals,

Mayr et al. (71) associated pituitary adenomas with low A frequenoy and high 0 frequenoy. MacM&han and Falusiak ( 68) stu d ied 1,387 leukemia patients and found them to be low in 0 of the ABO system.

McConnell et al. (75) and McConnell (73) reported that an excess of A occurred in oases of diabetes m ellitus.

Bingham (ll) in combining data from a number of centers disoovered that pernicious anema was oommoner in persons of group A than in 0 or

B, and Maoleod (67) found a high frequenoy of B and of AB in the same condition. Renwick and Lawler (91) gave evidence of linkage between the ABO loous and a loous controlling dystrophy of the nails and absenoe of the patella.

Toxemia of pregnanoy is also a condition found to be more preva­

lent in women of 0 blood group aocording to Pike and Diokens (87).

Linkage association between the loous oausing an anomaly of the red blood cell oalled elliptooytosis and the ABO system was suggested 11 by the work of Burks et a l* (33)* Strong evidence of linkage between this elliptocytosis locus and the Hh blood group was presented by

Lawler and Sandler (61), Goodall et a l* (48) and Marshall eib al* (69)*

However, Fuzii et al* (45) and Chalmers and Lawler (35) found no

linkage between this loous and any blood group loous*

In considering the selective significance of blood groups in humans one also needs to consider incompatibilities and their effects

upon individuals. Levine (62) reviewed work of seven workers in whioh

a shortage of A offspring from 0 type mothers existed. Gershowitz

et al* (46) proposed the following two possible causes of the shortage

of A offspring from 0 mothers* (a) immune antibodies oould damage the

fetus and result in loss or (b) seleotion, during preconception phase,

on spermatozoa themselves could account for such a result. The

presence in cervical secretions of hemagglutinins found by these workers support gamete seleotion as a means of accounting for the

distortions of the expected ratios in marriages of A fathers and 0

mothers. Also increased abortion rates were shown to be a faotor in

incompatible marriages. By contrast Bennett and Brandt (10) found no

deviations from expeoted frequencies in incompatible matings*

Landsteiner and Wiener’s (60) first description of the Rh blood

faotor and their subsequent oase histories of erythroblastosis fetalis

has led to knowledge of a clinically significant type of selectivity*

Their work describes the loss of life amongst Rh positive offspring

bora to Rh negative mothers. Yannet (126) showed that in addition to 12 loss of life, victims of mental deficiency had mothers of which 25 per cent ware Rh negative as compared with 12 to 15 per cent Rh negative mothers in the general population* Wiener and Sonn (124) reported two oases in whioh erythroblastosis oocurred from sensitization by the ABO system. Shields et al* (96) reported sex ratio seleotivity when a high proportion of males were bora amongst AB children of A mothers, and there was scarcity of A males from B mothers*

Blood Groups and Selection in Chiokens

The work in ohiokens has been concerned largely with the advantage of individuals heterozygous for a given loous as compared with those homozygous* Also studies have been performed to determine the most desirable combination of multiple alleles at suoh a loous.

Shultz and Briles (97) reported that in their findings birds heterozygous at the A loous were superior for egg produotion, while heterozygosity at the B locus played a vital roll in reproductive fitness and heterozygosity at the D' locus was unimportant* Briles et al. (22) reported that male birds heterozygous for the B locus were significantly heavier at nine weeks of age in two of the three lines tested* The increase was from seven to ten per cent* The females' weight was also heavier for the heterozygotes, but not signifioantly so* Briles (17) studied the hatchability of eggs produoed by matingB resulting in 0 , 50 , 75 and 100 per cent heterozygosity at the B loous and found that these hatohahilities were 46, 62 , 71 and 70 per oent respectively* He calculated that the probability of a heterozygous embryo hatching was 1.7 times that of a homozygous embryo. He found 13 also that the heterozygotes excelled by 5 to 10 per oent in body weight and 9 to 30 per cent in egg produotion# Briles and Krueger (23) made matings between homozygous parents, matings between homozygous and heterozygous parents and between birds which were eaoh heterozygous for two alleles# The first hatohability sequence results were 48.6,

67.4 and 54.5 per oent respectively and these values proved to be significant at the one per cent level of probability. Within the second hatch however there was no significant difference. Chick livability was 9 to 15 per cent greater at nine weeks for the group of birds comprised of heterozygous individuals.

Briles (18) studied the B loous and adult performance and found that the hen day produotion for female heterozygotes for the B loous consistently outranked either of the two homozygotes, but he found no difference at the age at which the first egg was laid. Krueger et al.

(57) showed that the use of heterozygous males resulted in two per oent higher livability and five per cent greater egg produotion among their offspring.

Crosses were made between inbred lines, by Briles (19), in which the males were of B2B7 makeup and the females B7b7* The B2b7 male offspring outweighed the homozygous males by 2 .8 per oent at 10 weeks of age and the heterozygous females outweighed the homozygous ones by

4.8 per oent. In this work the author demonstrated that heterozygosity at the B locus exerted an effeot on growth between line crosses as well as within line crosses. 14

Briles (13) reported that male ohiofcs which lacked faotor M24 had significantly greater body weight at six weeks of age than did

those whioh possessed this factor* However he did not find this to be true in females*

Of interest in the breeding work involving ohiokens is the faot

that heterozygosity appears to persist in spite of high rates of in-

breeding. Briles and McGibbons (25) found heterozygosity at two loci

in spite of 52 to 60 per oent inbreeding, and in other lines, Briles

(16) found heterozygosity when inbreeding was between 47 and 50 per

oent. Briles et al. ( 20) found three to eight alleles still in

existence in 28 lines with inbreeding below 65 per cent. Tfith the 16

lines over 65 per cent the alleles hod been reduced to two. In this

work inbreeding had reduced the number of alleles as expected, yet, in

all oases two alleles remained. Further indication of this was given

by Gilmour (47) in whioh he studied seven apparently non-linked looi

and found that they were s till segregating following fourteen gener­

ations of brother and sister matings.

An interesting parallel between the work in chickens and a

oondition in human blood groups was revealed when Briles (15) was able

to induoe hemolytic disease in chicks by immunizing the mother with

blood containing 0 a n tig e n .

The problems of chicken and cattle may differ somewhat since

Scheinberg (95) states that there are 39 pair of ohromosomes in

chickens but probably only seven of any genetio use. Also two systems,

A with nine alleles and E with six alleles, appear, according to

• 16

Briles (14), to bo closely linked. However sim ilarities do exist between cattle and ohioken antigenic patterns. Briles et al. ( 20) have shown that the B system in chickens is similar to the B system in cattle in that both are made up of large number of blood faotors.

Briles et al. (26) hold to the concept of multiple alleles rather than linkage groups in this system.

Application of Cellular Antigen Knowledge to Selection in Other Types of Livestock

One of the early and continuing uses of blood faotors in cattle has been their use in settling questions involving parentage of off­ spring which are candidates for registration. This has been especially necessary with widespread use of artificial insemination and the increased ohanoe of a female being bred to two different bulls on successive heat periods. Miller and Stormont (76) report that 95 per cent of disputed parentage oan be solved by blood typing from the eleven systems used. They report that the extensive alleles in the B system are critioally involved in 50 per cent of the determinations, C system in 25 per cent, the FV, Z, SU, AH, L and J from 5 to 10 per cent each, the H* and Z% 2 per cent each with the D system being of little u se .

A oondition common in oattle bisexed twins is that of the free- martin as described by Lillie (64). This oondition may be difficult to diagnose in young animals. Owen (83) relates that blood antigens may be used aB an indirect aid in this problem by giving evidence that vascular anastomosis has ooourred. He states that when twins of both sexes are born, if the heifer has identioal blood type as her brother she is likely to be a free martin, but that the difference of one antigen indicates a normal female* Owen et al. (84) showed also that this anastomosis, which brings about erythrocyte mosaioism, ooourred when quintuplet calves involving four males and one female were born*

Owen (83) and Stone and Irwin (104) pointed out instances in whioh this mosaicism could lead to a false conclusion since animals who are twins may fail to transmit antigens whioh they themselves exhibit. Stone and

Palm (106) also related an instance in whioh a oow was shown to transmit to offspring antigens whioh she expressed weakly.

Blood typeB of cattle have also been used by researohers to make studies of relationships and patterns of development. Gene frequency studies have been carried out by Owen et^ al. (85) ( 86) and Neimann-

Sorensen (80), and the latter concludes that breeds with the greatest difference in the frequency of blood factors are those which are also the most divergent in genetic history.

Since Ferguson (42) suggested that there was a very real possibility that antigens of the blood will be found associated genetioally with level of milk production, attempts at correlating several quantitative faotors with blood factors have ooourred. Irwin

(50) pointed out that either the recognition of genes themselves or the recognition of heterozygosity oould make a useful contribution to the breeding superior animals. Rendel (89) and Robertson (93) also 17 discussed the possibility that an association exists between hetero­ zygosity and production traits*

Dunlop (39) made a study of the association between 11 antigens and three highly heritable type characteristics. Of the 33 comparisons, only the I antigen and fore udder score showed an association whioh was significant at the five per oent level, although three other comparisons approached significance* Nair et al. (79) studied various type defects in presence of several antigens and groups of antigens, and within families the presence or absenoe of certain antigen groups was associated with the frequency of occurrence of type defeots. The authors would make no generalizations on the basis of this work however.

McClure (72) realized that breeds which appear and perform similarily also have similar frequency of factors. Holstein-Friesian cattle which have low fat test also have a low frequency of antigen A while Jerseys and Guernseys have high fat test and a high frequency of antigen A. In this author’s study he compared the highest testing cows in several herds of Holstein cattle with the lowest testers in the same herd. The highest testers averaged 4.33 per cent butterfat and had an A frequency of 0.22 while the low cows tested 3.36 per cent and had an A frequency of 0.27. However the author found this difference to be not significant. Nair (78) also studied blood antigens and butterfat test among Holstein oattle and found consistent and signifi­ cant associations with faotors and groups of faotors in the B system.

Factors G, Y, C1, Ef, S* and groups BYC’, GYC’, GYE’ were observed in 18 the presenoe of high f a t t e s t "while fa c to rs B, Ofy, and 0, and group

G10T were associated "with low butterfat test. The associations outside the B system which proved to be statistically significant were believed by the author to be chance correlations. Morton et al. (77) investigated ten loci in cattle and found no evidence of linkage among any of them. In studying 5,000 animals they found no effect of the presenoe or absence of any blood factor on fertility . Also they found no significant difference between the theoretical and observed homozygosity of blood factors of 187 Hoi steins with an average inbreed­ ing ooeffioient of 25 per oent. There was however a 10 per cent deficiency in the proportion of calves with the blood factor from

’incompatible' matings of positive sires with negative dams as compared with the reciprocal 'compatible' crosses. There was no difference in the total number of oalves born in incompatible matings.

Laben and Stormont (58) in their work with the B, F/V, and Z systems found an excess of heterozygotes in animals with coefficients of inbreeding from 0 to .56. They found no difference in milk production between oows homozygous at none to all three loci. Twelve daughters of one bull heterozygous for one B allele produced signifi- oantly more fat oorreoted milk than nine daughters whioh were homo­ zygous. Reproductive efficiency also was favored in the heterozygous category, but other factors such as birth weight and disposal age appeared not to be affected.

Plum (88) reported on his work involving matings between animals which differed in various number of antigens. He found that with 19 parents differing in one to five antigens* 46 per cent conception ooourred* If they differed in six to seven antigens 49 per cent conoeption resulted and if they differed in eight to nine antigens 49 per cent conoeption again ooourred; however, if the parents differed in

10 to 15 antigens 60 per cent conoeption resulted* These results do not seom to readily agree with the loss due to factor incompatibility generally reported*

Treece et a l. (121) reported that the absence of antigen L resulted in both a higher protein and higher casein fraction in the milk produced* These differences were of the magnitude of *14 to .17 percentage units and were highly significant (P<*01).

Other significant studies on antigens and reproduction inolude the work of Laing and Blakemore (59)* They found in a herd that had a high abortion rate that six of 44 cows had sera which reacted against the cells of the bull to which they were in oalf • Also they found that animals transfused with blood from bulls to which they were bred would, after two to three transfusions, abort. The authors postulated that the antibodies might be acting at the cotyledon sites* Kiddy et ai. (54) failed to confirm these findings when they found those heifers which they injected with bull sperm failed to show any demonstrable antibody development and when these heifers were bred to the bull which produced the sperm they conceived equally as well as non- immunized oows* Bull's blood was also injected intrauterinely into 12 heifers without antibodies being demonstrable in secretions of the reproductive organs, but six of the heifers did show antibodies in the 20 sera. However there was no indication that this treatment reduced fertility. This also did not follow the pattern of earlier work by

Kiddy at al. (53) in whioh isoimmune sera in rabbits reduoed litter size. Braend (12) reported no passage of blood factor antibodies through the piaoenta in oattle*

The work in other livestock has not been extensive but a few reports have appeared in literature regarding the significance of certain antigenic patterns. The jaundice foal in horses which results from transplacental isoimmunization of the mare and then a transmission of the antibodies to the foal by way of colostrum is described by

Ievine ( 63) •

Buxton and Brooksbank (34) report haemolytio disease of newborn pigs caused by isoimmunization of pregnancy with jaundice developing within 48 hours. In these pigs parental blood incompatibility was demonstrated •

Of interest is the report of Stormont et al. (119) that the free- martin condition also exists in sheep. They report 30 cases of this

oondition and give an estimate of five per cent placental anastomosis

in sheep. As in oattle, mosaicism of the eryfchroeyte antigenic pattern

also occurs.

Selection for Heterozygosity

Inasmuch as this study is to deal primarily with the comparison

between homozygous and heterozygous individuals at few looi. it seems

advisable to look for a few classic examples of the selective advantage

of heterozygosity. The selective advantage and greater adaptability of 21 the heterozygote seems to be in evidence in several works# Schultz

and Briles (97 ) found what they called balanced polymorphism. At the

A loous the heterozygote proved to be superior for egg production, while balance at the B locus was vital in reproductive fitness# The

authors postulated that this balance at the B locus had developed during course of evolution by natural selection while A balance was of

more recent origin due to artificial selection for production. They

also pointed out that balance at both loci favored the overall

performance of individuals.

Probably the classic example of balanced polymorphism is described

in the studies of the siokle oell trait in negro populations. This is

a trait described by Stern (107) as a dominant allele which causes the

red blood oell to assume a siokle shape when placed in medium of

deficient oaqrgen. A small percentage of individuals so affected

develop serious pathological siokle-ooll anemia. Allison (6) in h is

study of iiast Afrioan populations discovered that tribes in hyper-

endemic malarial regions have over 10 per cent frequency of siokle

alleles while others are below 10 per cent# From this he inferred

that heterozygotes carrying the siokle allele had a high resistance to

malaria and this gave them a selective advantage in certain areas.

Allison (7) in a later study found that malarial areas had 20 per cent

frequenoy of the siokle-oell allele# In the same study he discovered

another allele in this system whioh he called hemoglobin C and he

claimed that heterozygotes of normal-sickle-oell or normal-hemoglobin 22

C were at a selective advantage in this area, but that the heterozygotes made up of the other two alleles were at a disadvantage. Livingstone

( 6 6) in a recent study has found a high frequency of hemoglobin C

associated with a low inoidenoe of the sickle-cell gene and reports a

low frequency of sickle-cell in some malarial areas.

Ford (44) in his discussion of human blood types points out that

different blood groups seem to be at a selective disadvantage yet

usually a balanoe is reached such that all blood groups are represented.

Also he pointed out that the balance reaohed in one population may be

different from another.

One additional fragment of information regarding the advantage of

two or more alleles segregating at different loci is contained in the

work of Clarke et alt (38) in whioh they report the protection given

against erythroblastosis fetalis by incompatibility between parents in

the ABO system. These authors state that an ABO inoompatibile fetus

gives complete protection against Rh sensitization and they give a

possible explanation of how this proteotion is conferred. Earlier in

this review we have seen a situation which would select heavily against

the Rh negative faotor and also a situation which would put A and B

factors at a disadvantage and favor a high frequency of 0, however

additional information here shows how the presence of A and B alleles

w ith th e 0 brings about inoompatibilites whioh leads to a selective

advantage in situations involving Rh difficulties. MATERIAL AMD PROCEDURE

Animals

The Bourca of information was the animals in institutional herds whioh belong to the Ohio Department of Mental Hygiene and Correotion.

These animals are all Holstein-Friesian.

The data used in this study has been reoorded routinely by the personnel of the Morth Central Regional Dairy Cattle Breeding Project,

II .0.-2, beginning in the year of 1940.

For the purpose of this study all sire groups under the super­ vision of N.C. - 2 program were used, providing the sire had at least 20 daughters available whioh had one or more normal production records and had been blood tested by the oellular antigen laboratory.

Antigens

The antigens were identified by their presenoe on the erythrooytes of oattle by the method described by Ferguson (41).

Blood was collected from the jugular vein into an isotonio sodium

oitrate solution. The reagents for testing cells were from sera prepared mostly by isoimmunization and suitably absorbed as described by Lazear and Ferguson (62).

