Search

Nonadditive genetic effects are currently ignored in national genetic evaluations of farm animals because of ignorance of thelevel of dominance variance for traits of interest and the difficult computational problems involved. Potential gains fromincluding the effects of dominance in genetic evaluations include “purification” of additive values and availability ofpredictions of specific combining abilities for each pair of prospective parents. This study focused on making evaluation withdominance effects feasible computationally and on ascertaining benefits of such an evaluation for dairy cattle, beef cattle,and swine. Using iteration on data, computing costs for evaluation with dominance effects included costs could be less thantwice expensive as with only an additive model. With Method Â, variance components could be estimated for problemsinvolving up to 10 millions equations. Dominance effects accounted for up to 10% of phenotypic variance; estimates werelarger for growth traits. As a percentage of additive variance, the estimate of dominance variance reached 78% for 21-d litterweight of swine and 47% for post weaning weight of beef cattle. When dominance effects are ignored, additive evaluationsare “contaminated”; effects are greatest for evaluations of dams in a single large family. These changes in ranking wereimportant for dairy cattle, especially for dams of full-sibs, but were less important for swine. Specific combining abilitiescannot be included in sire evaluations and need to be computed separately for each set of parents. The predictions of specificcombining abilities could be used in computerized mating programs via the Internet. Gains from including the dominanceeffect in genetic evaluations would be moderate but would outweigh expenditures to produce those evaluations.

The dairy industry is known for its extensive use of artificial insemination, which has resulted in a population where most animals can be traced back to only a few sires. Due to their relatedness to the population, old influential sires could still contribute to the accuracy of genomic predictions. The objective of the study was to identify the impact of historically influential sires on the recent population. This was tested by constructing a genomic relationship matrix using recursion with different sets of sires. Differences in prediction accuracies with different sets are indicative of how important each set is. Recursion coefficients linking young animals to those sets reveal the relative importance of specific sires to the prediction accuracy of recent animals. The data included ∼10 million scores for stature and fore udder attachment (FUA) measured from 1983. Genotypes of 569,404 animals were available. Sire sets included the 100 most popular sires born within different time periods. Computations were with single-step genomic BLUP. In general, the younger sires had higher prediction accuracies than the oldest sires, even though they generally have fewer progeny. The accuracy of evaluation for stature was increased from 0.54 with the most popular sires born before 1981 to 0.69 with sires born from 2001 to 2010, while the accuracy for FUA increased from 0.47 to 0.61. The accuracy achieved using the overall 100 most used sires was 0.66 for stature and 0.58 for FUA. All 100 sires from each period were combined in a subset to determine the importance of each sire relative to all 400 animals in the combined subset. The highest relative impact of a sire that was born within the different time sets was 1.97 for Valiant (before 1981), 1.94 for Blackstar (1981 to 1990), 4.38 for Shottle (1991 to 2000), and 3.09 for Planet (2001 to 2010). The 3 sires among the 400 with the greatest impact were Shottle, Goldwyn (3.73), and Planet. The relative impact of a sire was not strongly related to the number of progeny. For instance, the relative impact of Durham with 34K progeny was 2.29, whereas the impact of O Man with 15K progeny was 3.13. The impact of a sire is also influenced by whether it was used as a sire of sires. Results show that younger sires are more relevant to the accuracy of breeding value prediction in the recent population., (© 2023.)

Over half a million Holsteins are being genotyped annually in the United States. The computational cost of including all genotypes in single-step genomic (ssG)BLUP is high, although it is feasible to conduct large-scale genomic prediction using an efficient algorithm such as APY (algorithm for proven and young). An effective method to further reduce the computing cost could be the use of indirect genomic predictions (IGP) for genotyped animals when they have neither progeny nor phenotypes. These young genotyped animals have no effect on the other genotyped animals and could have their genomic prediction done indirectly. The main objective of this study was to calculate IGP for various groups of genotyped animals and investigate the reduction in computing time as well as bias and accuracy of the IGP. We compared IGP with genomic (G)EBV for 18 linear type traits in US Holsteins, including 2.3 million (M) genotyped animals. The full data set consisted of 10.9M records for 18 linear type traits up to 2018 calving, 13.6M animals in the pedigree, and 2.3M animals genotyped for 79K SNP. For IGP, ssGBLUP included all genotyped animals except those with neither progeny nor phenotypes by year from 2014 to 2018 (i.e., the target animals). The SNP marker effects were computed based on GEBV for genotyped animals that had progeny, or phenotypes, or both. Further, IGP were calculated for target genotyped animals in each year group. For all genotyped animal groups from 2014 to 2018, the coefficients of determination (R 2 ) of a linear regression of GEBV on IGP were 0.960 for males and 0.954 for females for 18 traits on average. To reduce computing costs, the SNP marker effects were calculated based on GEBV from randomly selected genotyped animals from 15K to 60K. By randomly selecting a small number of genotyped animals, the computing time was dramatically reduced. As more genotyped animals were randomly selected to calculate SNP effects, R 2 was higher (more accurate) and the regression coefficient was lower (more inflated IGP). In a practical genomic evaluation in US Holsteins, to get sufficient contributions from GEBV, 25K to 35K is a rational number of genotyped animals that can be randomly selected to compute SNP effects and obtain accurate and unbiased IGP. Considering the computing time and both unbiasedness and accuracy of IGP, genomic evaluation can be conducted separately in GEBV for genotyped animals with phenotypes or progeny and in IGP for young genotyped animals. This can be a practical solution when conducting a large-scale genomic evaluation and would enable more frequent evaluation at lower cost, especially when many genotyped animals have neither phenotypes nor progeny., (© 2021.)