The test consisted of adding .05 co. of a three par oent cell

suspension to tubes containing 0 . 1 cc. of the serum reagent for each

antigenic factor. The tubes were incubated at room temperature (22 to

23 24

29° C.) for 15 to 30 minutes* The oells were next exposed to the notion of complement by adding *05 oo* of pooled rabbit serum* The hemolytio reactions mere read at three different times during the next three and one-half hours. The interpretation of the presence of the factor in an animal mas used in this study as it was diagnosed by the trained personnel in the laboratory*

Five looi were studied and the classification at each was made by

dividing animals into homozygous and heterozygous groups. In order to

do this it was neoessary to study the antigenio pioture of the sire

and dam of the animal in most instances* The six looi* A, F7, J, L,

SU and Z, were ohosen because of the small number of alleles detectable*

This brought greater ohanoe of having homozygous individuals to

compare and also inoreased the ease of classification*

At the A loous a category could be determined readily which was made up of animals showing a complete absence of any faotor* The

second group made up of heterozygotes (A/-* A/H, AH/- and H/-J could

be plaoed in most instances* This was especially simple when either

the sire or dam was laoking in any faotor at the A loous* In other

instanoes where both the sire and dam had the same factors as the off­

spring, it was impossible to separate the heterozygous from homozygous

dominant animals and for this reason the third category was composed

of the homozygous dominants and some heterozygotes*

Because of the nature of the F/V system all animals oould be

classified in a definite oategory* There is no allele known for the 25 absence of any factor so that the two categories used were the heterozygotes of F/V and homozygotes made up of V/V and F/F.

The conditions at the J loous offers some basis for dispute over its usefulness. Only cells were tested for the presence of the J

substanoe, therefore an animal not appearing to be positive for J might well have this soluble substanoe in the serum. The group

appearing to lack J therefore would be made up of animals actually

lacking J, and in addition, those in whioh it had not been absorbed

onto the cells from the serum. For the aforementioned reasons the J

locus was used only in an individual study against the nine variables

and not used when the antigens were grouped and studied.

At the L locus a situation existed again where only two known

alleles segregate and thus the system oould be divided into the

homozygous reoessive or absence of any factor (-/"')» ‘fche determinable

heterozygotes (L /-), and homozygous dominant plus the undetermined

heterozygotes (L/L and L/-).

In the STJ system animals were olassified into two groups. One

group included animals whioh did not have any factor (-/-) and the

seoond group included the heterozygotes which were determined easily

(usually S/- because of low frequency of U faotors). A few oases of

S/S and S/- undetermined were also grouped. Because H1 had not been

tested for regularity and had not been originally considered as part

of this system, it was not used in the analysis. 26

The Z loous was categorized in the same manner as the L loous*

This was necessary sinoe the laboratory was not determining the Z/- individuals by the rapidity of the reaotion. This does not affect the aoouraoy of comparison between -/- and Z/- individuals as determined by studying the matings except that it forced an elimination of a few heterozygous individuals which might have been classified by

serological determinations.

V ariables

The variables that were studied in relation to the antigen

categories were milk production, butterfat production, maturity index,

birth weight, condition at three months of age, amount of gain between

three and six months of age, amount of gain between three months of

age and three months after first calving, services per animal at

seoond breeding and birthweight of first offspring*

The milk production (variable 1) records were taken from herd

improvement records sponsored by the Holstein breed association or

from records from the standard dairy herd improvement records

supervised by the Ohio State University. The records were considered

normal if the record was of 150 days or more and the cow did not abort

and had no extreme sickness which drastioally upset her production*

The first record of each animal was used as an indication of her level

of production and these records were all standardized to twice a day

milking for 305 days and computed to a mature equivalent using the

factors presented by Kendrioks (52). The butterfat records 27

(v aria b le 2) were obtained from the same source and the same adjust­ ments as were used for milk production were again applied*

To determine if a difference in maturity rate existed, an index

(variable 3) as developed by Barr et al. ( 8) and similar to the one presented by Krishin (56) wbb used* This index is derived by dividing the standardised (2x-305 MB) first record into the standardised second rec o rd . Numbers here -were lim ite d beoause i t v&b necessary to have animals with normal first and second laotations plus the further stipulation of a normal dry period (30 to 100 days) between the two laotations•

Birth weight was estimated by heart girth measurement (variable 4) at a time as soon after birth as the regular visits of the personnel of the NC-2 project to the herds would permit. The age of these calves could vary from one to 15 days.

Condition at three months (variable 5) was measured on a 10 point scale and was an evaluation of the amount of fleshing that the animal had acquired at this age.

The two gain measurements were the simple differences between the routine heart girth measurements normally taken at the ages designated.

The difference in gain between three and six months is designated v a ria b le 6 and the difference between the three-month measurement and the one taken three months after first oalving is designated variable

7.

Services at the second breeding (variable 8) were taken to indicate reproductive ability of these animals. 28

Due to the faot that male oalves are not routinely measured, the firat offspring, if it were a female, was used to get an indication of the possible effect of antigenic composition on size of offspring

(variable 9).

A nalysis

The analysis involved a comparison between the heterozygous individuals and the definitely determined homozygotes. The general form used to calculate the mean difference between the heterozygous and homozygous animals is presented in Table 1. The category of homozygous dominant and undetermined heterozygotes was not included in the analysis for two reasons* (l) the frequency distribution of

animals in this category varied greatly between sire groups; in some

sire groups this type did not exist while in others it predominated,

and ( 2) it appeared to be too mixed a category to hold much signifi­

cance regardless of its result. A mean value was computed however as

a matter of interest and comparison.

The analysis of variance as described and outlined by Snedecor

(98) for studies involving disproportionate subsamples was used. This

involves first testing for interaction and allowing the computation to

diverge depending upon the presenoe or absence of interaction as

indicated by this preliminary analysis of variance.

The form for this calculation procedure is shown in Table 2* 29

TAB IE I

Pom for Calculating Unbiased Estimate of Difference

(nixng; Heterozygous Homozygous (* 1 ~XZ) (ni-Hig) ni xi ng xg D W W

Sire group 1 group 2 group 3

group 22

Unbiased estimate of difference »

TABLE 2 -=

Preliminary Analysis of Variance Fom

Sum of Mean DP Squares Square

Source of Variance

Sub Class Means

Zygosity

Sire Groups

Individuals 30

Interaction sum of squares £vr lest for significanoe Interaction sum of squares individual of in te ra c tio n a degree of freedom + mean square

The F table was used to denote significance.

In comparisons where no interaction existed the unbiased estimate of zygosity difference in mean values was calculated by the formula

I m / l Y f .

This mean d ifferen ce was te s te d by the form ula F » ( 2 ® ^ +

in d iv id u al mean square.

In cases where interaction did exist, the procedure followed was

that desoribed by Snedecor (98).

Each of the six looi had to be tested against each of the nine

variables so the presenoe or absence of interaction oould be determined.

Analysis of Combinations of Heterozygous Loci

In this part of the study each variable was studied to see if a

difference existed between the two following categoriest animals

heterozygous at not more than one and animals heterozygous at two or

more of the five loci studied (J loous was omitted). RESULTS ACID DISCUSSION

A Loous

A comparison involving the first variable, milk produotion levels, between animals known to be lacking the blood faotor A {-/-) and those oarrying A in the heterozygous state (A/-) oould be studied usin g 22 bulls whioh had production tested daughters in each of the two categories* A total of 734 daughters of these sires wore availa­ ble for analysis*

The simple average of the two categories revealed that the heterozygous individuals produced 12,679 pounds of milk compared with

12,353 for animals lacking the A factor. However, the distribution of daughters of the bulls were non-orthogonal, therefore it became necessary to calculate oorrected mean difference as described by

Snedeoor (98) for disproportionate subsamples* The calculation of the corrected mean is shown in Table 3. In this procedure the difference between the two oategory means for eaoh sire was derived and recorded as positive if the heterozygous animals exceeded the homozygous animals and as a negative number if the reverse was true. The weight­ ing faotor was calculated by the following formula* W a al n 2 where n i ni+n2 represents the number of heterozygous daughters of a bull and n 2 th e number of homozygous daughters*

31 32

TAB IB 3

Comparison of the Hilk Production of Animals Hatororygoua and Homosygous a t the A Locus

U ixn2) Hetero Homo (3TL-X2) (ni+n-j) S irs n i S i * 2 5?2 D IT

lbs. lbs. lbs. 2 20 13577 36 13818 - 241 1 2 .8 6 3 21 9976 28 9537 + 439 1 2 .0 0 8 16 10688 11 10889 - 201 6.62 10 10 13393 2 12560 + 833 1.67 11 33 13219 39 12688 + 531 17.88 12 8 10453 6 9906 + 547 3.43 14 4 14072 9 13681 + 491 2.77 18 23 12037 11 12190 - 153 7.44 19 6 12991 9 12411 + 580 3 .6 0 21 14 13614 17 12715 + 899 7.68 22 6 14276 7 12350 +1926 3.23 23 21 13659 32 13826 - 167 12.67 24 30 14329 35 13890 + 439 16.15 25 16 10319 10 11311 - 992 6.15 26 9 12095 9 11028 +1067 4.50 27 13 14298 37 12871 +1427 9.62 28 9 10068 22 10946 - 878 6.39 33 5 12360 19 11007 +1353 3.96 34 5 13136 9 11113 ♦2023 3.21 35 10 13287 17 13364 - 77 6.30 38 23 12314 16 11105 +1209 9.44 39 28 13568 23 12353 ♦1215 12.63 TO 170.09

2 WD a +78963.61 Unbiased estim ate of difference « +464.25 lbs. 33

The summation of the weight times the differenoe for each sire comparison amounted to 78963*61* This figure represents the differenoe between the positive SWD (98825.66) and the negative XwD (19862*04).

The corrected or unbiased estimate of differenoe was obtained by dividing the USD figure by the summation of the weights. This amounted to 464*25 pounds of milk that the mean of the heterozygous animals exceeded the mean of those homozygous.

According to this method of arriving at an unbiased estimate of differenoe and testing it by the use of analysis of variance, it is necessary to proceed differently, depending upon the existence of

interaction between sires and blood factor categories* Therefore it was mandatory to run a preliminary analysis of varianoe to arrive at

the individual mean square to use as a comparison against the corrected

mean square for interaction* The preliminary analysis is presented in

Table 4 .

The corrected interaction sum of squares is obtained from the

following formula*

£TT

The correoted interaction sum of squares amounted to 861,972 and

the mean square was 41,046* When oompared with the individual mean

square of 42,691 an F value of less than one existed and this was far

below the level of significance*

After determining that interaction was not of significance, an

analysis of variance using corrected sum of squares was calculated* 34

TABIE 4

Preliminary Analysis of Varianoe for Milk Production as Affected by the A Loous

Degrees of “STSToF” ’ Mean Freedom Squares SquareSouroe

S ire 21 12,465,729 593,606

Antigen 1 206,253 205,253

Interaction 21 1,068,716 50,891

In d iv id u al 690 29,457,401 42,691

T otal 733 43,197,099 36

This analysis of varianoe, corrected for disproportionality, is given in Table 6. The antigen sum of squares which is used to test the unbiased difference between category means of 464*25 pounds of milk is derived by the following formula* From Table 6 i t can be seen that the differenoe between antigen oategory means was sig­ nificant at the one per oent level of probability. The sire differenoe also was significant at the same level of probability.

liThen. butterfat production (VgJ was compared between animals heterozygous and those homozygous for the absence of the A faotor, all

sires again were represented in both categories and were represented

by 733 daughters. Table 6 shows the subsample means, the within sire

differences and the weighting oaloulations• The unbiased estimate of difference was 10.62 pounds of butterfat, with the heterozygous group

e x c e llin g .

The preliminary analysis of variance revealed an individual mean

square of 56.43. The corrected mean square for antigen oategory

differenoe was 247.58. This resulted in an F value of 4.39 whioh was

above the five per oent probability level for the degrees of freedom

represented by these data, but did not reach the one per oent value.

Table 7 presents the comparison of maturity indexes (Vg) between

the two groups divided on the basis of heterozygous and homozygous

absence of the A faotor. The neoessity of two reoords on an individual

in order to be able to develop a maturity index reduced the number of

animals from the total that was involved in the study of milk 36

TABIE 5

Analysis of Variance Corrected for Disproportionality for Milk Production as Affected by the A Loous

Degrees of Sum of ------He an ” ' ...... V Source Freedom Squares Square Value

S ire 21 12,597,064 599,860 14.05

A ntigen 1 336,588 336,588 7.88

Interaction 21 861,972 41,046 •96

In d iv id u al 690 42,691

F(.05) df 1 - 690 . 3.86 F ( .o i) d f 1 - 690 a 6.70 37

TABLE 6

Comparison of the Butterfat Production of Animals Heterozygous and Homozygous a t the A Loous

Hetero Homo ( X1-X2) ( n i+ ^ i S ire n l * 1 n 2 'W D W

lb s . lb s . lb s . 2 20 446 36 463 -17 1 2 .8 6 3 21 359 28 345 +14 1 2 .0 0 8 16 363 11 391 -28 6.52 10 10 461 2 410 +51 1.67 11 32 495 39 476 +19 17.33 12 8 371 6 341 +30 3.43 14 4 445 9 441 + 4 2.77 18 23 430 11 445 -15 7.44 19 6 458 9 458 0 3.60 21 14 458 17 455 + 3 7.68 22 6 523 7 480 +43 3.23 23 21 499 32 498 + 1 12.67 24 30 511 35 507 + 4 16.15 25 16 371 10 387 -16 6.15 26 9 438 9 410 +28 4.50 27 13 547 37 499 +48 9.62 28 9 339 22 380 -41 6.39 33 5 420 19 394 +26 3.96 34 5 430 9 364 +66 3.21 35 10 455 17 467 +12 6.30 38 23 510 16 453 +57 9.44 39 28 517 23 488 +29 12.63

329 404 169.55

2WD a +1800.27 Unbiased estimate of differenoe a +10.62 lbs. 38

TAB IE 7

Comparison of the Maturity Inderas of Animals Heterozygous and Homozygous a t th e A Loous

He te ro Homo (Si-xgJ (ni+n2 ; S ire n i xi * 2 ST D W

2 13 90.46 20 83.90 + 6.56 7.87 3 14 95.93 16 99.75 - 4.22 7.46 8 8 111.75 6 103.66 + 8.09 3.42 10 7 86.42 2 1 0 0 .0 0 -13.58 1.55 11 21 91.52 28 103.28 -11.76 1 2 .0 0 12 7 103.14 4 92.50 +10.64 2.54 14 3 87.00 6 92.66 — 5.66 2 .0 0 18 1 2 93.33 8 99.50 - 6 .2 0 4.80 19 4 99.00 3 87.66 +11.34 1.71 21 8 90.87 10 92.10 - 1.23 4.44 22 3 90.00 3 105.66 -15.66 1.50 23 10 80.80 21 87.33 - 6.53 6.77 24 20 94.50 11 101.81 - 7.31 7.09 25 13 104.69 8 1 0 1 .1 2 + 5.57 5.66 26 4 89.25 3 89.00 + .25 1.71 27 8 94.25 26 98.11 - 3.86 6 .1 1 33 4 100.75 13 104.53 - 3.78 3.05 34 3 96.66 5 103.40 - 6.74 1.62 35 8 95.87 10 98.90 - 3.03 4.44 38 9 97.00 2 83.00 +14.00 1.63 39 11 94.90 12 96.00 - 1 .1 0 5.73

190 217 93.10 -246.63 Unbiased estimate of differenoe a -2 .6 4 39 production. The twenty-two a ire a wore aaoh represented in both antigen oategorieB but the total daughters involved was only 407.

An unbiased estimate of differenoe was calculated to be 2.64 units greater for those laoking the A faotor. These units are derived by orediting a oow which makes the same production on a mature equiva­ lent basis in the second lactation as she did on first lactation with

100 units. The 2.64 units of differenoe in this comparison indicated that the homozygous group had not reached as great a part of their eventual level at two years of age as had the heterozygous individuals.

However when this differenoe was tested by the analysis of variance as desoribed in the two previous comparisons, an P value of 2.54 resulted, whioh approached, but did not reaoh, the five per cent level of significance•

The heart girth measurement at birth (V 4J was a variable in which recordings were made on most animals. Tnerefore 968 animals were a v aila b le fo r t h is comparison as shown in Table 8. The unbiased estimate of 0 * 0 1 centimeter greater heart girth for the homozygous individuals was not a significant difference.

The three-month of age body condition (V 5) study was limited to

21 sires since sire number 14 did not have daughters representing him in both antigen categories and was therefore eliminated from the study.

The testing of mean square representing estimate of the differenoe of

0.102 resulted in an P value of 3 . 0 0 . The superior body condition of the heterozygous animal approaohed but did not reaoh a significant le v e l. This i s shown in Table 9 . 40

TABLE 8

Comparison of Hearth Girth Measurements at Birth of Animals Heterozygous and Homozygous at the A Loous

(n im 2J Hetero Homo (x i-x 2) (ni+n2J Sire - n ! — * r »2 ...... *2 D W

cm. cm. cm. 2 14 77.15 24 77.33 - .18 8.84 5 26 79.65 29 77.79 +1.66 13.71 8 16 79.13 12 79.92 - .79 6.86 10 11 80.27 2 81.00 - .73 1.69 11 52 81.25 67 81.78 - .53 29.28 12 9 76.44 5 79.80 -3.36 3.21 14 4 77.25 9 74.56 +2.70 2.77 18 41 80.00 16 80.44 - .44 11.51 19 9 72.56 14 75.43 -2.87 5.48 21 18 83.00 18 82.00 +1.00 9.00 22 9 79.33 8 79.63 - .29 4.24 23 29 77.66 47 78.98 -1.32 17.93 24 56 81.36 79 81.05 + .51 32.77 25 18 79.83 8 79.88 - .04 5.54 26 25 76.96 26 78.08 -1.12 12.50 27 25 76.80 59 77.83 -1.03 17.56 28 13 80.38 28 77.68 +2.71 8.88 33 8 81.50 32 80.09 +1.41 6.40 34 5 76.60 7 78.71 -2.11 2.92 35 12 73.50 29 74.90 -1.40 8.49 38 30 80.27 27 79.78 + .49 14.21 39 30 80.63 31 79.42 ♦1.11 15.26

460 576 253.42

5!ra>. -2.513 Unbiased estim ate of difference = —0.01 cm. 41

table: 9

Comparison of the Body Condition Evaluations at Three Months of Age for Animals Heterozygous and Homozygous at the A Loous

(nixngj H etero Homo (ni+ng) S ire nl *1 n 2 ®2 DW

2 9 4.777 17 4.411 ♦.366 5.885 3 12 5.583 15 5.533 ♦.050 6.666 8 16 5.062 12 5.166 -.1 0 4 6.857 10 9 5.777 2 6.000 -.2 2 3 1.636 11 49 5.142 58 5.156 -.0 1 3 26.661 12 8 5.375 4 5.500 -.1 2 5 2.666 18 41 4.853 17 5.176 -.3 23 12.017 19 10 4.900 12 4.916 -.0 1 6 5.454 21 18 6.722 12 5.750 -.0 2 8 7.200 22 9 6.000 10 4.900 ♦.100 4.736 23 29 4.827 49 4.612 ♦ .215 18.217 24 53 5.566 76 5.328 ♦ .238 31.224 25 18 5.777 8 5.625 ♦ .162 5.538 26 25 5.240 25 4.880 +.360 12.500 27 25 4.960 61 5.049 -.0 8 9 17.732 28 13 4.615 28 4.821 -.2 0 6 8.878 33 9 5.777 35 5.485 +.292 7.159 34 5 5.000 8 5.125 -.125 3.076 35 12 4.666 30 4.833 -.1 6 7 8.574 38 SO 4.666 27 4.074 +. 592 14.211 39 31 4.483 31 4.226 +.258 15.500

431 537 222.287 ♦22.717 Unbiased estimate of difforenoe « ♦0.102 condition units 42

The amount of gain between the ages of three and six months (VgJ

for animals in each of the two categories is shown in Table 10* The

0.05 centimeter greater gain by the heterozygous group was too small to be significant and the same was true for the 0.337 centimeter

greater growth between the age of three months and the measurement

taken three months after the first calving (V 7J as shown in Table 11.

The services required for conception for second calving (Vq) was

compared in Table 12 and the 0.026 greater servioes required for

conception among the heterozygous individuals was determined to be not

significant.

In comparing the birth measurement of the female offspring of the

two year old animals (V 9J in the two categories only 19 sires had

daughters distributed such that they oould be used. The 0.11 centi­

m eter g re a te r siz e o f th e o ffsp rin g o f heterozygous dams was shown to

be not significant. These data are presented in Table 13.

Of all the comparisons made involving the A system, the most

important, both from the standpoint of eoonomical worth and probability

of a true difference, is that shown in the study of milk production.

The figure of 464 pounds of milk may not at first reflection appear to

be a large addition to cows already produoing at 1 2 ,0 0 0 pound level.

However, increments of genetic improvement in dairy oattle come slowly

and it appears that no other evaluation of animals could be used to

divide individuals within a herd from a common sire into two nearly

equal groups and have one group excel the other to Buoh a great extent.

Especially is this important sinoe the division according to blood 43

TABLE 10

Comparison of the Increase in Hearth Girths From Three to Six Months of Age in Animals Heterozygous and Homozygous a t th e A Loous

H etero ...Homo (x i-x 2 ) (ni+ n2J S ire n i x i n 2 D W

cm* ora. cm. 2 9 2.611 17 2.741 -.130 5.884 3 8 2.250 12 2.258 -.008 4.800 8 16 1.812 11 2.072 -.2 60 6.518 10 9 2.388 2 2.100 +.288 1.636 11 50 2.310 66 2.330 -.0 2 0 28.448 12 7 2.300 4 2.175 + .125 2.545 18 39 2.348 17 2.211 +.137 11.839 19 10 2.890 12 2.633 +.257 5.454 21 18 2.422 18 2.416 +.006 9.000 22 9 2.633 10 2.990 -.3 5 7 4.736 23 29 2.589 46 2.682 -.093 17.786 24 55 2.354 78 2.297 + .057 32.255 25 16 1.962 8 1.875 + .087 5.333 26 24 2.270 25 2.324 -.0 5 4 12.244 27 26 2.665 61 2.780 -.1 1 5 18.229 28 9 2.777 27 2.077 +.700 6.750 33 9 2.277 35 2.222 + .055 7.159 34 4 2.000 8 1.850 + .150 2.666 35 12 2.783 30 2.890 -.1 1 7 8.571 38 29 1.979 25 2.044 -.0 6 5 13.425 39 28 2.042 29 2.010 +.032 14.245

416 591 219.533 £ yvd a ♦1 .10 Unbiased estimate of differenoe - +0 .01 am. 44

TABLE 11

Comparison of the Increase in Heart Girths From Three Months of Age to Three Months After F irst Calving in Animals Heterozygous and Homozygous a t the A Locus

in ix n 2 ; H etero Homo (xi-X 2) (ni+ng) S ire n i x i »2 *2 D W

om. cm. om. 2 7 84.00 13 85.92 -1 .9 2 4.550 3 6 88.50 7 89.43 ™ .93 3.230 8 15 86.33 10 87.80 -1 .4 7 6.000 10 9 85.33 2 80.50 +4.83 1.636 11 31 81.16 40 79.65 +1.51 17.464 12 6 89.33 4 35.75 +3.58 2.400 18 15 86.06 7 83.14 +2.92 4.772 19 7 82.14 10 83.10 - .96 4.117 21 13 85.08 17 81.76 +3.32 7.366 22 5 80.60 6 86.67 -6 .0 7 2.727 23 22 86.06 27 85.78 + .27 12.122 24 35 83.26 42 81.40 +1.86 19.091 25 15 85.00 8 89.00 -4 .0 0 5.217 26 8 85.38 9 93.44 -8 .0 6 4.235 27 15 86.20 38 83.84 +2.36 10.754 28 9 91.78 21 87.63 +4.15 6.300 33 4 82.26 12 82.42 -0 .1 7 3.000 35 10 87.30 18 85.39 +2.41 6.428 38 25 92.60 15 94.87 -2.27 9.375 39 28 88.14 24 90.21 -2 .0 7 12.923

285 330 143.712 S tm m +48 • 52 Unbiased estimate of difference =* +0.337 om . 45

TAB IB 12

Comparison of Services per Conoeption for Animals Heterozygous and Homozygous a t the H Locus

Hetero Homo (n'l+ngj S ire n i afr n 2 *2 D W

2 14 1.214 28 1.964 - .750 9.333 3 20 3.000 21 2.761 + .239 10.243 8 16 3.062 11 1.454 +1.608 6.518 10 9 1.333 2 3.000 -1.667 1.636 11 32 1.937 37 2.270 - .333 17.159 12 7 2.285 6 2.000 + .285 3.230 14 4 1.500 7 1.714 - .214 2.545 18 21 3.000 11 3.181 - .181 7.218 19 6 1.666 9 1.444 ♦ .222 3.600 21 12 1.916 15 1.933 - .017 6.666 22 5 1.400 4 1.500 - .100 2.222 23 18 1.277 31 1.580 - .303 11.387 24 28 1.607 31 1.870 - .263 14.711 25 16 3.000 10 2.300 + .700 6.153 26 8 2.125 7 2.428 - .303 3.733 27 13 2.000 8 1.833 + .167 4.952 28 9 2.777 20 2.450 + .327 6.207 33 5 1.000 18 2.388 -1.388 3.913 34 5 2.400 9 1.444 + .956 3.214 35 10 1.400 17 1.529 - .127 6.296 38 20 2.050 15 2.466 - .416 8.571 39 27 3.148 24 1.916 +1.232 12.705

306 _ 341 152.212 a +3.898 Unbiased estim ate of difference » +0.03 services 46

TABIE 13

Comparison of the Ha art Girths of the First Offspring of Animals Heterozygous and Homozygous a t the A Locus

u v v Hetero Homo (XL-JC2) S ire n i *1 *2 ST DW

cm* cm. cm. 2 6 76.166 11 75.818 ♦ .348 3.882 3 12 74.666 13 76.846 -2.180 6.240 8 9 79.111 3 77.000 ♦2.111 2.250 10 7 79.000 2 83.000 -4.000 1.555 11 16 80.062 19 79.473 + .589 8.685 14 2 75.600 5 76.000 - .500 1.428 18 8 77.875 3 78.333 - .458 2.181 19 2 70.500 4 71.250 - .750 1.333 21 9 79.000 8 78.375 ♦ .626 4.235 23 9 77.777 16 75.562 ♦2.215 5.760 24 15 80.266 17 78.705 +1.561 7.968 26 6 79.333 4 81.000 -1.667 2.400 27 8 74.125 16 74.312 - .187 5.333 28 3 79.333 9 77.000 ♦2.333 2.225 33 3 80.000 7 79.142 ♦ .858 2.100 34 2 74.000 6 77.353 -3.333 1.500 35 4 72.500 10 74.100 -1.600 2.857 38 8 77.626 2 80.000 -2.375 1.600 39 9 79.777 12 78.583 +1.194 5.142

138 167 68.67 ♦7.287 Unbiased estimate of difference ■ ♦0.11 cm. 47 factor patterns could conceivably be made before the conception takes plaoe. The commonly accepted manner of selecting for superior off­ spring in a herd is by seleoting the daughters of cows above the average of the herd. If we accept a heritability estimate of milk production at .25 it would mean that the top half of the herd would have to average 3,682 pounds more milk than the lower half in order that 464 pounds more milk would be realised from the daughters of the top cows.

In addition it would seem that there is nothing to prevent the benefits resulting from blood factor selection from being superimposed upon the procedure of seleoting daughters from superior dams.

In noting the comparisons for individual sires in Table 3 it oan be seen that in seven of the 22 comparisons, the animals completely laokihg factor A exceeded their half sibs in production. This might be considered evidence pointing to the lack of usefulness of such a method. If we return again to dividing the mates of bulls into a high and a low half and calculate the production average of their respective daughters one should not be surprised to find the daughters of the low cows exceeding the daughters of the high c o w b seven twenty-

seconds of the times. Yet no one questions the validity of seleoting daughters from better dams.

A genetic explanation of why animals carrying a single dose of blood factor A should produce more milk than animals without the

factor would be desirable. If we oonsider the portion of the chromo­

some controlling this blood factor as a gene, then it is possible

that this gene has a pleiotropio effect. If this explanation is 48 correct, then animals of the gene tie complement A/A should be equal to

A/- animals if dominance is the gene relationship in milk produotion or they should be superior if the gene effeots are additive* A seoond possibility is that the A loous is linked to genes effecting milk produotion. Evidenoe of a linkage effeot oould be found if a positive relationship between milk produotion and the A factor existed in some families and a negative relationship ooourred in others suoh that a canceling out of effeots would ooour and no relationship would appear to exist -when a population of families was studied* S till another possible explanation oould be that the presenoe of two alleles at this loous indicates that the chromosomes on whioh they are carried originated from different sources and then presumably some genes affecting milk production would also be different and thus heterosls is being exhibited* If this latter explanation is true and the factors are of no importance themselves but serve only as markers to indicate the amount of heterozygosity that exists then presumably* animals homozygous dominant for A should not perform differently from animals homozygous for the absence of A* A study of the third oategory which contained animals having A but undetermined as to genotype (A/- or A/A) Beamed the logical prooedure to gain additional information which might be evidenoe for one of the aforementioned possibilities* A comparison was made between animals having this third oategory (A/A, A/-) and animals by the same sire that laoked A

(-/-) and is shown in Table 14* Also a comparison of this third 49

TABLE 14

Comparison of the Milk Production of Animals Homozygous Dominant and Homozygous R ecessive a t th e A Loous

A/A (Sl-xg} ------^ (n i« i2 ^ S ire »1 a 2 *2 D W

lb s . lb s . lb s . 2 41 13353 36 13818 - 465 19.17 10 27 12791 2 12560 + 231 1.86 12 21 10206 6 9906 + 300 4.67 14 6 14286 9 13581 + 705 3.60 18 3 13377 11 12190 +1187 2.36 19 13 12506 9 12411 + 95 5.32 22 6 13908 7 12350 +1558 3.24 25 27 10927 10 11311 - 384 7.30 26 14 11225 9 11028 + 197 5.48 34 19 11411 __ 9 11113 + 298 6.11

59.11 <5?© * +3906.39 Unbiased estimate of difference a +66*08 lbs* 50 oategory and animals heterozygous is shown in Table 15* Ten sires were available for this study and in the first comparison the animals in the mixed group (A/A, A/-) exoeeded the group laoking A by 66 pounds of milk •while the animals known to be heterozygous had 216 pounds more milk than the third oategory (A/A, A/-). Neither of these two figures was statistically significant; however, the intermediate position of the mixed group oould be construed as evidenoe that the A

gene itself was not exerting a pleiotropic effect or else the popu­

lation with its high frequency of A faotors should have excelled the

known heterozygous group# The bulls that had daughters in the mixed

group were the bulls carrying heterozygous A and it would be in this

group one would logioally look for evidence of extreme positive and

negative associations if linkage were the cause of the effects*

Apparently there was no more indication of this occurring in thiB

group than among the remaining sires which were completely laoking in

blood factor A themselves and the A factor in their offspring came

from the variety of cows to whioh they were mated* The general

inorease in heterozygosity as indicated by heterozygosity at the A

loous appears to be the explanation whioh best fits this information*

If the difference in milk produotion as it is affeoted by the A

loous in this group of oattle is shown by further studies to be a

characteristic of all Holstein-Friesian oattle or of all dairy animals,

then it might well be worthy of consideration in breeding plans* The

productive life expeotanoy of a dairy ocw appears to be about four

years, therefore the 464 additional pounds of milk per lactation 51

TABLE 16

Comparison of the Milk Produotion of Animals Heterozygous and Homozygous Dominant a t the A Loous

InjT O jJ V- A/A (nx+n2i S ire n i x i n 2 *2 D W

lb s . lb s . lb s . 2 20 13577 41 13353 + 224 13.44 10 10 13393 27 12791 + 602 7.30 12 8 10453 21 10206 + 247 5.79 14 14 14072 6 14286 - 214 4.20 18 23 12037 3 13577 -1340 2.65 19 6 12991 13 12506 + 485 4.11 22 6 14276 6 13908 + 368 3.00 25 16 10319 27 10927 - 608 10.05 26 9 12095 14 11225 + 870 5.48 34 5 13136 19 11411 +1725 3.96

59.98 & D * +12971. Unbiased estimate of differenoe a +216.26 lbs* 62 produced, by the animals bearing heterozygous A oould be roughly estimated to be worth $75.00. Under suoh a program* females of the blood factor complement -/- should be bred to sires homozygous for the

A faotor* while females bearing A should be mated to bulls lacking A«

All of the -/- females would then bear heterozygous offspring as would the females homozygous for factor A. Those bearing A whioh are heterozygous for this factor should have 50 per oent heterozygous off­ spring whioh is the maximum to be expected from any type of sire. This then would leave the homozygotes among the offspring absent of the factor so that they could be easily identified and mated for the next generation. Indications are that it may be possible to Identify hetero­ zygous animals by typing procedures whioh would permit an outstanding sire heterozygous for factor A to be mated to females shown in the laboratory also to be heterozygous without any greater loss of hetero­ zygosity in the offspring than when a homozygous A absent bull was u sed .

The study involving the second variable* butter fat production* would have been expeoted to follow the same pattern as milk production.