The objective of this study was to clarify how bias in genomic predictions is created by investigating a relationship among selection intensity, a change in heritability (Δh 2 ), and assortative mating (ASM). A change in heritability, resulting from selection, reflects the impact that the Bulmer effect has on the reduction in between-family variation, whereas assortative mating impacts the within-family variance or Mendelian sampling variation. A partial data set up to 2014, including 841K genotyped animals, was used to calculate genomic predictions with a single-step genomic model for 18 linear type traits in US Holsteins. A full data set up to 2018, including 2.3 million genotyped animals, was used to calculate benchmark genomic predictions. Inbreeding and unknown parent groups for missing parents of animals were included in the model. Genomic evaluation was performed using 2 different genetic parameters: those estimated 14 yr ago, which have been used in the national genetic evaluation for linear type traits in the United States, and those newly estimated with recent records from 2015 to 2018 and those corresponding pedigrees. Genetic trends for 18 type traits were estimated for bulls with daughters and cows with phenotypes in 2018. Based on selection intensity and mating decisions, these traits can be categorized into 3 groups: (a) high directional selection, (b) moderate selection, and (c) intermediate optimum selection. The first 2 categories can be explained by positive assortative mating, and the last can be explained by negative assortative or disassortative mating. Genetic progress was defined by genetic gain per year based on average standardized genomic predictions for cows from 2000 to 2014. Traits with more genetic progress tended to have more "inflated" genomic predictions (i.e., "inflation" means here that genomic predictions are larger in absolute values than expected, whereas "deflation" means smaller than expected). Heritability estimates for 14 out of 18 traits declined in the last 16 yr, and Δh 2 ranged from -0.09 to 0.04. Traits with a greater decline in heritability tended to have more deflated genomic predictions. Biases (inflation or deflation) in genomic predictions were not improved by using the latest genetic parameters, implying that bias in genomic predictions due to preselection was not substantial for a large-scale genomic evaluation. Moreover, the strong selection intensity was not fully responsible for bias in genomic predictions. The directional selection can decrease heritability; however, positive assortative mating, which was strongly associated with large genetic gains, could minimize the decline in heritability for a trait under strong selection and could affect bias in genomic predictions., (The Authors. Published by Elsevier Inc. and Fass Inc. on behalf of the American Dairy Science Association®. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).)

The objectives of this study were to investigate bias in genomic predictions for dairy cattle and to find a practical approach to reduce the bias. The simulated data included phenotypes, pedigrees, and genotypes, mimicking a dairy cattle population (i.e., cows with phenotypes and bulls with no phenotypes) and assuming selection by breeding values or no selection. With the simulated data, genomic estimated breeding values (GEBV) were calculated with a single-step genomic BLUP and compared with true breeding values. Phenotypes and genotypes were simulated in 10 generations and in the last 4 generations, respectively. Phenotypes in the last generation were removed to predict breeding values for those individuals using only genomic and pedigree information. Complete pedigrees and incomplete pedigrees with 50% missing dams were created to construct the pedigree-based relationship matrix with and without inbreeding. With missing dams, unknown parent groups (UPG) were assigned in relationship matrices. Regression coefficients (b 1 ) and coefficients of determination (R 2 ) of true breeding values on (G)EBV were calculated to investigate inflation and accuracy in GEBV for genotyped animals, respectively. In addition to the simulation study, 18 linear type traits of US Holsteins were examined. For the 18 type traits, b 1 and R 2 of GEBV with full data sets on GEBV with partial data sets for young genotyped bulls were calculated. The results from the simulation study indicated inflation in GEBV for genotyped males that were evaluated with only pedigree and genomic information under BLUP selection. However, when UPG for only pedigree-based relationships were included, the inflation was reduced, accuracy was highest, and genetic trends had no bias. For the linear type traits, when UPG for only pedigree-based relationships were included, the results were generally in agreement with those from the simulation study, implying less bias in genetic trends. However, when including no UPG, UPG in pedigree-based relationships, or UPG in genomic relationships, inflation and accuracy in GEBV were similar. The results from the simulation and type traits suggest that UPG must be defined accurately to be estimable and inbreeding should be included in pedigree-based relationships. In dairy cattle, known pedigree information with inbreeding and estimable UPG plays an important role in improving compatibility between pedigree-based and genomic relationship matrices, resulting in more reliable genomic predictions., (Copyright © 2019 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

The variance of gametic diversity ( σ gamete 2 ) can be used to find individuals that more likely produce progeny with extreme breeding values. The aim of this study was to obtain this variance for individuals from routine genomic evaluations, and to apply gametic variance in a selection criterion in conjunction with breeding values to improve genetic progress. An analytical approach was developed to estimate σ gamete 2 by the sum of binomial variances of all individual quantitative trait loci across the genome. Simulation was used to verify the predictability of this variance in a range of scenarios. The accuracy of prediction ranged from 0.49 to 0.85, depending on the scenario and model used. Compared with sequence data, SNP data are sufficient for estimating σ gamete 2 Results also suggested that markers with low minor allele frequency and the covariance between markers should be included in the estimation. To incorporate σ gamete 2 into selective breeding programs, we proposed a new index, relative predicted transmitting ability, which better utilizes the genetic potential of individuals than traditional predicted transmitting ability. Simulation with a small genome showed an additional genetic gain of up to 16% in 10 generations, depending on the number of quantitative trait loci and selection intensity. Finally, we applied σ gamete 2 to the US genomic evaluations for Holstein and Jersey cattle. As expected, the DGAT1 gene had a strong effect on the estimation of σ gamete 2 for several production traits. However, inbreeding had a small impact on gametic variability, with greater effect for more polygenic traits. In conclusion, gametic variance, a potentially important parameter for selection programs, can be easily computed and is useful for improving genetic progress and controlling genetic diversity., (The Authors. Published by FASS Inc. and Elsevier Inc. on behalf of the American Dairy Science Association®. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).)

The objective of this study was to compare genetic trends from single-step genomic BLUP (ssGBLUP) and traditional BLUP models for milk production traits of US Holsteins. Phenotypes were 305-d milk, fat, and protein yields from 21,527,040 cows recorded between January 1990 and August 2015. The pedigree file included 29,651,623 animals and was limited to 3 generations back from recorded or genotyped animals. Genotypes for 764,029 animals were used, and analyses were by a 3-trait repeatability model as used in the US official genetic evaluation. Unknown-parent groups were incorporated into the inverse of a relationship matrix (H -1 in ssGBLUP and A -1 in BLUP) with the QP transformation. For ssGBLUP, 18,359 genotyped animals were randomly chosen as core animals to calculate the inverse of the genomic relationship matrix with the APY algorithm. Computations took 6.5 h and 1.4 GB of memory for BLUP, and 13 h and 115 GB of memory for ssGBLUP. For genotyped sires with at least 10 daughters, the average genetic levels for predicted transmitting ability (PTA) and genomic PTA were similar up to 2008, with a higher level for ssGBLUP later (approximately by 36 kg for milk, 2.1 kg for fat, and 1.1 kg for protein for bulls born in 2010). For genotyped cows, the average genetic levels were similar up to 2006, with a higher level for ssGBLUP (approximately by 91 kg for milk, 3.6 kg for fat, and 2.7 kg for protein for cows born in 2012). For all cows, the average levels were slightly higher for ssGBLUP, with much smaller differences than for genotyped cows. Trends for BLUP indicate bias due to genomic preselection for genotyped sires and cows. For official evaluations released in December 2016, traditional PTA had the same trend as multiple-step genomic PTA for both genotyped bulls and cows except for the youngest bulls, who had traditional PTA slightly lower than genomic PTA. For genotyped bulls born in recent years, genetic gain for official traditional and genomic evaluations was similar in contrast to ssGBLUP and BLUP differences. Official PTA for cows were adjusted so that the Mendelian sampling variance was comparable with that for bulls, and those adjustments likely removed bias due to genomic preselection from traditional PTA, especially for genotyped cows. The ssGBLUP method seems to account partially for that bias and is computationally suitable for national evaluations., (The Authors. Published by FASS Inc. and Elsevier Inc. on behalf of the American Dairy Science Association®. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/3.0/).)