In general that is true, however the superiority expressed in the heterozygous individuals amounted to ten pounds of butter fat. This is

somewhat less than would have been expeoted considering the difference that existed in milk produotion. Also this difference did not present

a level of significance as high as milk produotion. This suggests

that* in spite of the faot that the heterozygous animals produoed more

milk and butterfat* the homozygous individuals probably carried the 53 higher percentage of butterfat. This also would not be out of line with observations that in many inbreeding studies the inbred animals, whioh are supposedly more homozygous, frequently test higher than those randomly bred,

The third variable was of considerable interest in that it might give some insight into the genetic aotion involved. Again referring to work involving inbreeding, animals inbred and therefore more homozygous have shown a decided tendenoy to mature slower than heterozygous or randomly bred contemporaries. The calculation of the maturity index in this study come about by dividing the first laotation record, figured to a mature equivalent into the second -lactation also computed on a mature equivalent basis. By this method an animal which was slow maturing would have a value above one (coded to 100 for this study ) while a rapidly maturing animal would have her largest mature equiva­

lent record at two years of age and therefore would have a maturity

index of less than one. To fu lfill the hypothesis that this was a

case of generally increased heterozygosity, this comparison snould

reveal that the homozygous individuals carried a higher average index

than did their heterozygous half sibs. The homozygous (-/-) anim als

did exceed by 2.64 unitsj nevertheless, this difference was not great

enough to be significant at the five per cent level but did carry an F

value of 2.54.

The remaining variables appear not to be important enough to

justify individual attention except possibly for the score of body

condition at three months which favored the heterozygotes by 0.102 54 units and -which, when tested, had an F value of 3.00 and was just short of significance.

The summary of the A loous appears to deserve oomment. I t oan be noted in examining Table 16 that most of the mean differences have been in favor of heterozygous animals even though some have not been at significant levels. The exceptions are variable 3, the maturity index, which fits the pattern better as a negative quantity and variable 4, the birth measurement, whioh has a slight negative value.

The eighth variable, services per conception, also is a positive value but I have no opinion as to whether this adds to or detracts from the

general pattern.

F/V Loous

The genotype of the individual can be determined by laboratory means, therefore the F/V would seem to be a very easily studied system.

However the low frequenoy of the V factor in this population eliminated

any opportunity of acquiring a useful sample of animals carrying V in

a homozygous oondition. Therefore the comparison was made between

animals carrying the F/V genotype and a group made up by pooling the

homozygous individuals whether they were F/F or V/V•

Hone of the variables had significant differences between the

homozygous and heterozygous groups, th e re fo re only th e summary of t h i s

locus is being inoluded in this text. The within sire comparisons for

each of the variables at this loous are to be found in the Appendix

Tables 24 through 32. 66

TABLE 16

Summary of Unbiased Differences, for the Hine Variables, Between Animals Heterozygous and Homozygous a t the A Loous

S ire s Hetero Homo Unbiased V ariable Involved *1 n J> D ifference 1 22 330 404 4464*25 pounds of milk v*

2 22 329 404 ♦ 10.62 pounds of butterfat *

3 21 190 217 - 2*64 m atu rity index u n its

4 22 460 576 M 0.01 centimeters

5 21 431 537 4- 0.10 condition units

6 21 416 491 + 0.01 centimeters

7 20 285 330 4- 0.34 centimeters

8 22 305 341 4- 0.03 services per conception

9 19 138 167 4> 0.11 centim eters

4W P<.01 * P<.05 56

The pattern or trends at this loous as shown in the summary in

Table 17 are not very olear cut.

The largest F value obtained for any of the comparisons at this

locus m s only 1.7 and this does not approaoh a level of significance

of five per cent. This faot would suggest that, within the sires

studied, the comparison between females heterozygous for blood faotors

F and V and those homozygous for either of the faotors resulted in no

difference for the nine variables studied. These results and

conclusions are not too disturbing when it* s recalled that in the

ohioken work certain loci were shown to be associated with certain

variables and not with others.

J Loous

The interpretation of any associations at the J locus might not

be dear. Animals whioh lacked the J substance on their cells may

have had it in their serum. If one of the postulations of Stone and

Irwin (102) be truo that three alleles occur (Jos on cells and in

serum, JjB in serum only, and j a absence of factor) then the animals

designated as heterozygous may have been either Jcs/js or Jos/ja and

the other group laoking the factor on cells may have been composed of

three genotypeB J b/J s, j a/ja or Js/ja, if heterozygosity m s at work

at this loous, then the difference would be reduced due to the fact

that the homozygous indicated group m s oontaminated with some

heterozygous individuals. 57

table: 17

Summary o f the Unbiased D iffe re n c e s, fo r th e Nine V a ria b le s, Between Animals Heterozygous and Homozygous a t th e F/V Loous

S ire s ifeuero Homo Unbiased V ariable Involved Pi P2 . D ifferen ce 1 15 167 513 -205*80 pounds of milk

2 15 168 513 - 6*20 pounds of butterfat

5 13 77 268 - 1.91 maturity units

4 16 219 638 - 0.29 centimeters

5 16 216 602 - 0.08 body oondition units

6 16 220 606 •f 0.17 centimeters

7 15 142 395 - 0.91 oentimeters

8 15 150 412 - 0.01 services per conception

9 12 66 177 + 0.27 centimeters 58

The roaults at this loous presented no significant differences* therefore the results of the within sire oomporisons for each -variable are p resen ted in Appendix Tables 55 through 41* The g en eral summary

of these comparisons at the J loous is presented in Table 18* An

inspection of this table strongly suggests that the animals are divided into two groups whioh might actually be performing differently

providing some refinements and further ganotypio determinations are made* Of the seven variables studied whioh would indicate 'fitness1

or ability to perform well* six had a difference favoring the

heterozygous group* The second variable was particularly impressive

in that the heterozygous group exceeded their half Bibs by an average

difference of 11.52 pounds of butterfat. This represented a larger

difference than was obtained at the A locus* however the difference was

not statistically significant due to a reduced number of total animals

and also because they were not balanced between the two categories*

Variables 4* 6 and 7* along with the aforementioned variable 2* all

had P values which approached significant levels. It would appear

that with the testing for all alleles at this loous and soma

realignment of individuals on this additional information, that there

is a very good possibility that true differences in performance exists

between homozygous and heterozygous animals at the J locus*

L Locus

The results obtained in studying the L loous proved to be one of

the more interesting series of comparisons* 59

TABLE 18

Summary of tbs Unbiased Differences, for the Nine Variables, Between Animals Heterozygous and Homozygous a t the J Loous

S ire s H etero Homo Unbiased V ariable Involved D ifference fli *9 1 15 126 356 +232*69 pounds of milk

2 15 126 356 + 11.52 pounds of butterfat

3 13 70 130 - 0.68 maturity units

4 18 196 603 + 0.59 centimeters

5 17 191 576 - 0.01 condition units

6 17 187 586 + 0.55 centimeter

7 16 111 346 + 1.05 oentimeters

8 16 120 349 + 0.13 services per conception

9 14 51 149 + 0.27 centim eter 60

A summary of the nine variables at this loous is given in Table

19. The first three variables involving milk production, fat pro­ duction and the maturity index presented mean differences whioh favored the heterozygotes but were not significantly different.

However, variable 4, the heart girth measurement, differed between categories by 0.76 of a centimeter. This difference proved to be h ighly sig n ific a n t (P<.OiJ when te s te d .

In contrast to the results obtained at the A locus, the animals that were completely lacking the L factor performed better than did the animals oarrying the heterozygous genotype. An explanation of why animals lacking L factor might grow a little more while in the uterus is not readily apparent. A possibility would be that the portion of the chromosome responsible for the L factor might be producing a substance inhibitory to growth, or failing to produoe a substanoe necessary for most rapid growth. The position of the mixed group made up of animals having L, but undetermined as heterozygosity or homo­ zygosity appeared again to be the likely source of evidenoe as to what was happening. I f th e segment of the chromosome producing L was also producing a substance inhibitory to growth, then the mixed group should equal the heterozygous group, if simple dominance were the mode of

inheritance, or be depressed to an extent greater than were the heterozygotes if the effects of L were additive. A comparison between animals of the mixed category (L/L, L/-J and those known to be heterozygous (L/-J was made. For this study 14 sires with 319 61

TAB112 19

Summary of the Unbiased. Differences, for the Uine Variables* Between Animals Heterozygous and Homozygous at the L Loous

6 ire s lle tero ttomo Unbiased V ariable Involved . . “I n J> D ifferen ce 1 18 141 524 +211.99 pounds of milk

2 18 141 523 + 7.17 pounds of butterfat

3 16 71 288 + 1.04 maturity units

4 19 213 751 - 0.76 centimeter

5 18 207 687 - 0.05 condition unit

6 18 194 686 + 0.1 6 cen tim eter

7 17 124 406 - 1.53 centimeters #

8 18 141 473 + 0.01 service per conception

9 13 83 163 - 0.05 oentiraeter

F<.01 # P<.05 62 daughters were available. The weighted mean differenoe showed the mixed group to be smaller in size than were the heterozygotes by 0.61 centimeter. This difference did not prove to be significant. However, the magnitude of this difference suggests that if this mixed category were clearly defined and only homo2ygous animals were inoluded that the depressing effeots of the L factor might well be additive*

Body condition, whioh is variable 5, also was less among animals having L. However this difference did not approach significant levels.

Variable 6 is the growth measurements between three and six months and showed a small but insignifioant amount of greater growth in heterozygous L animals.

The study of variable 7, whioh includes the amount of growth between three months of age and the three months after calving measure- ment added further evidenoe to the depressing effeot of L on growth.

Animals heterozygous for L added 1.53 centimeters I osb heart girth in this period of time than did animals laoking in L. This differenoe proved to have significance at the five psr cent level of probability.

Service per conception had a small positive but insignificant

value with the heterozygote requiring more services. The heart girth measurement of the first offspring if a female, also showed a slightly

lower value among the animal carrying one dose of L when compared with

those laoking L. The detailed comparisons at this locus are presented

in Appendix Tables 42 through 50. 63

SU Loous

Results at the SU loous revealed that the measurements of all variables differed little between animals heterozygous and those apparently homozygous for the observed alleles* The summary of these comparisons i s shown in Table 20*

The oonolusion that oan be moat logically drawn from this series of comparisons is that, among the animals studied, the presence of heterozygosity at this locus exerted no effect upon the nine variables studied*

The within sire comparisons for eaoh variable and the calculations of the weighted mean difference are presented in Appendix Tables 51 through 59.

Z Loous

The summary of the comparisons at the Z locus is presented in

Table 21 of this text and the detailed comparison for each variable is to be found in Appendix Tables 60 through 68*

At the Z loous no patterns or trends were apparent and with one exoeption differences were far below the level of significance.

The amount of inorease in heart girth between three and six months did prove to have a differenoe between categories which was significant at the five per cent level. Heterozygosity may again be exerting an effeot at this locus on this early growth measurement but the laok of any such indications on other variables suggests that this may well have been a chance differenoe. As laboratory tests are available on 64

TAB IS 20

Summary of the Unbiased Differences, for the Nine Variables, Between Animals Heterozygous and Homozygous at the SU Loous

S ire s Be te r o Homo Unbiased Variable Inyo I've d n^ Difference

1 19 186 566 -3.07 pounds of milk

2 19 185 566 -4.77 pounds of butterfat

3 13 90 228 -0.61 maturity unit

4 19 264 785 -0.02 centimeter

5 19 253 743 +0.08 condition unit

6 18 245 681 +0.14 oentimeter

7 16 163 434 +0.16 centimeter

8 18 174 476 +0.04 service per conception

9 12 73 136 -0.05 centimeter 65

TABUS 21

Summary of the Unbiased. Differences, for the Nine Variables, Between. Animals Heterozygous and Homozygous a t the Z Locus

S ire s Hetero Homo Unbiased V ariable Involved n i *2 D ifference

1 20 209 374 -63*48 pounds of milk

2 20 209 374 - 3.84 pounds o f b u tte r f a t

0r» 18 112 192 + 0.56 maturity unit

4 21 298 554 - 0.11 centimeter

5 20 273 514 + 0.002 oondition unit

6 20 268 511 +0.72 centimeter

7 18 182 302 - 0.03 centim eter gain

8 20 192 348 +0.13 service per conception

9 17 90 149 - 0.02 centimeter 66 th ese animals to determ ine a l l the ho te r o zygotes, numbers may become available whioh -will resolve this question.

Combined Looi

An attempt was made to study the effect on animal performance of heterozygosity at several looi as compared with heterozygosity at a few looi. 'Hi© five looi, A, F/V, L, STJ and Z were used in these compari­ sons. Animals homozygous at all five looi and those heterozygous at only one loous were grouped in one category while animals heterozygous at two, three, four or five looi were in another category. The within

sire comparisons of animals within the two categories were made for eaoh of the nine variables and they are presented in Appendix Tables

69 through 77. A summary of th ese comparisons are presented in Table

22. From this summary table no general differences are apparent and none of the individual difference* are of such magnitude as to be

significant.

A summary of a ll comparisons between animals heterozygous and

homozygous at the six looi is presented in Table 23. 67

TABUS 22

Summary of the Unbiased Differences, for Nine Variables, Between Animals Heterozygous at Two or More Looi and Animals Heterozygous at Not More Than One Locus

H etero H etero S ire s (2 ,3 ,4 ,5 ) ( o , i ; Unbiased V ariable Involved n 2 D ifferen ce 1 16 157 187 -229.45 pounds of milk

2 16 156 187 - 7.46 pounds of butterfat

3 12 118 75 - 3.27 maturity index units

4 16 249 305 + 0.26 centimeter

5 16 244 290 - 0.07 oondition unit

6 16 238 282 > 0.43 centimeter

7 15 154 158 - 0.17 centim eter

8 15 145 165 ♦ 0.14 service per conception

9 10 49 60 + 0.24 oen time ter I

TAB 13 23

Summary of All Comparisons Between Animals Heterozygous and Homozygous a t th e S ix Looi

V ariable Vl " V2 V3 V4 V5 Vs V7 va * w ■ Locus Lbs* Lbs* U nits cm* U n its cm* cm* Sue* cm*

A +464.25H# 4-10.62# -2 .6 4 -.0 1 4-.10 4-.05 + .34 + .03 +.11

F/7 -205.80 - 6.20 -1 .9 0 -.2 9 -.0 8 4-.17 - .91 -.0 1 +.27

J 4-232*69 4-11.52 — .68 +• 59 -.0 1 4-. 55 +1.05 + .13 +.27

L 4-211*99 4- 7.17 4-1.04 - .7 -.0 5 4-.16 -1.53 # + .01 -.0 5

SU - 3.07 - 4.77 - • 61 -.0 2 4-.08 +.14 + .16 + .04 -.0 5

Z - 63.48 - 3.84 4. .56 -.1 1 4-.002 +.72# - .03 +.13 -.0 2

* (P<.06j ■H# (P<*Ol)

09a SUMMARY AND CONCLUSIONS

An investigation was undertaken to see if a population of Holstein-

Friesian oattle performed differently depending upon the existence of heterozygosity or homozygosity at looi oontrolling several blood fao to rs*

The A, F/Y, J, L, SU and Z looi were each studied in relation to the following performance variables* milk production, butterfat pro­ duction, maturity index, heart girth at birth, body condition at three months of age, heart girth inorease from three to six months, heart girth gain from three months to three months after first calving, services per conception at second breeding and heart girth measurement of first calf if female*

The animals used were in the institutional herds of Ohio under supervision of the Department of Mental Hygiene and Correction*

The comparisons between the heterozygous and homozygous

(recessive) individuals were made within sire, and an overall unbiased estimate of mean differences was oaloulated and tested by the analysis of variance*

Animals heterozygous at the A loous (A/-) produced, on the average, 464*26 pounds of milk more than did animals homozygous (-/-) at this locus* This difference proved to be highly significant

(P<*Ol)* The heterozygous group also excelled in butterfat produced

69 70 by 10*6 pounds. Ibis also was significant (P<.05). Three hundred thirty heterozygous and 404 homozygous daughters representing 22 sires ware available in both the milk and butterfat production studies. All other comparisons at this loous were below the value neoessary for significance. However, animals shown to be heterozygous matured more rapidly as indicated by the maturity index, and the difference of 2.64 maturity unitB approaohed a significant level.

The study of the L locus revealed thaL individuals laoking the factor were 0.76 centimeter larger at birth and had 1.53 centimeters more growth between three months of age and three months after calving.

The former was significant at the .01 level and the latter at .05. Mo other variable differed significantly when the study of this loous was com pleted.

Animals heterozygous at the Z loous (Z/-) grew 0.72 centimeter more between three and six months than did animals laoking Z (-/-) *

This was the only significant comparison involving this system.

No significant differences were found in comparisons made at the

F/V, J or SU looi. Also groups made up of animals heterozygous at two or more loci and animalB heterozygous at not more than one loous did not differ for any of the traits studied.

The following conclusions would seem to follow from the information aoquireds

1. Within the population studied, animals exhibiting hetero­ zygosity at the A locus produced significantly more milk and butterfat than did animals lacking the A factor. 71

£• If this is shown to be a oharaoteristio of all dairy oattl© p o p u latio n s» i t oould be as e ffe o tiv e a method. of s e le c tin g immature daughters of a sire as any method now available.

5. Animals carrying L factor in their blood grew less rapidly and were smaller at birth than animals lacking the faotor*

4* Heterozygosity at the Z locus resulted in animals that grew significantly more between three and six months of age*

5* The P/7, J, and STJ looi had no significant effect upon the traits studied*

6* Animals heterozygous at two or more loci did not perform differently from animals heterozygous at not more than one loous* BIBLIOGRAPHY

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120. Todd, C», and White, R. G. On the Haemolybio Immune Isolysins of the Ox and Their Relation to the Question of Individuality and Blood Relationship. Journal of Hygiene, 10* 185, 1910.