The objective of this study was to investigate the feasibility of genomic evaluation for cow mortality and milk production using a single-step methodology. Genomic relationships between cow mortality and milk production were also analyzed. Data included 883,887 (866,700) first-parity, 733,904 (711,211) second-parity, and 516,256 (492,026) third-parity records on cow mortality (305-d milk yields) of Holsteins from Northeast states in the United States. The pedigree consisted of up to 1,690,481 animals including 34,481 bulls genotyped with 36,951 SNP markers. Analyses were conducted with a bivariate threshold-linear model for each parity separately. Genomic information was incorporated as a genomic relationship matrix in the single-step BLUP. Traditional and genomic estimated breeding values (GEBV) were obtained with Gibbs sampling using fixed variances, whereas reliabilities were calculated from variances of GEBV samples. Genomic EBV were then converted into single nucleotide polymorphism (SNP) marker effects. Those SNP effects were categorized according to values corresponding to 1 to 4 standard deviations. Moving averages and variances of SNP effects were calculated for windows of 30 adjacent SNP, and Manhattan plots were created for SNP variances with the same window size. Using Gibbs sampling, the reliability for genotyped bulls for cow mortality was 28 to 30% in EBV and 70 to 72% in GEBV. The reliability for genotyped bulls for 305-d milk yields was 53 to 65% to 81 to 85% in GEBV. Correlations of SNP effects between mortality and 305-d milk yields within categories were the highest with the largest SNP effects and reached >0.7 at 4 standard deviations. All SNP regions explained less than 0.6% of the genetic variance for both traits, except regions close to the DGAT1 gene, which explained up to 2.5% for cow mortality and 4% for 305-d milk yields. Reliability for GEBV with a moderate number of genotyped animals can be calculated by Gibbs samples. Genomic information can greatly increase the reliability of predictions not only for milk but also for mortality. The existence of a common region on Bos taurus autosome 14 affecting both traits may indicate a major gene with a pleiotropic effect on milk and mortality., (Copyright © 2017 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

The objectives of this study were to develop and evaluate an efficient implementation in the computation of the inverse of genomic relationship matrix with the recursion algorithm, called the algorithm for proven and young (APY), in single-step genomic BLUP. We validated genomic predictions for young bulls with more than 500,000 genotyped animals in final score for US Holsteins. Phenotypic data included 11,626,576 final scores on 7,093,380 US Holstein cows, and genotypes were available for 569,404 animals. Daughter deviations for young bulls with no classified daughters in 2009, but at least 30 classified daughters in 2014 were computed using all the phenotypic data. Genomic predictions for the same bulls were calculated with single-step genomic BLUP using phenotypes up to 2009. We calculated the inverse of the genomic relationship matrix GAPY(-1) based on a direct inversion of genomic relationship matrix on a small subset of genotyped animals (core animals) and extended that information to noncore animals by recursion. We tested several sets of core animals including 9,406 bulls with at least 1 classified daughter, 9,406 bulls and 1,052 classified dams of bulls, 9,406 bulls and 7,422 classified cows, and random samples of 5,000 to 30,000 animals. Validation reliability was assessed by the coefficient of determination from regression of daughter deviation on genomic predictions for the predicted young bulls. The reliabilities were 0.39 with 5,000 randomly chosen core animals, 0.45 with the 9,406 bulls, and 7,422 cows as core animals, and 0.44 with the remaining sets. With phenotypes truncated in 2009 and the preconditioned conjugate gradient to solve mixed model equations, the number of rounds to convergence for core animals defined by bulls was 1,343; defined by bulls and cows, 2,066; and defined by 10,000 random animals, at most 1,629. With complete phenotype data, the number of rounds decreased to 858, 1,299, and at most 1,092, respectively. Setting up GAPY(-1) for 569,404 genotyped animals with 10,000 core animals took 1.3h and 57 GB of memory. The validation reliability with APY reaches a plateau when the number of core animals is at least 10,000. Predictions with APY have little differences in reliability among definitions of core animals. Single-step genomic BLUP with APY is applicable to millions of genotyped animals., (Copyright © 2016 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

To include feed-intake-related traits in the breeding goal, accurate estimates of genetic parameters of feed intake, and its correlations with other related traits (i.e., production, conformation) are required to compare different options. However, the correlations between feed intake and conformation traits can vary depending on the population. Therefore, the objective was to estimate genetic correlations between 6 feed-intake-related traits and 7 conformation traits within dairy cattle from 2 countries, the Netherlands (NL) and the United States (US). The feed-intake-related traits were dry matter intake (DMI), residual feed intake (RFI), milk energy output (MilkE), milk yield (MY), body weight (BW), and metabolic body weight (MBW). The conformation traits were stature (ST), chest width (CW), body depth (BD), angularity (ANG), rump angle (RA), rump width (RW), and body condition score (BCS). Feed intake data were available for 1,665 cows in NL and for 1,920 cows in US, from 83 nutritional experiments (48 in NL and 35 in US) conducted between 1991 and 2011 in NL and between 2007 and 2013 in US. Additional conformation records from relatives of the animals with DMI records were added to the database, giving a total of 37,241 cows in NL and 28,809 in US with conformation trait information. Genetic parameters were estimated using bivariate animal model analyses. The model included the following fixed effects for feed-intake-related traits: location by experiment-ration, age of cow at calving modeled with a second order polynomial by parity class, location by year-season, and days in milk, and these fixed effects for the conformation traits: herd by classification date, age of cow at classification, and lactation stage at classification. Both models included additive genetic and residual random effects. The highest estimated genetic correlations involving DMI were with CW in both countries (NL=0.45 and US=0.61), followed by ST (NL=0.33 and US=0.57), BD (NL=0.26 and US=0.49), and BCS (NL=0.24 and US=0.46). The MilkE and MY were moderately correlated with ANG in both countries (0.33 and 0.47 in NL, and 0.36 and 0.48 in US). Finally, BW was highly correlated with CW (0.77 in NL and 0.84 in US) and with BCS (0.83 in NL and 0.85 in US). Feed-intake-related traits were moderately to highly genetically correlated with conformation traits (ST, CW, BD, and BCS) in both countries, making them potentially useful as predictors of DMI., (Copyright © 2016 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

The objective of this study was to investigate genotype by environment interactions for culling rates and milk production in large and small dairy herds in 3 US regions, using genotypes, pedigree, and phenotypes. Single nucleotide polymorphism (SNP) marker variances were also estimated in these different environments. Culling rates including cow mortality were based on 6 Dairy Herd Improvement termination codes reported by dairy producers. Separate data sets for culling rates and 305-d milk yield were created for large and small dairy herds in the US regions of the Southeast (SE), Southwest (SW), and Northeast (NE) for the first 3 lactation cows that calved between 1999 and 2008. Genomic information from 42,503 SNP markers on 34,506 bulls was included in the analysis to predict genomic estimated breeding value (GEBV) of culling rates and 305-d milk yield with a single-step genomic BLUP using a bivariate threshold-linear model. Cow replacement rates in large SE and NE herds were higher. Heritability estimates of culling rates ranged from 0.03 to 0.11, but the differences were small between large and small herds and among the 3 US regions. Genetic correlations between culling rates and 305-d milk yield were medium to high for cows sold for poor production and reproduction problems. Correlations of GEBV for culling rates among the 3 US regions ranged from 0.34 to 0.92 and were lower between the SW and the other regions, especially in small herds. Correlations of GEBV between large and small herds ranged from 0.44 to 0.90 and were lower in the SW. These results indicate genotype by environment interactions of cow culling rate between the US regions and between large and small herds. Correlations of top 30 SNP marker effects for culling rates between 2 US regions ranged from 0.64 to 0.98 and were higher than those of more SNP marker effects except for a culling reason "sold for dairy purpose." Those correlations between large and small herds ranged from 0.67 to 0.98. High correlations of top SNP marker effects on culling reasons between the US regions and between large and small herds suggest that major markers can be useful for selection in different environments. The SNP variance shown in a marker gene segment on chromosome 14 was strongly associated with milk production in large and small herds in the NE but not in the SE and SW. Marker genes on chromosome 14 also showed a strong association with cow culling rates due to poor production and mortality in large herds in the NE., (Copyright © 2015 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

The purpose of this study was to evaluate the accuracy of genomic selection in single-step genomic BLUP (ssGBLUP) when the inverse of the genomic relationship matrix (G) is derived by the "algorithm for proven and young animals" (APY). This algorithm implements genomic recursions on a subset of "proven" animals. Only a relationship matrix for animals treated as "proven" needs to be inverted, and the extra costs of adding animals treated as "young" are linear. Analyses involved 10,102,702 final scores on 6,930,618 Holstein cows. Final score, which is a composite of type traits, is popular trait in the United States and was easily available for this study. A total of 100,000 animals with genotypes were used in the analyses and included 23,000 sires (16,000 with >5 progeny), 27,000 cows, and 50,000 young animals. Genomic EBV (GEBV) were calculated with a regular inverse of G, and with the G inverse approximated by APY. Animals in the proven subset included only sires (23,000), sires+cows (50,000), only cows (27,000), or sires with >5 progeny (16,000). The correlations of GEBV with APY and regular GEBV for young genotyped animals were 0.994, 0.995, 0.992, and 0.992, respectively Later, animals in the proven subset were randomly sampled from all genotyped animals in sets of 2,000, 5,000, 10,000, 15,000, and 20,000; each sample was replicated 4 times. Respective correlations were 0.97 (5,000 sample), 0.98 (10,000 sample), and 0.99 (20,000 sample), with minimal difference between samples of the same size. Genomic EBV with APY were accurate when the number of animals used in the subset is between 10,000 and 20,000, with little difference between the ways of creating the subset. Due to the approximately linear cost of APY, ssGBLUP with APY could support any number of genotyped animals without affecting accuracy., (Copyright © 2015 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

Assigning unknown parent groups (UPG) in mixed-model equations using single-step genomic BLUP was investigated to reduce bias and to increase accuracy in genomic estimated breeding values (GEBV). The original UPG were defined based on the animal's birth year and the sex of the animal's unknown parents. Combining the last 2 UPG for the animals' birth years and separating the UPG for US and non-US Holsteins were considered in the redefinition. A full data set in the 2011 national genetic evaluation of final score in US Holsteins was used to calculate estimated breeding values (EBV) for validation, and a subset of the 2011 data, which excluded phenotypes recorded after 2007, was used to calculate GEBV for all animals, including 34,500 genotyped bulls. The EBV and GEBV in 2007 were compared with EBV in the 2011 full data. The last group effects for unknown sires and dams were overestimated with the GEBV model using the reduced 2007 data. The genetic trends from EBV in 2011 and GEBV in 2007 with the original UPG in the last few years demonstrated inflation, whereas GEBV with the redefined UPG by combining the last 2 groups showed deflation. On the other hand, the redefined UPG by separating for US and non-US Holsteins reduced the bias in GEBV. Regression coefficients smaller than unity for GEBV for young genotyped bulls with no daughters in 2007 on progeny deviations in 2011 also indicated inflation. The redefining of UPG reduced bias and slightly increased accuracy in GEBV for both US and non-US genotyped bulls. Rank correlations between GEBV in 2007 and in 2011 with the redefined UPG were higher than those with no UPG and the original UPG, especially for non-US bulls. Redefining of UPG in genomic evaluation could improve reliability of GEBV and provide correct genetic trends., (Copyright © 2014 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