121. Treeoe, *1. M., Gilmore, L. 0., Washburn, R. G., and Feohheimer, H. S. Evidence of Inherited Influences Affecting Casein Yield and Percentage as Measured by Relationships to Sire Groups and Cellular Antigens. J . Dairy Soi., 42* 922, 1959.

122.Wiener, A. S. History of Rhesus Blood Types. Journal of the History of Medicine and Allied Soienoe, 7* 369, 1952.

123. Wiener, A. S., and Sonn, E. B. Parthogenesis of Congenital Hemolytio Disease Erythroblastosis Fetalis. American Journal of Diseases of Children, 71* 25, 1946.

124. ______. Permeability of the Human Placenta to Isoantibodies. Journal “of Laboratory and Clinical Medicine, 31* 1020, 1946.

125. Woolf, B. On Estimating the Relationship Between Blood Groups and Disease. Annals of Human GenetioB, 19* 251, 1954-55.

126. Yannet, H. The Importance of the Rh Factor in Mental Deficiency. Hew York Academy of Medioine, 20* 512, 1944. XICIKSHdV 83

TAB IE 24

Comparison of the Milk Production of Animals Heterozygous and Homozygous at the F/V Locus

(niocnj,; H etero Homo (TL-X2) < »1«2) S ire *1 X l n2 *T D W

Lbs* Lbs* Lbs* 2 22 13388 75 13626 - 238 17.01 10 6 13366 33 12854 ♦ 512 5.07 11 27 12812 45 13003 - 191 16.88 14 3 13380 16 14006 - 626 2.53 19 3 11540 25 12704 -1164 2.68 21 11 12879 20 13265 - 376 7.10 22 5 13636 14 13384 + 252 3.68 23 9 14507 44 13607 + 900 7.47 25 8 11903 50 10912 + 991 6.90 27 9 12620 41 13379 - 759 7.58 28 14 10290 17 11022 - 732 7.68 33 5 9358 19 11797 -2439 3.98 34 13 10921 20 12027 -1106 7.88 35 14 13227 13 13451 - 224 6.74 38 18 12018 21 11647 + 371 13.03

167 513 116.01 Zw> = -23882 Unbiased estimate of difference = -205.80 lbs* 84

TAB IE 26

Comparison of the Butterfat Production of Animals Heterozygous and Homozygous a t th e F/V Loous

tm x n jJ Hetero Homo (*L“X2) (ni+ n2J S ire *1 x i n 2 * T D W

Lbs* Lbs* Lbs* 2 22 446 75 456 -10 17.01 10 6 450 33 438 +12 5.07 11 26 482 45 486 - 4 16.87 14 3 430 16 452 -22 2.53 19 3 427 25 457 -30 2.68 21 11 445 20 463 -18 7.10 22 5 504 14 501 + 3 3.68 23 9 504 44 497 + 7 7.47 25 8 446 50 376 +70 6.90 27 9 490 41 517 -27 7.38 28 14 353 17 380 -27 7.68 33 5 348 19 413 -66 3.98 34 13 358 20 396 -38 7.88 35 14 455 13 471 -16 6.74 38 18 497 21 478 +19 13.03

168 513 116.01 ZttD * -719.30 Unbiased estim ate of difference =»-6.2 lbs* 85

TABLE 26

Comparison of the Maturity Indexes of Animals Heterozygous and Homozygous a t the F/V Locus

Hetero Homo ( ^ - x g ) (ni+n25 S ire n i x i »2 * r D W

2 13 86.6 45 89.5 - 2.9 10.1 10 2 88.0 25 93.4 - 5.4 1.9 11 18 97.2 31 98.8 - 1.6 11.4 14 2 105.5 10 93.6 +11.9 1.7 21 3 81.3 15 93.6 -12.3 2.5 22 3 100*3 6 90.8 + 9.5 2.0 25 4 76.0 27 86.6 -10.6 3.5 25 4 83.3 39 101.1 -17.8 3.6 27 7 103*4 27 95.6 + 7.8 5.6 33 3 106.0 14 102.8 + 3.2 2.5 34 5 104.6 13 100. b + 4.1 3.6 3b 9 95.6 9 99.6 - 4.0 4.5 38 4 93.5 7 95.0 — 1.5 2.5

77 268 55.4

3 -105.8 Unbiased e stim ate of difference a -1.91 86

TABIE 27

Comparison of the Heart Girth Measurements at Birth of Animals Heterozygous and Homozygous at the F/V Loous

Hetero Homo (*1-*2J S ire n l ^1 nz x2 D w

om. cm. cm. 2 15 75.9 49 77.8 -1 .9 11.3 10 6 80.0 34 82.4 -2 .4 5.1 11 45 81.2 74 81.8 - .6 28.0 14 4 76.3 16 76.6 — .3 3.2 19 5 76.8 35 74.5 +2.3 4.4 21 12 81.3 24 83.1 -1 .8 8.0 22 8 79.5 17 79.7 - .2 5.4 23 15 79.1 61 78.3 + .8 12.0 25 18 80.1 56 80.3 - .2 13.6 26 3 77.3 75 77.7 - .4 2.9 27 13 76.5 72 77.7 -1 .2 11.0 28 16 80.1 25 77.5 +2.6 9.8 33 8 79.3 32 80.7 -1 .4 6.4 34 14 79.6 17 78.7 + .9 7.7 3b 21 74.7 20 74.2 + .5 10.2 38 26 79.7 31 80.3 — .6 14.1

219 638 153.1 Z w . -44.22 Unbiased estimate of difference «. -0.288 cm. 87

TABLE 28

Comparison of the Body Condition Evaluations at Three Months of Age for Animals Heterozygous and. Homozygous a t the F/V Loous

— - (n,xny/ Hetero Homo (ni-**^ Sire ni Ti njj 'Scg ® ^

2 9 4.11 36 4.94 -.83 7.2 10 3 6.00 27 5.70 -.7 0 2.7 11 43 5.12 64 5.17 -.0 5 25.7 14 2 5.00 2 5.50 -.5 0 1.0 19 4 4.75 35 5.03 -.28 3.6 21 11 5.55 19 5.84 -.29 7.0 22 8 b.38 20 4.75 + .63 5.7 23 17 4.76 61 4.67 + .09 13.3 25 17 5.24 56 5.57 -.33 13.0 26 3 5.33 77 5.01 +.32 2.9 27 13 4.85 74 5.05 -.20 11.1 28 16 4.63 25 4.84 -.19 9.8 33 8 5.25 36 5.61 -.3 5 6.5 34 14 5.15 19 5.11 + .04 8.1 35 22 4.86 20 4.70 + .16 10.5 38 26 4.54 31 4.26 +.28 14.1

216 602 142.2 2EtY)D a -11.07 Unbiased estimate of difference a -0.082 condition unit 88

TABIE 29

Comparison of the Inorease in Heart Girths Prom Throe to Six Months of Age in Animals Heterozygous and Homozygous a t F/V Loous

(.n-jxnoJ Hetero Homo (xi-X2J Sire n i *1 n2 x2 D W

om. cm. cm. 2 8 30.87 37 27.45 +3.42 6.57 10 3 25.33 28 22.28 +3.05 2.70 11 45 23.71 71 22.90 + .81 30.14 14 2 26.00 2 24.50 +1.50 1.00 19 4 26.25 35 27.85 -1.60 3.59 21 12 23.83 24 24.37 - .54 8.00 22 8 29 .Ou 19 26.84 +2.16 5.63 23 16 26.18 59 26.54 — .36 12.59 25 18 22.16 53 19.79 +2 *37 13.44 26 6 22 .33 76 23.46 -1.13 5.56 27 13 27.07 75 27.56 - .49 11.08 28 16 20.12 24 21.45 -1,33 9.60 33 8 21.25 36 22.58 -1.33 6.54 34 13 18.92 19 19.36 - .44 7.72 35 22 28.72 20 28.45 + . 27 10.48 38 26 19.07 28 21.03 —1.96 13.48

220 606 149.12 a +25.87 Unbiased estim ate of difference a +0.173 am. 89

TABLE 30

Comparison of the Increase in Heart Girths From Three Months of Age to Three Months After F irst Calving in Animals Heterozygous and Homozygous a t th e F/V Loous

U i i a 2j H etero Homo (x i-x 2) (nj+ngj S ire n l *1 n 2 *2 D vr

cm* om. cm. 2 7 89.0 30 85.1 +3.9 5.67 10 3 83.7 27 84.7 -1 .0 2.70 11 27 81.6 44 79.5 +2.1 16.73 19 3 74.3 28 83.6 -9 .3 2.71 21 11 81.5 19 84.2 -2 .7 6.97 22 5 84.2 11 81.9 +2.3 3.44 23 7 87.0 42 85.7 +1.3 6.00 25 10 81.3 46 87,4 -6 .1 8.21 26 2 91.5 25 90.0 +1.5 1.85 27 10 83.4 43 84.8 -1 .4 8,11 28 12 89.2 18 88.7 + .5 7.20 33 4 79.3 12 83.3 -3 .5 3.00 34 9 83.3 14 83.0 + .3 5.48 35 14 85.0 14 87.5 —2.5 7.0 38 18 91.8 22 94.8 -3 .0 9.90

142 395 93.20 Z tcd . -84.88 Unbiased e stimate of difference » —0.910 cm. 90

TABIE 31

Comparison of the Services Per Conception for Animals Heterozygous and Homozygous a t th e F/V Locus

(nixn2) Hetero Homo ( xi 2 ) (nl+n2) S ire n i 5 l n 2 *2 D W

2 17 2.176 51 1.764 +.412 12.75 10 5 1.400 32 1.687 -.2 87 4.33 11 24 2.333 45 2.000 +.333 15.65 14 3 1.000 14 1.571 -.571 2.47 19 3 1.333 24 1.375 -.042 2.67 21 8 2.125 19 1.842 + .283 5.63 22 4 1.750 11 1.454 +.296 2.93 23 9 1.222 40 1.525 -.303 7.35 25 8 2.375 50 2.580 -.2 0 5 6.89 27 9 1.111 40 2.050 -.939 7.35 28 14 2.500 15 2.600 -.100 7.24 33 4 1.750 19 2.157 -.4 0 7 3.30 34 12 2.250 20 2.450 -.2 00 7.50 35 14 1.642 13 1.307 +.335 6.74 38 16 2.250 19 2.210 + .040 8.69

150 _ 412 101.39 ZW a -1 .3 7 Unbiased estimate of difference a -0.014 service per conception 91

TABLB 32

Comparison of the Heart Girths of the First Offspring of Animals Heterozygous and. Homozygous a t the F/V Locus

tm z n s) Hetero Homo (5?L-x2) (nj+ngJ S ire n i x i * 2 ■3T D W

om. om. cm. 2 6 76*50 27 75.33 +1.17 4.90 10 2 79.00 19 79.95 - .95 1.81 11 12 78.92 23 80.17 -1 .2 5 7.67 21 8 79.63 9 77.89 +1.74 4.24 23 3 77.67 22 76.18 +1.49 2.64 25 5 79.60 21 78.62 + .98 4.04 27 4 73.00 20 74.50 -1.50 3.33 28 4 78.50 8 77.13 +1.37 2.67 33 2 79.00 8 79.50 - .50 1.60 34 9 75.33 7 74.14 +1.19 3.94 35 7 73.57 7 73.71 - .14 3.50 38 4 78.00 6 78.17 - .17 2.40

66 177 42.74 £ wd * +11.35 Unbiased estimate of difference ■ +0.265 cm. 92

TABIE 33

Comparison of the Milk Production of Animals Heterozygous and Homozygous at the J Locus

(nixnjJ Hetero Homo (T1-X2) (n i+n2; Sire n i $L n2 *2 D W

Lbs. Lbs. Lb3. 2 14 13041 14 13605 - 464 7.00 10 7 12627 12 13000 - 382 4.42 11 7 13836 43 12913 + 923 6.02 18 12 12545 24 12035 + 510 8.00 19 7 13604 11 11944 +1660 4.28 21 2 15235 29 12976 +2259 1.87 24 14 14468 51 13990 + 478 10.98 25 11 10755 41 10905 - 150 8.67 26 9 11326 23 11449 - 123 f 6.47 28 5 12296 16 10763 +1533 3.81 33 4 12325 7 10314 +2011 2.55 34 10 11045 23 11829 - 784 6.97 35 5 11868 9 13529 -1661 3.21 38 7 12319 15 11347 + 972 4.77 39 12 12742 38 13037 - 295 9.12

126 356 88.14 £m> = +21376 Unbiased estimate of difference = +232.69 lbs. 93

TAB IE 34

Comparison of the Butterfat Production of Animals Heterozygous and Homozygous at the J Loous

(nixnaJ H etero Homo (X1 -X2 ) (ni+ n2; S ire n i x i n 2 5 T D W

Lbs. Lbs. Lbs. 2 14 438 14 441 - 3 7.00 10 7 427 10 447 -20 4.42 11 7 520 43 483 +37 6.02 18 12 450 24 432 +18 8.00 19 7 480 11 428 ♦52 4.28 21 2 515 29 452 +63 1.87 24 14 526 51 504 +22 10.98 25 11 372 41 375 - 3 8.67 26 9 420 23 412 + 8 6.47 28 5 410 16 368 +42 3.81 33 4 438 7 387 +51 2.55 34 10 377 23 383 - 6 6.97 35 5 394 9 477 -83 3.21 38 7 511 15 470 +41 4.77 39 12 498 38 501 - 3 9.12

126 356 88.14 ZlW a +1015.05 Unbiased estim ate of difference a +11.52 lbs. 94

TABLE 35

Comparison of tho Maturity Indexes of Animals Heterozygous and Homozygous a t th e J Loous

(nTxnrJ Hetero Homo (^L-xg) (n!+n2J S ire nl *1 n 2 x2 D W

2 10 87.6 9 99.7 —l

70 —, 130 46.3 Z w = -31.55 Unbiased estim ate of difference ■ -0.68 95

TABLE 36

Comparison of the Heart Girth Measurements at Birth of Animals Heterozygous and Homozygous at the J Loous

ln-|Xn9) Hetero Homo (n'i+n2i S ire n2 D W

om* cm. cm. 2 8 76.9 15 77.1 - .2 5.22 8 2 80.5 8 80.4 + .1 1.60 10 7 81.1 12 80.4 + .7 4.42 11 16 82.5 70 81.2 +1.3 13.02 18 20 81.1 41 79.6 +1.5 13.44 19 10 75.2 12 75.8 - .6 5.45 21 4 81.0 32 82.7 -1 .7 3.56 23 5 78.2 29 78.4 - .2 4.26 24 20 81.8 115 81.1 + .7 17.04 25 14 80.5 45 80.1 + .4 10.68 26 20 78.3 58 77.5 + .8 14.87 27 10 78.0 26 77.1 + .9 7.22 28 7 77.1 19 80.0 -2 .9 5.12 33 8 78.1 16 80.2 -2 .1 5.33 34 10 81.6 21 77.9 +3.7 6.77 35 8 73.8 15 74.7 - .9 5.22 38 12 81.3 23 79.6 +1.7 7.88 39 14 79.8 46 80.0 - .2 10.73

195 ~ 603 141.83 ZrWD - +69.3 Unbiased estimate of difference * +0.588 cm. 96

TABLE 37

Comparison of the Body Condition Evaluations at Three Months of Age of Animals Heterozygous and HomozygouB a t th e J Locus

U tXHo) H etero Homo (Y1-X2J (n i+n 2) S ire n i n 2 *2 D W

2 6 4,83 14 4.79 + .04 4 .2 10 6 5.67 11 5.73 - .06 3.88 11 15 5.20 62 5.21 - .01 12.08 18 21 5.19 40 4.78 + .41 13.77 19 9 5.22 11 4.73 + .49 4.95 21 4 5.76 26 5.73 + .02 3.47 23 5 3.60 30 4.63 -1 .0 3 4.29 24 19 5.58 110 5.40 + .18 16.20 25 14 5.43 45 6.58 - .15 10.68 26 21 5.24 59 4.95 + .29 15.49 27 10 4.30 26 4.65 - .35 7.22 28 7 4.43 19 4.84 - .41 5.11 33 8 5.63 17 5.41 + .22 5.44 34 11 5.09 22 5.14 - .05 7.33 35 8 4.76 15 4.86 - .11 5.22 38 12 4.33 23 4.52 - .19 7.89 39 15 4.00 46 4.46 - .46 11.31

191 „ 576 138.50 = -1.69 Unbiased estimate of difference a -0.012 97

TAB1E 38

Comparison of the Increase in Heart Girth Prom Three to Six Months of Age in Animals Heterozygous and Homozygous a t the J Locus