Data sets of US Holsteins, Israeli Holsteins, and pigs from PIC (a Genus company, Hendersonville, TN) were used to evaluate the effect of different numbers of generations on ability to predict genomic breeding values of young genotyped animals. The influence of including only 2 generations of ancestors (A2) or all ancestors (Af) was also investigated. A total of 34,506 US Holsteins, 1,305 Israeli Holsteins, and 5,236 pigs were genotyped. The evaluations were computed by traditional BLUP and single-step genomic BLUP, and computing performance was assessed for the latter method. For the 2 Holstein data sets, coefficients of determination (R(2)) and regression (δ) of deregressed evaluations from a full data set with records up to 2011 on estimated breeding values and genomic estimated breeding values from the truncated data sets were computed. The thresholds for data deletion were set by intervals of 5 yr, based on the average generation interval in dairy cattle. For the PIC data set, correlations between corrected phenotypes and estimated or genomic estimated breeding values were used to evaluate predictive ability on young animals born in 2010 and 2011. The reduced data set contained data up to 2009, and the thresholds were set based on an average generation interval of 3 yr. The number of generations that could be deleted without a reduction in accuracy depended on data structure and trait. For US Holsteins, removing 3 and 4 generations of data did not reduce accuracy of evaluations for final score in Af and A2 scenarios, respectively. For Israeli Holsteins, the accuracies for milk, fat, and protein yields were the highest when only phenotypes recorded in 2000 and later were included and full pedigrees were applied. Of the 135 Israeli bulls with genotypes (validation set) and daughter records only in the complete data set, 38 and 97 were sons of Israeli and foreign bulls, respectively. Although more phenotypic data increased the prediction accuracy for sons of Israeli bulls, the reverse was true for sons of foreign bulls. Also, more phenotypic data caused large inflation of genomic estimated breeding values for sons of foreign bulls, whereas the opposite was true with the deletion of all but the most recent phenotypic data. Results for protein and fat percentage were different from those for milk, fat, and protein yields; however, relatively, the changes in coefficients of determination and regression were smaller for percentage traits. For PIC data set, removing data from up to 5 generations did not erode predictive ability for genotyped animals for the 2 reproductive traits used in validation. Given the data used in this study, truncating old data reduces computation requirements but does not decrease the accuracy. For small populations that include local and imported animals, truncation may be beneficial for one group of animals and detrimental to another group., (Copyright © 2014 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

Currently, the US Department of Agriculture Animal Improvement Programs Laboratory utilizes a multi-step procedure in genomic evaluations for US Holstein bulls and cows, with adjustments for cows. We used a single-step procedure to investigate whether adding cows' genotypes could increase reliability in genomic breeding values for bulls while minimizing bias. The first data set to 2007 was used to calculate genomic estimated breeding values (GEBV) for animals, including young genotyped bulls with no daughters and young cows (heifers) with no records in 2007. The second data set to 2011 was used to calculate GEBV for the same animals, including those young bulls with daughters and young cows with records in 2011. Genotypes (42,503 single nucleotide polymorphism markers) for 34,506 bulls and 5,235 cows from 356,413 bulls and 9,245,619 cows in pedigree were used to calculate single-step GEBV (ssGEBV) and multi-step GEBV (msGEBV). Regression coefficients of 2007 GEBV on 2011 progeny deviations and coefficients of determination were used as indicators of bias and reliability in 2007 GEBV for bulls with no daughters and for cows with no records in 2007, using bull genotypes only and using bull and cow genotypes. Parent averages were also calculated from estimated breeding values of parents to compare with GEBV. For genotyped bulls, inflation was larger for ssGEBV than for msGEBV, whereas reliability was higher for ssGEBV. Using all genotyped bulls and cows, reliabilities were increased by 2 to 3%. Use of genotypes of high-profile cows improves reliability in ssGEBV and msGEBV for bulls., (Copyright © 2013 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

Reliability of predictions from single-step genomic BLUP (ssGBLUP) can be calculated by matrix inversion, but that is not feasible for large data sets. Two methods of approximating reliability were developed based on the decomposition of a function of reliability into contributions from records, pedigrees, and genotypes. Those contributions can be expressed in record or daughter equivalents. The first approximation method involved inversion of a matrix that contains inverses of the genomic relationship matrix and the pedigree relationship matrix for genotyped animals. The second approximation method involved only the diagonal elements of those inverses. The 2 approximation methods were tested with a simulated data set. The correlations between ssGBLUP and approximated contributions from genomic information were 0.92 for the first approximation method and 0.56 for the second approximation method; contributions were inflated by 62 and 258%, respectively. The respective correlations for reliabilities were 0.98 and 0.72. After empirical correction for inflation, those correlations increased to 0.99 and 0.89. Approximations of reliabilities of predictions by ssGBLUP are accurate and computationally feasible for populations with up to 100,000 genotyped animals. A critical part of the approximations is quality control of information from single nucleotide polymorphisms and proper scaling of the genomic relationship matrix., (Copyright © 2013 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

Background: Artificial insemination and genetic selection are major factors contributing to population stratification in dairy cattle. In this study, we analyzed the effect of sample stratification and the effect of stratification correction on results of a dairy genome-wide association study (GWAS). Three methods for stratification correction were used: the efficient mixed-model association expedited (EMMAX) method accounting for correlation among all individuals, a generalized least squares (GLS) method based on half-sib intraclass correlation, and a principal component analysis (PCA) approach., Results: Historical pedigree data revealed that the 1,654 contemporary cows in the GWAS were all related when traced through approximately 10-15 generations of ancestors. Genome and phenotype stratifications had a striking overlap with the half-sib structure. A large elite half-sib family of cows contributed to the detection of favorable alleles that had low frequencies in the general population and high frequencies in the elite cows and contributed to the detection of X chromosome effects. All three methods for stratification correction reduced the number of significant effects. EMMAX method had the most severe reduction in the number of significant effects, and the PCA method using 20 principal components and GLS had similar significance levels. Removal of the elite cows from the analysis without using stratification correction removed many effects that were also removed by the three methods for stratification correction, indicating that stratification correction could have removed some true effects due to the elite cows. SNP effects with good consensus between different methods and effect size distributions from USDA's Holstein genomic evaluation included the DGAT1-NIBP region of BTA14 for production traits, a SNP 45kb upstream from PIGY on BTA6 and two SNPs in NIBP on BTA14 for protein percentage. However, most of these consensus effects had similar frequencies in the elite and average cows., Conclusions: Genetic selection and extensive use of artificial insemination contributed to overlapped genome, pedigree and phenotype stratifications. The presence of an elite cluster of cows was related to the detection of rare favorable alleles that had high frequencies in the elite cluster and low frequencies in the remaining cows. Methods for stratification correction could have removed some true effects associated with genetic selection.