(ni xa 0) Hetero Homo (xi-xfcj (ni+ngj S ire n f ------W »2 “ ' ' "2 D W

om. cm. om. 2 7 29.85 13 29.69 + .16 4.55 10 6 23.00 12 22.33 + .67 4.00 11 15 23.73 69 23.11 + .62 12.32 18 21 24.42 39 22.48 +1.94 13.65 19 9 28.11 11 25.90 +2.21 4.95 21 4 23.00 32 24.34 ~ .34 3.56 23 5 27.80 29 26.79 +1.01 4.26 24 20 22.90 113 23.26 - .36 16.99 25 12 19.75 44 20.09 - .34 9.43 26 21 23.14 58 23.51 - .37 15.42 27 10 29.90 27 27.40 +2.50 7.30 28 7 21.57 19 20.10 +1.47 5.11 33 8 22.75 17 22.29 + .46 5.44 34 11 20.09 21 18.71 +1.38 7.22 35 8 28.87 15 28.80 + .07 5.22 38 11 20.27 23 19.21 +1.06 7.44 39 12 19.41 44 20.31 - .90 9.43

187 586 136.20 * +74.54 Unbiased estimate of difference a +0.547 om. 98

TABUS 59

Comparison of* the Increase in Heart Girths Prom Three Months of Age to Three Months After First Calving in Animals Heterozygous and Homozygous a t th e J Locus

U ixnjj) Hetero Homo (T i-xgj (nl+ngj S ire n i x i n2 D W

cm. om. cm. 2 6 88.7 13 85.8 +2.9 4.11 10 7 90.3 11 81.8 +8.5 4.28 11 6 79.2 44 80.9 -1 .7 5.28 18 10 85.1 14 84.2 + .9 5.83 19 7 85.6 11 79.8 +5.8 4.28 21 2 80. 0 28 83.4 -3 .4 1.87 24 7 81.6 61 82.4 - .8 6.28 25 11 87.8 36 87.3 + .5 8.43 26 9 90.4 18 89.9 + .5 6.00 27 4 86.0 11 85.7 + .3 2.93 28 5 94.4 15 89.3 +5.1 3.75 33 4 80.0 5 83.4 -3 .4 2.22 34 7 84.8 16 82.4 +2.4 4.87 35 5 87.4 10 84.4 +3.0 3.33 38 7 93.1 16 94.1 -1 .0 4.87 39 14 88.0 37 89.2 -1 .2 10.16

111 346 78.49 £.m> . +82.30 Unbiased estimate of difference » +1.05 cm. 99

TABIS 40

Comparison of the Services For Conception for Animals Heterozygous and Homozygous a t the J Loous

T n ix n 2J H etero Homo (Yl-X2> S ire »1 *1 n 2 w D

2 7 1,571 12 1.750 - .179 4.42 10 7 1.714 11 1.545 + .169 4.28 11 8 3.166 42 2.119 ♦1.047 6.72 18 12 2.583 22 3.136 - .553 7.76 19 7 1*857 10 1.300 ♦ .557 4.12 21 2 2.000 25 1.920 + .080 1.85 24 14 1.714 45 1.755 — .041 10.68 25 11 1.545 41 3.024 -1.479 8.67 26 9 2.555 17 2.235 ♦ .320 5.88 27 2 3.000 10 1.600 +1.400 1.67 28 5 4.200 15 2.333 +1.867 3.75 33 4 2.000 6 1.833 + .167 2.40 34 9 3.222 34 2.043 +1.179 7.12 35 5 1.400 9 1.222 + .178 3.21 38 7 3.000 11 1.818 +1.182 4.28 39 11 2.363 39 2.666 - .303 8.58

120 ~ 349 85.39 £m > * +11 • 02 Unbiased estimate of difference = +0*129 services 100

TABLE 41

Comparison of Heart Girths of the F irst Offspring of Animals Heterozygous and Homozygous at the J Locus

w H etero Homo {x i-x z )

om. om. cm. 2 3 74.67 7 76.14 -1.47 2.10 10 5 78.00 6 79.83 -1.83 2.73 11 4 83.00 21 79.66 +3.34 3.36 18 3 76.33 10 78.90 -2 .5 7 2.31 19 2 75.00 7 72.86 +2.14 1.56 23 2 78.50 5 79.80 -1 .3 0 1.43 24 7 80.43 25 79.16 +1.27 5.47 25 4 76.25 17 79.18 -2.93 3.24 26 2 82.00 14 78.29 +3.71 1.75 28 2 80.60 6 77.83 +2.67 1.50 33 2 77.50 2 80.50 -3.00 1.00 34 5 75.20 11 74.64 + .56 3.44 35 3 73.33 5 74.40 -1 .0 7 1.88 39 _7 80.57 13 78.62 +1.95 4.55

51 149 45.51 S w t* +12.42 Unbiased estimate d iffe re n c e S3 +0 1.27 cm. 101

TABIE 42

Comparison of the Milk Production of Animals Heterozygous and Homozygous a t the L Loous

U ix n 9; He ta ro Homo (*L-*2) (ni+ n2) S ire n i *1 n2 *2 D W

Lbs* Lbs. Lbs. 2 5 13884 57 13318 + 566 4.60 3 2 9250 39 9678 - 428 1.90 8 3 10686 13 10808 - 122 2.44 10 10 13368 24 12780 + 588 7.06 11 9 13072 60 12930 + 142 7.83 12 7 10836 27 10150 + 686 5.56 18 8 11249 21 12747 -1498 5.79 21 6 13678 11 12840 + 838 3.88 23 11 13532 21 14140 - 598 7.21 24 9 14S62 56 14001 + 661 7.75 25 28 11066 24 10560 + 506 12.92 26 3 10193 25 11351 -1158 2.68 27 2 11970 39 13338 -1368 1.90 28 4 11518 27 10569 + 949 3.48 33 3 11273 21 11291 - 18 2.63 54 7 12070 13 1119 5 + 875 4.55 38 9 11271 30 11983 - 712 6.92 39 15 13475 16 12145 +1330 7.74

141 524 96.84 2 . TO) * +20529.7 Unbiased estim ate of difference = +211.99 lbs. 102

TABIE 43

Comparison of the Butterfat Production of Animals Heterozygous and Homozygous a t th e L Loous

ln-|xn?; H etero Homo (ni+ n2) S ire nl n 2 *2 D W

Lbs* Lbs* Lbs* 2 5 480 57 442 +38 4.60 3 2 335 39 351 -16 1.90 8 3 383 13 375 + 8 2.44 10 10 445 24 435 +10 7.06 11 9 488 59 485 + 3 7.83 12 7 373 27 351 +22 5.56 18 8 395 21 461 -66 5.79 21 6 475 11 445 ♦30 3.88 23 11 506 21 507 - 1 7.21 24 9 624 56 506 ♦18 7.75 25 28 381 24 367 +14 12.92 26 3 360 25 412 -52 2.68 27 2 460 39 516 —56 1.90 28 4 418 27 360 +58 3.48 33 3 410 21 398 ♦12 2.63 34 7 383 13 373 -10 4.55 38 9 476 30 490 -14 6.92 39 15 513 16 471 +42 7.74

141 « 523 96.84 2* wd « +694.46 Unbiased estimate of difference » +7.17 l b s . 103

TAB IB 44

Comparison of the Maturity Indexes of Animals Heterozygous and Homozygous at the L Loous

^ x n * ; He ta ro Homo (t i -* 2; (n 1+n 2; S ire n l *L n2 ■*2 D W

2 2 110.5 38 88.7 ♦21.8 1.90 3 2 102.0 25 98.2 ♦ 3.8 1.85 8 2 132.0 8 109.0 +23.0 1.60 10 7 85.9 16 96.4 -10.5 4.77 11 3 92.7 44 97.8 - 5.1 2.81 IE 4 107.8 18 98.3 ♦ 9.5 3.27 18 4 95.3 12 104.6 - 9.3 3.00 21 3 94.0 6 88.7 + 5.3 2.00 23 4 87.3 12 79.9 ♦ 7.4 3.00 24 7 90.4 24 98.8 - 8.4 5.42 25 18 101.9 25 99.7 ♦ 2.2 10.47 27 2 100.0 25 98.7 ♦ 1.3 1.85 33 2 105.5 15 103.1 + 2.4 1.76 34 4 102.3 7 104.1 - 1.8 2.56 38 3 101.3 8 91.9 + 9.4 2.18 39 _4 95.5 6 98.2 - 1.71 2.40

71 288 50.83 cLiW a ♦52.76 Unbiased estimate of difference = +1.04 104

TABIE 45

Comparison of the Heart Girth Measurements at Birth of Animals Heterozygous and Homozygous a t the L Locus

(nixn2) Hetero Homo (2i-352) (ni+n2) S ire nl *1 ng.. _____ x2 D W om* om. cm. 2 4 77.5 44 77.2 + .3 3.66 3 2 76.0 40 78.5 -2 .5 1.90 8 4 80.0 15 78.7 4*1. 3 3.16 10 11 81.5 23 81.9 — .4 7.44 11 16 81.5 100 81.5 13.79 12 7 75.1 29 78.8 -3 .7 5.64 18 16 79.1 39 80.5 -1 .4 11.34 21 8 82.4 12 83.7 -1 .3 4.80 22 2 78.5 15 79.8 —1.3 1.76 23 22 77.8 28 79.3 -1 .5 12.32 24 22 80.9 113 81.2 - .3 18.41 25 31 80.3 28 80.1 + .2 14.71 26 14 77.2 56 78.2 -1 .0 13.07 27 3 77.3 71 77.5 - .2 2.88 28 6 75.0 35 79.1 -4 .1 5.12 33 3 82.3 31 80.5 +1.8 2.74 34 7 77.0 12 80.3 —3.3 4.42 38 15 80.6 42 79.8 + .8 11.05 39 20 79.9 18 80.8 - .9 9.47

213 751 147.70 ZsYJD a -112.82 Unbiased estim ate of difference a -0 .7 6 cm. 105

TABIE 46

Comparison of the Body Condition Evaluations at Three Months of Age of Animals Heterozygous and Homozygous at the L Loous

{n im z ) Hetero Homo ( X1-X2) (n]+n2) S ire n l x l *2 *2 DW

2 4 5.00 36 4.72 +.28 3.60 8 4 4.75 15 5.13 -.3 8 3.15 10 11 5.64 19 5.63 +.01 6.97 11 15 5.20 92 5.14 + .06 12.90 12 4 6.25 22 5.32 +.93 3.38 18 16 4.56 39 5.05 -.49 11.34 21 8 5.63 10 5.60 +.13 4.44 22 2 5.00 17 4.82 + .18 1.79 23 22 4.36 29 4.72 -.3 6 12.51 24 22 5.45 107 5.42 +.03 18.24 25 31 5.65 28 5.50 +.15 14.71 26 14 4.71 58 5.12 -.4 1 13.10 27 3 5.33 72 4.93 +.40 2.88 28 6 4.50 35 4.80 -.3 0 5.12 33 3 5.33 34 5.62 -.29 2.75 34 7 5.14 13 5.23 -.0 9 4.55 38 15 4.53 42 4.33 + .20 11.05 39 20 4.45 19 4.47 -.02 9.74

194 687 142.22 -7.35 Unbiased estimate of difference = -0.05 106

TABLE 47

Comparison of the Increase in Heart Girths From Three to Six Months of Age in Animals Heterozygous and Homozygous a t th e L Locus

H etero Homo (x i-x z ) (nx+n2J S ire n l xx a 2 x2 D W

cm* am. cm. 2 4 28.50 36 28.30 + .20 3.60 8 4 17.00 14 20.07 -3.07 3.11 10 11 22.00 20 22.90 - ,90 7.10 11 15 24,86 99 23.02 ♦1.84 13.03 12 4 16.50 19 20.78 -4.28 3.30 18 16 24.56 39 22.82 +1.74 11.35 21 8 24.50 12 24.91 — .41 4.80 22 2 30.00 16 28.12 +1.88 1.78 23 22 26.40 27 27.37 - .97 1.21 24 21 23.71 112 23.11 + .60 17.68 25 31 19.90 25 20.00 - .10 13.84 26 14 24.28 57 23.01 +1.27 13.08 27 3 27.66 73 27.38 + .28 2.88 28 6 21.83 34 20.76 +1.07 5.10 33 3 22.33 34 22.55 - .22 2.76 34 6 19.00 13 19.53 - .53 4.11 38 6 19.00 39 20.51 -1.51 5.20 39 18 19.11 17 21.00 -1.89 8.74

194 686 122.67 S i© * +19 • 53 Unbiased estimate o f d iffe re n c e =>+0.159 cm. 107

TAB IE 48

Comparison of the In or ease in Heart Girths From Three Months of Age to Three Monthb After Calving in Animals Heterozygous and Homozygous a t th e L Loous

H etero Homo (ni+n.2 ) S ire n i XL n2 x 2 D W

cm* cm. cm. 2 4 85*25 29 86.34 -1.09 3.51 8 3 83.00 13 87.38 -4.38 2.44 10 9 83*22 20 85.50 -2.28 6.21 11 8 80*75 61 80.56 + .19 7.07 12 4 82.00 18 85.61 -3.61 3.27 18 4 80.00 14 86.36 -6 .3 6 3.11 21 6 80*33 10 83.70 -3.37 3.75 23 12 85.50 18 84.72 + .78 7.20 24 10 82.10 67 82.27 - .17 8.70 25 25 85.56 22 89.60 —4.04 11.70 26 2 96.00 21 89.57 +6.43 1.83 27 2 82.50 42 85.26 -2 .7 6 1.90 28 4 87.25 26 89.12 -1 .8 7 3.47 33 2 81.00 13 82.38 -1 .3 8 1.73 34 4 79.00 9 81.11 -2 .1 1 2.77 38 9 90.88 7 94.19 -3.31 3.94 39 16 88.81 16 87.19 +1. 62 8.00

124 — 406 80.60 ^Y!D a -122.96 Unbiased estimate of difference = -1.53 cm. 108

TABUS 49

Comparison of the Services Per Conception for Animals H eterozygous and Homozygous a t th e L Locus

tn-ixn.,) H etero Homo (ni+ n2J S ire n i T l n2 x2 D W

2 4 2.00 38 1.58 + .42 3.62 3 2 1.50 31 2.74 -1 .2 4 1.88 8 3 3.00 13 2.46 + .54 2.44 10 10 1.80 21 1.57 + . 23 6.77 11 8 1.13 58 2.17 -1 .0 4 7.03 12 7 3.00 27 3.00 5.56 18 7 2.71 21 3.19 — .48 5.25 21 3 2.67 11 2.00 ♦ .67 2.36 23 9 2.22 20 1.45 + .77 6.21 24 9 1.78 50 1.74 + .04 7.63 25 28 2.96 24 2.04 + .92 12.92 26 21 2.00 20 2.40 - .40 10.24 27 2 2.00 30 1.89 + .11 1.90 28 3 2.33 26 2.58 - .25 2.69 33 3 1.33 20 2.20 - .90 2.61 34 7 2.43 13 2.69 — .26 4.55 38 9 2.89 26 2.00 + .89 6.69 39 14 2.14 16 3.13 - .99 7.47

141 _ 473 97.82 •1-1.254 Unbiased estimate of difference a +0*012 service 109

TABLE 50

Comparison of the Heart Girths of the First Offspring of Animals Heterozygous and Homozygous a t th e L Loous

H etero Homo ( * i- x 2; (ni+ngj S ire n i x i *2 X2 D W

om* cm. om. 2 3 72.00 19 76.32 -4.32 2.59 10 8 80.63 11 79.27 +1.36 4.63 11 4 80.75 29 79.55 +1.20 3.52 12 3 78.67 11 79.09 - .42 2.36 18 3 78.67 6 79.33 — . 66 2.00 21 5 77.80 6 77.17 + .63 2.73 23 7 75.43 8 77.25 -1.82 3.73 24 7 80.14 25 79.24 + .90 5.47 25 13 77.85 7 80.29 -2.44 4.55 27 21 76.50 16 74.00 +2.50 9.08 28 3 74.33 9 78.67 —4.34 2.25 34 3 74.33 7 75.57 -1 .2 3 2.10 38 78.67 7 77.86 + .81 2.10

83 141 47.11 £ w = -2.22 Unbiased estimate of difference a -0 .0 5 om.

7 110

TAB IS 51

Comparison of the Milk Production of Animals Heterozygous and Homozygous a t the SU Loous

l n , ® 0 Hetero Homo S ire n i 95“ B2 D

Lbs* Lbs* Lbs* 5 5 9404 36 9855 - 451 4.39 8 6 9405 21 11160 -1755 4*66 10 2 13740 36 12899 + 841 1.89 11 42 13007 2 12060 + 947 1*91 12 10 10648 25 10036 + 612 7.14 14 4 13768 11 14037 - 269 2*93 19 3 11697 25 12685 - 988 2.68 21 5 13694 26 13012 + 682 4*19 23 3 12643 47 13770 -1127 2.82 24 9 13280 56 14224 - 944 7.75 25 25 10847 19 11093 - 246 10.80 26 2 13675 30 11264 •*■2411 1.88 27 31 13391 18 12806 ♦ 585 11.39 28 4 10890 27 10663 + 227 3.48 33 3 12580 21 11104 ♦1476 2.63 35 2 14340 25 13255 +1085 1.85 38 7 11531 32 11881 - 350 5.74 39 7 12816 44 13053 - 237 6.04

186 566 97.00 * -298*46 Unbiased estimate of difference a -3 .0 7 lb s . I l l

TABIE 52

Comparison of the Butterfat Production of Animals Heterozygous and Homozygous a t the SU Loous

Hetero Homo (x i-x g ) (ni+n25 S ire »1 T l *2 *2 D W

Lbs* Lbs. Lbs* 2 16 457 65 457 0 12.84 3 5 344 36 357 -13 4.39 8 6 325 21 389 -64 4.66 10 2 420 36 440 -20 1.89 11 41 487 2 460 ♦27 1.91 12 10 364 25 354 +10 7.14 14 4 460 11 445 +15 2.93 19 3 423 25 458 -35 2.68 21 5 480 26 451 +29 4.19 23 3 427 47 501 -74 2.82 24 9 492 56 512 -20 7.75 25 25 372 19 397 -25 10.80 26 2 485 30 410 +75 1.88 27 31 516 18 494 ♦22 11.39 28 4 345 27 371 -26 3.48 33 3 453 21 392 ♦61 2.63 35 2 535 25 457 +78 1.85 38 7 473 32 489 -16 5.74 39 __ 7 483 44 507 -24 6.04

185 566 97.00 Z.v© »-463*13 Unbiased estimate of difference = -4*77 lbs* 112

TABUS 5 3

Comparison of the Maturity Indexes of Animals Heterozygous and Homozygous a t the SIT Loous

U l* n 2J Hetero ....Homo (Ti-XgJ (ni«i2J S ire n i xi n2 D W

2 11 85.18 39 87.56 - 2.38 8.58 3 4 91.75 22 99.32 - 7.57 3.38 8 4 115.00 10 105.60 + 9.40 2.86 10 2 94.50 25 92.84 + 1.66 1.85 12 6 100.50 16 99.88 + .62 4.36 14 3 92.33 7 90.57 + 1.76 2.10 21 4 75.55 14 96.14 -20.59 3.11 24 4 93.26 27 97.41 - 4.16 2.58 25 21 102.42 14 95.86 + 6.56 8.40 27 22 96.30 12 98.76 - 2.45 7.76 33 3 102.00 14 103.64 - 1.64 2.47 38 3 100.33 8 92.25 + 8.08 3.63 39 _3 92.33 20 95.95 - 3.62 2.61

90 228 53.69 ^ 7 © a -32.50 Unbiased estimatei of difference a -0.605 113

TABLE 54

Comparison of the Heart Girth Measurements at Birth of Animals Heterozygous and Homozygous a t the STJ Loous

(ai3m97 He te r o Homo (TL-*2J (n^+ngj Sire n i x i *2 x i“ D W

om* om. om. 2 10 77.40 47 77.27 + .13 8.24 8 6 80.00 22 79.32 + .68 4.71 10 2 81.00 37 82.08 -1.08 1.90 11 75 82*05 2 81.50 + .55 1.96 12 10 79.20 27 77.81 +1.39 7.30 14 4 76.25 12 77.33 -1.08 3.00 18 4 82.00 60 80.00 +2.00 3.75 19 3 76.00 37 74.65 +1.35 2.78 21 5 80.20 31 82.87 -2.67 4.31 23 4 80.50 69 78.36 +2.14 3.78 24 20 80.80 115 81.24 - .44 17.04 25 29 80.28 20 80.00 + .28 11.84 26 3 81.67 75 77.55 +4.12 2.88 27 50 76.68 31 78.55 -1.87 19.14 28 5 79.00 36 78.47 + .53 4.39 33 9 78.44 31 80.94 -2.50 6.98 35 3 75.66 37 74.46 +1.20 2.78 38 11 80.73 46 79.87 + .86 8.88 39 11 79.27 50 80.12 - .85 9.01

264 785 124.65 2?m> = -1.88 Unbiased estimate of difference > —0.015 om. 114

TABIE 55

Comparison of Body Condition Evaluations at Three Months of Age of Animals Heterozygous and Homozygous a t the SU Loous

^n-j 300;,) H etero Homo (x i-* 2 ) (n]_+n2) S ire n i n2 *2 D W

2 8 5.00 37 4.73 + .27 6.58 8 6 4.67 22 5.23 -.6 6 4.71 10 2 6.00 27 5.63 + .37 1.86 11 73 5.15 2 5.50 -.3 5 1.95 12 7 5.71 19 5.37 +.34 5.12 14 2 5.00 2 5.50 -.50 1.00 18 2 4.50 60 4.95 — .45 1.94 19 3 5.00 35 5.00 00 2.76 21 5 5.60 25 5.76 -.1 6 4.17 23 4 4.75 70 4.66 + .09 3.78 24 19 5.26 110 5.45 -.1 9 16.20 25 29 5.52 20 5.60 -.0 8 11.84 26 3 4.67 77 5.04 -.3 7 2.89 27 50 5.16 32 4.81 +.35 19.51 28 5 4.80 36 4.75 + .05 4.39 33 9 5.77 35 5.49 +.28 7.16 35 4 5.26 37 4.76 +.49 3.61 38 11 4.82 46 4.28 + .54 8.88 39 11 4.27 51 4.37 -.1 0 9.05

253 743 115.35 8 +8.81 Unbiased estimate of difference « +0.076 - 115

TABLE 56

Comparison of the Inoreaee in the Hoart Girths From Three to Six MonthB of Age in Animals Heterozygous and Homozygous a t the SU Locus

Im xn,,) H etero Homo (TL-r2; (n-j+ng) S ire »1 *L n 2 *2 D W

cm. cm. cm. 2 6 27.33 38 28.28 - .95 5.18 8 6 18.00 21 19.52 -1.5 2 4.67 10 2 22.00 28 22.71 - .71 1.87 11 74 23.18 2 21.50 +1.68 1.95 12 6 20.16 17 20.00 + .16 4.43 14 2 29.00 2 21.50 +7.50 1.00 19 3 30.00 35 27.31 +2.69 2.76 21 5 25.40 31 24.00 +1.40 4.31 23 4 24.75 67 26.43 -1.68 3.77 24 19 23.63 114 23.14 + .49 16.29 25 27 20.25 19 20.36 — .11 11.15 26 3 25.66 76 23.32 +2.34 2.89 27 50 27.24 33 27.48 - .24 19.88 28 5 17.20 35 21.45 -4.2 5 4.38 33 9 23.00 35 22.17 + .83 7.16 35 4 30.00 37 28.27 +1.73 3 . 61 38 10 20.90 44 19.90 +1.00 8.15 39 10 20.20 47 20.27 - .07 8.25

245 681 111.70 uWD a +15.16 Unbiased estim ate of difference » +0.135 cm. 116

TAB1E 57

Comparison of the Increase in Heart Girths From Three Months of Age to Three Months After Calving in Animals Heterozygous and Homozygous a t th e SU Loous

(nTxajJ Hetero Homo (*1-X2> (n!+n2) S ire ■ n i ------‘SET *2 ..■‘5 T D W

om. cm. cm. 2 6 85.83 31 85.84 - .01 5.03 8 6 83.33 19 88.05 -4.72 4.56 10 2 92.50 27 84.00 +8.50 1.86 11 42 81.26 2 81.00 + .26 1.91 12 6 85.67 16 84.69 + .98 4.36 14 2 70.00 19 3 87.00 27 82.04 +4.96 2.70 21 5 82.60 25 83.32 - .72 4.17 23 3 84.33 43 86.14 -1.81 2.80 24 11 83.82 66 81.98 +1.84 9.43 25 22 87.64 18 87.77 - .13 9.90 27 31 84.84 20 85.75 +1.09 12.16 28 4 87.50 26 89.08 -1.58 3.47 33 4 80.75 12 82.92 -2.17 3i00 35 3 86.00 25 86.26 - .26 2.68 38 8 92.25 32 93.75 -1.50 6.40 39 7 89.29 45 89.07 + . 22 6.06

165 434 80.58 Z w a +12.69 Unbiased estimate of difference = +0.157 cm. 117

TABLE 68

Comparison of the Services Par Conception for Animals Heterozygous and Homozygous at the SU Locus

Hetero Homo (ni+n2J S ire n i *1 nz x2 D W

2 12 1.08 47 2.00 - .92 9.72 3 3 3.33 30 2.46 + .87 2.73 8 6 3.00 21 2.24 + .76 4.67 11 40 2.18 21 1.50 + .68 1.90 12 10 2.90 24 3.04 " .14 7.06 14 4 1.00 10 1.70 - .70 2.86 19 3 1.67 24 1.33 + .34 2.67 21 4 1.25 23 2.04 - .79 3.41 23 3 1.33 43 1.51 - .18 2.80 24 8 1.75 51 1.7b .00 6.90 25 25 2.88 19 2.58 + * 30 10.79 26 2 3.00 24 2.29 + .71 1.85 27 30 1.83 17 1.65 + .18 10.85 28 4 4.25 25 2.28 +1.97 3.44 33 3 2.33 20 2*06 + .28 2.61 35 2 1.50 25 1.48 + .02 1.85 38 8 1.50 27 2.44 - .94 6.17 39 __ 7 2.86 44 2.52 + .34 6.04

174 _ 476 81.42 Z tvd » +3.29 Unbiased estimate of difference a +0.04 service 118

TAB IE 59

Comparison of tha Heart Girths of the First Offspring of Animals Heterozygous and Homozygous at the SU Loous

lm Hetero Homo (x±-X2) (ni+n2} S ire * 1 35T n2 *2 D W

am. am. cm. 2 6 73.17 21 75.95 -2.78 4.67 8 3 77.6? 9 78.89 -1.22 2.25 11 17 79.35 2 78.50 + .85 1.79 12 6 80.33 8 78.00 +2.33 3.43 14 2 72.50 4 76.25 -3.75 1.33 19 2 71.50 9 72.56 -1.06 1.64 21 5 76.60 12 79.58 —2 .98 3.53 24 6 79.67 26 79.38 + .29 4.88 25 9 79.78 7 77.71 +2.07 3.94 27 13 74.38 11 74.09 + .29 5.96 38 2 79.00 8 77.88 +1.12 1.60 39 __2 8 0 , d0 19 78.95 +1.55 1.81

73 136 35.19 ZW a -1.82 Unbiased estimate of difference a -0.0b om. 119

TAB1E 60

Comparison of the Milk Production of Animals Heterozygous and Homozygous at the Z Loous

H etero Homo (xx-x2) O i+ n g ; S ire n i *1 &2 ^2 D w

Lbs* Lbs* Lbs* 2 14 12865 18 13541 - 676 7.876 3 13 9404 15 9719 - 315 6.964 8 7 10479 20 10873 - 394 5.135 10 5 13U52 10 13082 - 30 3.333 11 31 12895 41 12960 - 65 17.652 12 6 10167 29 10220 - 53 4.971 14 11 13909 8 13905 + 4 4.632 18 7 10741 30 12530 -1789 5.675 19 b 10806 7 13131 -2325 2.917 21 6 13307 8 12991 + 316 3.428 22 6 12992 2 14245 -1252 1.500 23 13 12694 10 14724 -2030 5.652 24 24 14733 41 13719 >1014 17.723 25 12 12103 21 10522 +1581 7.636 26 5 11206 27 11453 - 247 4.218 27 6 12188 14 13548 - 360 4.200 28 8 9604 22 10909 -1305 5.876 34 4 13703 15 11703 +2000 3.158 38 11 11970 16 11303 + 367 6.519 39 15 13242 20 12815 + 427 8.571

209 374 127.635 2 m - -8105.96 Unbiased estim ate of differenoe = -63*48 lbs. TAB IE 61

Comparison of the Butterfat Production of Animals Heterozygous and Homozygous at the Z Locus

( n ^ H etero Homo (xi-X2) (ni+ n2j S ire n i x i n 2 *2 D W

Lbs. Lbs* Lbs* 2 14 421 18 452 -31 7.875 3 13 344 15 357 -13 6.964 8 7 364 20 378 -14 5.185 10 b 474 10 443 +31 3.333 11 30 480 41 488 * 8 17.652 12 6 356 29 357 - 1 4.971 14 11 455 8 440 +15 4.632 18 7 381 30 450 -69 5.675 19 5 404 7 464 -60 2.917 21 6 473 8 449 +24 3.428 22 6 490 2 495 - 5 1.500 23 13 468 10 520 -52 5.652 24 24 524 41 500 +24 17.723 25 12 405 21 365 +40 7.636 26 5 408 27 416 - 8 4.218 27 6 470 14 521 -51 4.200 28 8 328 22 377 -49 5.876 34 4 438 15 386 +52 3.158 38 11 493 16 466 +27 6.519 39 15 521 20 496 +25 8.571

209 _ 374 127.655 = -490.99 Unbiased estimate of difference = —3*845 lbs* 121

TABLE! 62

Comparison of the Maturity Inderos of Animals Heterozygous and Homozygous a t the Z Loous

(nixn2J H etero Homo (n i* n 2J S ire »1 *2 w DW

2 8 91.37 14 87.64 + 3.73 5.09 3 7 109.57 10 96.10 +13.47 4.12 8 3 116.67 11 106.00 +10.67 2.36 10 4 82.50 8 95.13 -12.63 2.67 11 25 95.20 24 101.42 - 6.22 12.22 12 3 98.00 19 100.37 - 2.37 2.59 14 8 96.75 4 93.25 + 3.50 2.67 18 4 99.50 18 94.89 + 4.61 3.27 19 2 133.30 3 78.33 +54.97 1.20 21 2 85.00 4 88.25 - 3.25 1.33 22 2 80.00 23 5 81.00 7 77.14 + 3.86 2.92 24 14 95.21 17 98.24 - 3.03 7.68 25 11 95.13 15 96.53 - 1.35 6.35 26 2 93.00 12 91.66 + 1.34 1.71 27 3 94.33 10 90.40 + 3.93 2.31 33 13 79.62 34 3 102.67 7 107.86 - 5.19 2.10 35 4 102.50 38 3 95.33 4: 88.00 + 7.33 1.71 39 5 89.60 10 97.30 - 7.70 3.33

112 __ 192 65.63 2LWD = +36.64 Unbiased estimate of differonoe =*+.558 122

TABIE 63

Comparison of the Heart Girth Measurements at Birth of Animals Heterozygous and Homozygous a t the Z Loous

H etero Homo f a - 'Z z ) (ni+n2<) Sire n i XI »2 ^2 D TrSf

cm. om. cm. 2 10 76.20 17 77.59 -1.39 6.30 3 19 78.53 17 78.59 - .06 8.97 8 7 79.00 21 79.62 - .62 5.25 10 6 79.33 10 82.80 -3 .4 7 3.53 11 49 81.84 70 81.34 + .50 28.82 12 7 75.86 30 78.73 -2 .8 7 5.68 14 11 75.64 9 77.56 -1.92 4.95 18 14 80.29 50 79.96 + .33 10.94 19 5 72.80 10 76.00 -3.20 3.33 21 7 80.71 9 81.89 -1.18 3.93 22 6 78.33 4 82.50 -4.1 7 2.40 23 12 76.58 16 79 .26 -2.67 6.86 24 52 81.23 83 81.14 + .09 31.97 25 14 80.21 23 79.96 + .25 8.70 26 14 77.71 63 77.71 - 8.86 27 12 79 .00 17 77.29 +1.71 7.03 28 10 80.00 30 77.73 +2.27 7.50 33 7 81.71 13 79.62 +2.09 4.55 34 4 78.50 14 78.85 - .30 3.11 38 15 80.07 22 79.00 +1.07 8.92 39 17 8U.24 25 79.72 + .52 10.12

298 554 172.86 brHD a -18.73 Unbiased estimate of differenoa a -0.108 om. 123

TABLE 64

Comparison of the Body Condition Evaluations at Three Months of Age of Animals Heterozygous and Homozygous at the Z Loous

(nixngj H etero Homo (ni+ngj S ire »1 *1 *2 *2 D W

2 8 4.625 13 4.846 -.221 4.95 3 15 5.733 8 5.375 +.368 5.22 8 7 5.285 21 5.047 +.238 5.25 10 6 5.500 7 5.571 -.0 7 1 3.23 11 45 5.311 62 5.032 + .279 26.07 12 5 5.000 21 5.571 -.5 7 1 4.04 18 14 4.928 50 4.920 + .008 10.94 19 5 4.600 9 5.111 -.511 3.21 21 6 6.500 6 5.883 -.3 8 3 3.00 22 7 4.472 4 5.250 -.7 7 8 2.55 23 13 5.000 16 4.687 +.313 7.17 24 49 5.428 80 5.425 +.003 30.39 25 14 5.428 23 5.521 -.0 9 3 8.70 26 14 4.857 65 5.061 -.2 0 4 11.52 27 11 4.909 19 5.052 -.1 4 3 6.96 28 10 4.400 30 4.866 -.4 6 6 7.50 33 7 5.428 15 5.666 -.238 4.78 34 4 5.500 16 5.125 + .375 3.20 38 16 4.600 22 4.045 +.555 8.92 39 18 4.277 25 4.400 -.123 10.47

273 514 268.07 + .49 Unbiased e stimate of differenoe « +0.002 124

TABLE 65

Comparison of the Inorease in Heart Girths From Three to Six Months of Age in Animals Heterozygous and Homozygous at the Z Loous