The study documents the procedures used to estimate genetic correlations among countries for overall conformation (OCS), overall udder (OUS), overall feet and legs (OFL), and body condition score (BCS) of Holstein sires. Major differences in traits definition are discussed, in addition to the use of international breeding values (IBV) among countries involved in international genetic evaluations, and similarities among countries through hierarchical clustering. Data were available for populations from 20 countries for OCS and OUS, 18 populations for OFL, and 11 populations for BCS. The IBV for overall traits and BCS were calculated using a multi-trait across-country evaluation model. Distance measures, obtained from genetic correlations, were used as input values in the cluster analysis. Results from surveys sent to countries participating in international genetic evaluation for conformation traits showed that different ways of defining traits are used: the overall traits were either computed from linear or composite traits or defined as general characteristics. For BCS, populations were divided into 2 groups: one scored and evaluated BCS, and one used a best predictor. In general, populations were well connected except for Estonia and French Red Holstein. The average number of common bulls for the overall traits ranged from 19 (OCS and OUS of French Red Holstein) to 514 (OFL of United States), and for BCS from 17 (French Red Holstein) to 413 (the Netherlands). The average genetic correlation (range) across countries was 0.75 (0.35 to 0.95), 0.80 (0.41 to 0.95), and 0.68 (0.12 to 0.89) for OCS, OUS, and OFL, respectively. Genetic correlations among countries that used angularity as best predictor for BCS and countries that scored BCS were negative. The cluster analysis provided a clear picture of the countries distances; differences were due to trait definition, trait composition, and weights in overall traits, genetic ties, and genotype by environment interactions. Harmonization of trait definition and increasing genetic ties could improve genetic correlations across countries and reduce the distances. In each national selection index, all countries, except Estonia and New Zealand, included at least one overall trait, whereas none included BCS. Out of 18 countries, 9 have started genomic evaluation of conformation traits. The first were Canada, France, New Zealand, and United States in 2009, followed by Switzerland, Germany, and the Netherlands in 2010, and Australia and Denmark-Finland-Sweden (joint evaluation) in 2011. Six countries are planning to start soon., (Copyright © 2012 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

The objective of this study was to quantify the gains in genetic potential of replacement females that could be achieved by using genomic testing to facilitate selection and culling decisions on commercial dairy farms. Data were simulated for 100 commercial dairy herds, each with 1,850 cows, heifers, and calves. Parameters of the simulation were based on the US Holstein population, and assumed reliabilities of traditional and genomic predictions matched reliabilities of animals that have been genotyped to date. Selection of the top 10, 20, 30, …, 90% of animals within each age group was based on parent averages and predicted transmitting abilities with or without genomic testing of all animals or subsets of animals that had been presorted by traditional predictions. Average gains in lifetime net merit breeding value of selected females due to genomic testing, minus prorated costs of genotyping the animals and their unselected contemporaries, ranged from $28 (top 90% selected) to $259 (top 20% selected) for heifer calves with no pedigrees, $14 (top 90% selected) to $121 (top 10% selected) for heifer calves with known sires, and $7 (top 90% selected) to $87 (top 20% selected) for heifer calves with full pedigrees. In most cases, gains in genetic merit of selected heifer calves far exceeded prorated genotyping costs, and gains were greater for animals with missing or incomplete pedigree information. Gains in genetic merit due to genomic testing were smaller for lactating cows that had phenotypic records, and in many cases, these gains barely exceeded or failed to exceed genotyping costs. Strategies based on selective genotyping of the top, middle, or bottom 50% of animals after presorting by traditional parent averages or predicted transmitting abilities were cost effective, particularly when pedigrees or phenotypes were available and a relatively small proportion of animals were to be selected or culled. Based on these results, it appears that routine genotyping of heifer calves or yearling heifers can be a cost-effective strategy for enhancing the genetic level of replacement females on commercial dairy farms. Increasing the accuracy of predicted breeding values for young females with genomic testing might lead to synergies with other management tools and strategies, such as propagating genetically superior females using advanced reproductive technologies or selling excess females that were generated by the use of sex-enhanced semen., (Copyright © 2012 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

Background: Genome-wide association analysis is a powerful tool for annotating phenotypic effects on the genome and knowledge of genes and chromosomal regions associated with dairy phenotypes is useful for genome and gene-based selection. Here, we report results of a genome-wide analysis of predicted transmitting ability (PTA) of 31 production, health, reproduction and body conformation traits in contemporary Holstein cows., Results: Genome-wide association analysis identified a number of candidate genes and chromosome regions associated with 31 dairy traits in contemporary U.S. Holstein cows. Highly significant genes and chromosome regions include: BTA13's GNAS region for milk, fat and protein yields; BTA7's INSR region and BTAX's LOC520057 and GRIA3 for daughter pregnancy rate, somatic cell score and productive life; BTA2's LRP1B for somatic cell score; BTA14's DGAT1-NIBP region for fat percentage; BTA1's FKBP2 for protein yields and percentage, BTA26's MGMT and BTA6's PDGFRA for protein percentage; BTA18's 53.9-58.7 Mb region for service-sire and daughter calving ease and service-sire stillbirth; BTA18's PGLYRP1-IGFL1 region for a large number of traits; BTA18's LOC787057 for service-sire stillbirth and daughter calving ease; BTA15's CD82, BTA23's DST and the MOCS1-LRFN2 region for daughter stillbirth; and BTAX's LOC520057 and GRIA3 for daughter pregnancy rate. For body conformation traits, BTA11, BTAX, BTA10, BTA5, and BTA26 had the largest concentrations of SNP effects, and PHKA2 of BTAX and REN of BTA16 had the most significant effects for body size traits. For body shape traits, BTAX, BTA19 and BTA3 were most significant. Udder traits were affected by BTA16, BTA22, BTAX, BTA2, BTA10, BTA11, BTA20, BTA22 and BTA25, teat traits were affected by BTA6, BTA7, BTA9, BTA16, BTA11, BTA26 and BTA17, and feet/legs traits were affected by BTA11, BTA13, BTA18, BTA20, and BTA26., Conclusions: Genome-wide association analysis identified a number of genes and chromosome regions associated with 31 production, health, reproduction and body conformation traits in contemporary Holstein cows. The results provide useful information for annotating phenotypic effects on the dairy genome and for building consensus of dairy QTL effects.