(n jsn g ; H otero Homo (Sl-Sii) U l+ n 2) S ire n i *2 *2 Q W

om. cm. cm. 2 9 27.22 11 27.18 + .04 4.95 3 10 22.50 6 22.83 - .33 3.75 8 7 21.14 20 18.50 +2.64 5.19 10 6 24.66 7 22.28 +2.38 3.23 11 48 23.83 68 22.77 +1.06 28.14 12 4 19.00 19 20.26 -1 .2 6 3.30 14 3 26.33 18 14 21.64 49 23.67 -2.03 10.88 19 5 25.80 9 26.88 -1.08 3.21 21 7 24.85 9 24.11 + .74 3.94 22 7 26.71 4 26.75 - .04 2.55 23 13 26.76 15 25.26 +1.50 6.96 24 51 23.78 82 22.85 + .93 31.44 25 12 20.33 23 20.04 + .29 7.89 26 14 23.92 64 25.28 + .64 11.49 27 12 27.41 19 27.36 + .05 7.35 28 10 21.20 29 20.68 + .52 7.44 33 7 22.00 15 20.86 +1.14 4.77 34 4 19.00 15 18.26 + .74 3.16 35 10 28.80 38 14 20.57 21 19.09 +1.48 8.40 39 14 22.78 24 20.16 +2.62 8.84

268 511 166.88 Z to = +120.21 Unbiased estimate of difference =* +0.720 cm. 125

TAB IE 66

Comparison of the Increase in Heart Girths From Three Months of Age to Three Months After Calving in Animals Heterozygous and Homozygous a t th e Z Loous

nl m 2 | Hetero Homo (T l-^2) (n]+n2J S ire m x l n2 w D W

om. om. om. 2 9 87.33 9 88.51 -1.1 8 4.50 3 7 86.57 4 92.75 -6.18 2.54 8 7 86.14 18 87.22 -1.08 5.04 10 5 84.20 8 83.00 +1.20 3.08 11 29 80.10 42 80.45 - .35 17.15 12 4 85.50 18 84.83 + .67 3.27 18 5 87.00 19 84.00 +3.00 3.96 19 5 77.20 7 80.71 -3.51 2.92 21 6 85.17 7 81.43 +3.74 3.23 22 5 77.80 2 81.50 —3*70 1*43 23 13 86.31 8 86.00 + .31 4*95 24 29 82.83 48 81.89 + .94 18.08 25 9 87.89 18 88*33 — .44 6.00 26 5 88.40 22 90.50 -2.10 4.07 27 7 85.43 13 84.08 +1.35 4.55 28 7 90.71 22 88.36 +2.35 5.31 38 12 93.58 16 93.81 - .23 6.86 39 16 88.25 20 88.80 - .55 8.89

182 302 106.80 ZiflD = -2.649 Unbiased estimate of difference « -0.025 om. 126

TABIE 67

Comparison of the Servioes Per Conception for Animals Heterozygous and Homozygous a t the Z Loous

(ni3m2) Hetero Homo C*L-*2) (ni+ngj S ire n i n2 ...... *2 D W

2 12 2.00 14 1.43 + .57 6.46 3 13 2.31 12 3.00 " .69 6.25 8 7 2.00 20 2.55 - .55 5.19 10 5 1.60 10 2.10 - .50 3.33 11 29 2.10 40 2.13 - .03 16.81 12 6 2.00 28 3.21 -1.21 4.94 14 10 1.30 7 1.71 - .41 4.12 18 8 3.43 28 2.79 + .64 6.22 19 5 1.40 7 1.57 - .17 2.92 21 5 2.20 8 1.75 + .45 3.08 22 4 1.50 2 2.00 - .50 1.33 23 8 1.27 9 1.22 + .05 4.24 24 23 2.17 36 1.47 + .70 14.03 25 12 2.58 21 2.71 - .13 7.64 26 4 1.25 22 2.55 -1.30 3.38 27 12 2.74 14 2.29 + .45 6.46 28 7 3.29 21 2.24 +1.05 5.25 34 4 2.75 14 2.79 - .04 3.11 38 11 2.82 13 2.00 ♦ .82 5.96 39 13 3.08 22 2.27 + .81 8.17

192 348 118.89 Z m =. +15.33 Unbiased estim ate of difference = +0.128 servioes 127

TABIE 68

Comparison of the Heart Girths of the First Offspring of -Animals Heterozygous and Homozygous at the Z Locus

Ul»X2> E etero Homo U i-x g ) (ni+n23 S ire *1 **L n2 ■*2 D W

cm. cm. cm. 2 4 77.50 8 75.13 +2.37 2.66 3 7 76.29 9 77.22 - .93 3.94 8 3 80.00 9 78.11 +1.89 2.25 10 4 78.50 5 79.80 -1 .3 0 2.22 11 18 79.56 17 79.94 - .38 8.74 12 2 80.00 12 78.83 ♦1.17 1.71 14 5 77.20 3 74.00 +3.20 1.88 19 2 71.50 3 75.00 -3 .5 0 1.20 21 4 77.50 5 80.20 -2 .7 0 2.22 23 6 77.50 4 75.50 +2.00 2.40 24 15 79.20 17 79.65 - .45 7.97 25 2 79.00 11 77.73 +1.27 1.69 26 2 79.00 14 78.71 + . 29 1.75 27 4 74.25 6 74.17 + .08 2.40 28 2 80.00 9 77.11 +2.89 1.64 34 2 74.00 8 74.50 - .50 1.60 39 7 77.29 7 79.43 -2 .1 4 3.50

90 1^9 49.77 SwD » -0.98 Unbiased estim ate of difference a -0.0196 am. 128

TABLE 69

Comparison of the Milk Production of Animals Heterozygous at Two or More Loci and Animals Heterozygous a t Hot More Than One Locus

Hetoro Hetero (n-ixn^J (2 ,3 ,4 ,5 ) (0 ,1 ) (■5q.-X2) (ni+n2) S ire n i n2 W D W

Lbs* Lbs* Lbs* 2 8 13326 5 12952 + 374 3.08 3 4 10323 19 9491 + 832 3.30 8 8 1U473 8 11166 - 693 4.00 11 37 12771 5 13866 -1095 4.40 12 2 10105 11 10203 - 98 1.69 18 8 10627 16 13016 -2389 5.33 22 2 15460 2 10210 +5250 1.00 23 8 12720 7 13868 -1148 3.73 24 21 14204 44 14267 - 63 14.22 25 10 11743 3 13347 -1604 2.31 26 2 10925 13 11409 - 484 1.73 27 10 12313 8 13353 -1040 4.44 28 9 10373 21 10641 - 268 6.30 35 4 14100 2 13655 + 445 1.33 38 14 11945 13 11252 + 693 6.74 39 10 13202 10 11929 +1273 5.00

157 187 68.60 a —•157*0.3 Unbiased estim ate of difference « -•229*45 lbs* 129

TABIE 70

Comparison of the Butterfat Produotion of Animals Heterozygous at Two or More Looi and Animals Heterozygous at Hot More Than One Locus

Hetero Hetero (2*3*4. 5) (0 ,1 ) (Si-X2) (ni+ng) S ire n i YT H2 ^2 D W

Lbs* Lbs* Lbs. 2 8 443 5 434 + 9 3.08 S 4 388 19 346 + 42 3.30 8 8 362 8 398 - 36 4.00 11 37 479 5 530 — 51 4.40 12 2 350 11 361 - 11 1.69 18 8 374 16 476 -102 5.33 22 2 555 2 405 +150 1.00 23 8 479 7 503 - 24 3.73 24 21 514 44 507 + 7 14.22 25 10 408 3 457 - 49 2.31 26 2 370 13 421 - 51 1.73 27 10 481 8 513 - 32 4.44 28 9 345 21 372 - 27 6.30 35 4 500 2 495 + 5 1.33 38 14 498 13 455 + 43 6.74 39 10 522 10 460 ♦ 62 5.00

156 187 68.60 S © =* -bll*44 Unbiased estimate of difference = -7.455 lbs. ISO

TABIE 71

Comparison of the Maturity Indexes of Animals heterozygous at Two or More Looi and Animals Heterozygous at itfot More Than One Locus

Hetero Hetero in,™,,) (2 ,3 ,4 ,5 ) ( 0,1 ) (*L-*2J (ni+njJ S ire » i Yl n2 *2 D W

2 6 88.0 5 88.6 - .6 2.70 3 2 100.5 11 101.3 - 1.3 1.69 8 4 112.5 4 109.5 + 3.0 2.00 11 21 94.5 2 95.5 - 1.0 1.83 12 2 110.0 9 96.9 4-13.1 1.64 18 3 98.3 10 105.3 - 7.0 2.31 23 4 76.0 4 80.5 - 4.5 1.33 24 14 92.7 17 100.7 - 8.0 7.68 25 8 96.1 2 103.0 — 6.9 1.60 27 7 92.7 4 94.8 - 2.1 2.55 38 2 81.5 4 88.8 - 7.3 1.33 39 __ 2 97.0 _3 99.7 - 2.7 1.20

118 75 27.80 m =, -91.10 Unbiased estimate of difference = -3.27 131

TABIE 72

Comparison, of tho Heart Girth Measurements at Birth of Animals Heterozygous a t Two or Moro Loci and Animals Heterozygous at Not More Than One Loous

H etero Hetero (n ix a 2j (2 ,3 ,4 ,5 ) (0 ,1 ) U i~ x 2) (n x+n2) S ire *1 *1 n 2 *2 0 W

om* om. om. 2 8 76*8 5 77.6 - .8 3.08 3 6 78.8 20 78.3 + .5 4.62 3 9 78.8 11 79.4 - .6 4.95 11 65 81.9 9 82.3 - .4 7.90 12 2 70.5 11 78.9 -8 .4 1.69 18 19 80.2 28 80.3 - .1 11.32 23 9 76.4 10 80.3 -3 .9 4.74 24 46 81.3 89 81.1 + .2 30.32 25 10 79.9 2 79.5 + .4 1.67 26 8 77.4 35 78.0 - .6 6.51 27 16 79.1 10 76.4 +2.7 6.15 28 12 79.9 28 77.6 +2.3 8.40 33 4 83.8 13 79.7 +4.1 5.06 35 4 75.3 4 74.8 + .5 2.00 38 19 80.2 18 78.7 +1.5 9.24 39 12 80.6 12 80.1 + .5 6.00

249 305 111*65 -CTO) =. +29.06 Unbiased estim ate of difference a +.26 cm. 132

TABIE 73

Comparison of the Body Condition Evaluations at Throe Months of Age of Animals Heterozygous at Two or More Loci and Animals heterozygous at hot More Than One Loous

H etero H etero IrijXngJ (2 j 314* 5) (O.iJ (n-t+n2j S ire ni ^ r n2 *2 D W

2 6 4.33 5 5.00 -.6 7 2.72 3 6 5.50 16 5.56 -.0 6 4.36 8 9 4.41 11 5.09 -.6 8 4.95 11 64 b . l l 8 5.38 -.2 7 7.11 18 19 4.74 29 5.00 -.2 6 11.48 22 2 b.00 2 5.00 .00 1.00 23 9 4.78 10 4.70 + .08 4.74 24 44 5.46 85 5.41 + .05 28.99 25 10 5,60 2 5.00 + .60 1.67 26 8 5.00 35 5.11 -.1 1 6. 52 1 o 27 15 5.00 10 5.10 . 6.00 28 12 4.50 28 4.86 -.3 6 8.40 33 4 5.50 15 5.73 —. 23 3.16 35 4 5.25 4 4.50 + .75 2.00 38 19 4.37 18 4.17 + .20 9.24 39 13 4.62 12 4.15 + .47 6.24

244 ^ 290 108.58 2Twd = - 6.31 Unbiased estim ate of difference =* -0.068 133

TABLE 74 ,

Comparison of the Increase in Heart Girths From Three to Six Months of Age in Animals Heterozygous at Two or More Loci and Animals Heterozygous at Not More Than One Loous

Hetero Hetero C^xnjj) (2 ,3 , 4, b) ( 0,1 ) { y \ - r 2) (ni+n2) d ire n l *1 n2 *2 D W

om. cm. om. 2 7 27.71 4 25.50 ♦2.21 2.40 3 5 22.60 11 22.64 - .04 3.44 8 9 18.00 10 20.80 -2.80 4.74 11 65 23.12 9 23.35 - .21 7.91 18 18 24.33 29 21.90 +2.93 11.11 22 2 26.50 2 30.00 -3.50 1.00 23 9 26.55 9 25.44 +1.11 4.50 24 44 24.23 89 22-81 +1.42 29.44 2b 9 19.22 4 20. bO -1.28 1.64 26 6 23.62 34 22.59 +1.03 5.10 27 16 26.62 10 28.30 -1.68 6.15 28 12 19.92 27 21.22 -1 .3 0 8.31 33 4 20.25 15 21.60 -1 .3 5 3.16 35 4 29.50 4 27.75 +1.75 2.00 38 19 19.47 16 19.^4 - .47 8.69 39 10 21.50 11 21.09 + .41 5.24

ZlSD = ♦45.40 Unbiased estim ate of difference o +0.433 cm. 134

TABIE 75

Comparison of tne Increase in Heart Girth From Three Months of Age to Three Months After Calying in Animals Heterozygous at Two or More Loci and Animals Heterozygous at Not More Than One Locus

Hetero Hetero (2 ,3 ,4 ,5 ) (o ,D (Ti-Xg) tn,+ n S ire n i XI n 2 ■ * r D vr

om* cm. cm. 2 7 88.70 2 85.50 +3.20 1.56 3 4 86.75 7 90.00 -3.25 2.55 8 8 84.75 8 88.13 -3.38 4.00 11 37 80.89 6 81.83 -0.94 5.16 18 3 94.00 12 84.42 +9.58 2.40 23 8 86.10 5 83.00 +3.10 3.08 24 24 83.80 53 81.53 +2.27 16.52 25 8 84.25 2 93.50 -9.25 1.60 26 3 89.00 11 89.27 - .27 2.36 27 11 83.82 8 85.63 -1.81 4.63 28 8 89.63 21 88.67 + .96 5.79 33 2 74.50 4 82.75 -8.25 1.33 35 4 90.50 2 86.50 +4.00 1.33 38 16 92.44 7 95.27 -2.83 4.87 85.09 89.20 39 - 1 1 10 -4.11 5.24 164 158 62.40 -10.86 Unbiased estim ate of difference = -0.174 cm. 135

TABIE 76

Comparison of the Services Per Conception For Animals Heterozygous at Two or More Loci and Animals Heterozygous a t Hot More Than One Locus

Hetero Hetero (Dlm 2/ ( 2 ,3 ,4 ,b; (0 ,1 ) C*l-x2) (ni+ng) S ire n l H2 *2 D W

2 7 1.57 5 1.00 + .57 2.92 3 5 1.80 15 2.67 - .87 3.75 8 8 1.91 8 2.50 - .59 4.00 11 35 2.11 5 1 • 60 + .51 4.38 12 2 2.50 10 2.20 + .30 1.67 18 7 2.86 16 3.38 - .52 4.87 23 5 1.60 6 1.17 + .43 2.73 24 20 1.6U 39 1.82 - .22 13.22 25 10 3.50 3 1.00 +2.50 2.31 26 2 2.00 10 2.20 - .20 1.67 27 8 1.13 8 2.50 -1.37 4.00 28 9 3.22 19 2.16 ♦1.06 6.11 3b 4 1.25 2 1.00 + .25 1.33 38 15 2.47 8 2.38 + .09 5.22 39 8 3.75 11 2.18 +1.57 4.63

145 165 62.81 ZjW = ♦9.01 Unbiased estim ate of difference = ♦0.143 servioes 136

TABLE 77

Comparison of the Heart Girths of the First Offspring of Animals Heterozygous at Two or More Loci and Animals Heterozygous at Not More Than One Locus

Hetero H etero (2 ,3 ,4 , ( 0 , 1 ) S ire n i X l n2 *2 D

om* om. om. 2 4 74.20 4 78.25 -4.05 1 .0 0 3 3 77.00 10 77.50 - .50 2.30 8 3 80.67 4 79.60 +1.17 1.71 11 14 78.79 2 80.00 -1.21 1.75 18 2 78.00 5 79.20 -1 .2 0 1.43 23 3 79.33 4 75.50 +3 . 03 1.71 24 13 80.15 19 78.95 +1.20 7.72 27 5 73.60 2 75.50 -1.90 1.43 28 2 76.50 8 77.37 - .87 1.60 38 2 78.00 4 77.50 + .50 1.33

49 60 21.98 Jw"V® sr +5.34 Unbiased estimate of difference = +0.24 om* AUTOBIOGRAPHY

I, Harry Lionel Barr, was born near Morristown, Ohio, February

15, 1922* I received my secondary school education in the publio sohools of Bamesville, Ohio. I pursued my undergraduate training at

Georgetown University and Ohio State University, the latter of which granted me the Bachelor of Science degree in 1954. I was appointed

Instructor in the Department of Dairy Science in January, 1955, and have continued in this position while earning the Master of Science degree, and while completing the requirements for the degree of Doctor of Philosophy*

137