Currently, the USDA uses a single-trait (ST) model with several intermediate steps to obtain genomic evaluations for US Holsteins. In this study, genomic evaluations for 18 linear type traits were obtained with a multiple-trait (MT) model using a unified single-step procedure. The phenotypic type data on up to 18 traits were available for 4,813,726 Holsteins, and single nucleotide polymorphism markers from the Illumina BovineSNP50 genotyping Beadchip (Illumina Inc., San Diego, CA) were available on 17,293 bulls. Genomic predictions were computed with several genomic relationship matrices (G) that assumed different allele frequencies: equal, base, current, and current scaled. Computations were carried out with ST and MT models. Procedures were compared by coefficients of determination (R(2)) and regression of 2004 prediction of bulls with no daughters in 2004 on daughter deviations of those bulls in 2009. Predictions for 2004 also included parent averages without the use of genomic information. The R(2) for parent averages ranged from 10 to 34% for ST models and from 12 to 35% for MT models. The average R(2) for all G were 34 and 37% for ST and MT models, respectively. All of the regression coefficients were <1.0, indicating that estimated breeding values in 2009 of 1,307 genotyped young bulls' parents tended to be biased. The average regression coefficients ranged from 0.74 to 0.79 and from 0.75 to 0.80 for ST and MT models, respectively. When the weight for the inverse of the numerator relationship matrix (A(-1)) for genotyped animals was reduced from 1 to 0.7, R(2) remained almost identical while the regression coefficients increased by 0.11-0.26 and 0.12-0.23 for ST and MT models, respectively. The ST models required about 5s per iteration, whereas MT models required 3 (6) min per iteration for the regular (genomic) model. The MT single-step approach is feasible for 18 linear type traits in US Holstein cattle. Accuracy for genomic evaluation increases when switching ST models to MT models. Inflation of genomic evaluations for young bulls could be reduced by choosing a small weight for the A(-1) for genotyped bulls., (Copyright © 2011 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

A national data set of artificial inseminations in US Holsteins was used to obtain genetic evaluations for conception rate (CR). The objective of this study was to investigate the feasibility and resulting accuracy from using all available phenotypic, pedigree, and genomic information. Evaluations were performed by regular BLUP or by BLUP with the traditional pedigree and genomic relationships combined in a unified single-step procedure (SSP). Genetic parameters of CR in the first 3 parities were estimated with data from New York State only. Heritability estimates were around 2% and genetic correlations between CR in different parities were >0.73. The R(2) obtained with the SSP were almost twice as large as those achieved with regular BLUP. Computing the SSP took 2h, and it was 33% slower than a regular BLUP. A multiple-trait evaluation of CR using the SSP is both possible and advantageous., (Copyright © 2011 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

The purpose of this study was to find ways of reducing changes of sire predicted transmitting ability for type's final scores (PTATs) from the first to second crop of daughters. The PTATs were estimated from two datasets: D01 (scores recorded up to 2001) and D05 (scores recorded up to 2005). The PTAT changes were calculated as the difference between the evaluations based on D01 and D05. The PTATs were adjusted to a common genetic base of all evaluated cows born in 1995. The single-trait (ST) animal model included the fixed effects of the herd-year-season-classifier, age by year group at classification, stage of lactation at classification, registry status of animals, and additive genetic and permanent environment random effects. Unknown parent groups (UPGs) were defined based on every other birth year starting from 1972. Modifications to the ST model included the usage of a single record per cow, separate UPGs for first and second crop daughters, separate UPGs for sires and dams, and deepened pedigrees for dams with missing phenotypic records. Also, the multiple-trait (MT) model treated records of registered and grade cows as correlated traits. The mean PTAT change, for all of the sires, was close to zero in all of the models analyzed. The estimated mean PTAT change for 145 sires with 40 to 100 first crop and ≥ 200 second crop daughters was -0.33, -0.20, -0.13, -0.28, and -0.12 with ST, only first records, only last records, updated pedigrees, and allowing separate parent groups (PGs) for sires and dams after updating the pedigrees, respectively. The percentages of sires showing PTAT decline were reduced from 74.5 (with ST) to 57.3 by using only the last records of cows, and to 56.4 by allowing separate UPGs for sires and dams after updating the pedigrees. Though updating of the pedigrees alone was not effective, separate UPGs for sires together with additional pedigree was helpful in reducing the bias.

The first national single-step, full-information (phenotype, pedigree, and marker genotype) genetic evaluation was developed for final score of US Holsteins. Data included final scores recorded from 1955 to 2009 for 6,232,548 Holsteins cows. BovineSNP50 (Illumina, San Diego, CA) genotypes from the Cooperative Dairy DNA Repository (Beltsville, MD) were available for 6,508 bulls. Three analyses used a repeatability animal model as currently used for the national US evaluation. The first 2 analyses used final scores recorded up to 2004. The first analysis used only a pedigree-based relationship matrix. The second analysis used a relationship matrix based on both pedigree and genomic information (single-step approach). The third analysis used the complete data set and only the pedigree-based relationship matrix. The fourth analysis used predictions from the first analysis (final scores up to 2004 and only a pedigree-based relationship matrix) and prediction using a genomic based matrix to obtain genetic evaluation (multiple-step approach). Different allele frequencies were tested in construction of the genomic relationship matrix. Coefficients of determination between predictions of young bulls from parent average, single-step, and multiple-step approaches and their 2009 daughter deviations were 0.24, 0.37 to 0.41, and 0.40, respectively. The highest coefficient of determination for a single-step approach was observed when using a genomic relationship matrix with assumed allele frequencies of 0.5. Coefficients for regression of 2009 daughter deviations on parent-average, single-step, and multiple-step predictions were 0.76, 0.68 to 0.79, and 0.86, respectively, which indicated some inflation of predictions. The single-step regression coefficient could be increased up to 0.92 by scaling differences between the genomic and pedigree-based relationship matrices with little loss in accuracy of prediction. One complete evaluation took about 2h of computing time and 2.7 gigabytes of memory. Computing times for single-step analyses were slightly longer (2%) than for pedigree-based analysis. A national single-step genetic evaluation with the pedigree relationship matrix augmented with genomic information provided genomic predictions with accuracy and bias comparable to multiple-step procedures and could account for any population or data structure. Advantages of single-step evaluations should increase in the future when animals are pre-selected on genotypes., (Copyright 2010 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

The purpose of this study was to estimate trends in conception rate (CR) of Holsteins in the southeastern United States over time across month by milk production level and month by days in milk (DIM) subclasses. Data were obtained from Dairy Records Management Systems (Raleigh, NC) and included service records from 10 states (Virginia, Kentucky, North Carolina, South Carolina, Tennessee, Georgia, Florida, Alabama, Mississippi, and Louisiana). After eliminating records with lactation >1 and uncertain and extreme records (records without calving or birth date, with days to service after calving <21 or >250, or without next calving date), the final data set included 827,802 artificial insemination service records for 424,513 cows born from 1985 to 2000, and in 2,953 herds. Effects included in the model were year of birth (1985 to 1989, 1990 to 1994, 1995 to 2000), DIM class, milk production level (high, medium, low based on SD), service month, the covariate of cow age at calving, and 2- and 3-way interactions. Over time, an increase was observed for milk production and an overall decline in CR occurred. Examination of month by milk production subclass least squares means showed that in cool months (November to April) the deterioration of CR over time was small for low and medium milk production cows and virtually none for high-producing cows. However, in other months (May to June), there was a large decline over time in CR for cows in all milk production level subclasses. The trends in CR by DIM subclasses were examined for the months of February, May, June, and August. There was a general increase in CR with increasing DIM for all months within all birth-year groups. The months of February and August were somewhat similar for CR up to 175 DIM for the different birth-year groups. Much larger differences over time were observed for the months of May and June, and it appeared that for these 2 mo, cows in recent periods did not return to the same level of performance as cows in earlier periods. It may be that there has been a decline over time in the ability of cows to handle the onset of heat stress or the switch to pasture-based management systems.

The objective of this study was to estimate genetic correlations between conception rates (CR) and test-day (TD) milk yields in Holsteins for different days in milk (DIM) in small and large herds. The data included 217,213 first-parity service records of 94,984 cows in New York State between 1999 and 2003. The CR was defined as the outcome of a single insemination. Conception rate and TD milk were analyzed using a series of threshold-linear models with fixed effects that included herd-test-date for TD milk and herd-year for CR, age, service month, cubic regressions on DIM using Legendre polynomials and with random effects that included herd x sire interaction, sire additive genetic and permanent environments with quadratic random regressions on DIM, service sire for CR, and residual. Variance components were estimated using a Bayesian method via Gibbs sampling. Herds were categorized into small (< or =80 cows) and large operations. Large herds produced a higher TD milk (34.0 vs. 30.8 kg), had lower CR (29.5 vs. 34.4%), began breeding earlier (75 vs. 92 d to first service), and had fewer days open (138 vs. 145 d). The average CR was 20% at 50 DIM, increased to about 38% at DIM 100, and then leveled off. Estimated genetic correlations between CR and TD milk stayed around -0.15 for small herds but changed from positive (0.3) at 60 DIM to negative (-0.3) at 120 DIM for large herds. Genetic correlations for CR between small and large herds were highest at 80 DIM and lowest at 140 DIM. The chi-square test showed that the frequency of service records was significantly different during a given week for 71% of large herds and for 15% of small herds, suggesting more timed artificial insemination services in large herds. For the top 15% of cows for milk, fertility peaked around 100 DIM in large herds and at around 100 and 170 DIM in small herds. It seems that optimum breeding practices in large herds of breeding cows earlier are already followed.

The purpose of this study was to investigate the compounded impact on conception rates (CR) of the effects of milk production, service month, and days in milk (DIM) by using recent artificial insemination records of Holsteins in New York (NY) and Georgia (GA). Dairy Herd Improvement records were obtained from Dairy Records Management Systems in Raleigh, North Carolina. After removing records with lactations >1 and uncertain and extreme records (records without a calving or birth date, with days to service after calving of <21 or >250, and without the next calving date), the final data set comprised 298,015 service records for 160,879 cows and 23,366 service records for 12,184 cows in NY and GA, respectively, from 2000 to 2003. The analytical model included DIM class, milk-production level, service month, the covariate of cow's age at calving, and all 2-way interactions. The 2 states were analyzed separately. In general across the 2 states, CR declined as milk production increased, and CR declined during the hottest months. Conception rate was similar in NY and GA, at approximately 55% from December to April. In NY, CR declined by approximately 10% in May and June and mostly recovered by July. In GA, the CR started declining in May, bottomed at 31% in September, and did not recover until December. The difference in CR between high- and low-producing cows was 7% in NY and 6% in GA. That difference was the strongest from June to July in GA (15%) and was more uniform in NY. The increase in CR with increasing DIM varied across service season. The CR was nearly flat from 50 to 125 DIM in NY for all seasons, except for a large increasing trend in spring. In GA, there was also an increasing trend in fall. Conception rates were similar in NY and GA between December and May, and were strongly influenced by heat stress in GA from June to November. A decline in CR for reasons other than heat stress was present in both states in late spring. High production resulted in a faster decline of the CR in GA under heat stress. Models analyzing service records should include the DIM x season x region interaction.

Heat stress was evaluated as a factor in differences between regional evaluations for milk yield in the United States. The national data set (NA) consisted of 56 million first-parity, test-day milk yields on 6 million Holsteins. The Northeastern subset (NE) included 12.5 million records on 1.3 million first-calved heifers from 8 states, and the Southeastern subset (SE) included 3.5 million records on 0.4 million heifers from 11 states. Climatic data were available from 202 public weather stations. Each herd was assigned to the nearest weather station. Average daily temperature-humidity index (mean THI) 3 d before test date was used as an indicator of heat stress. Two test-day repeatability models were implemented. Effects included in both models were herd-test date, age at calving class, frequency of milking, days in milk x season class, additive genetic (regular breeding value) and permanent environmental effects. Additionally, the second model included random regressions on degrees of heat stress (t = max[0, mean THI - 72]) for additive genetic (breeding value for heat tolerance) and permanent environmental effects. Both models were fitted with the national and regional data sets. Correlations involved estimated breeding values (EBV) from SE and NE for sires with >or=100 and >or=300 daughters in each region. When heat stress was ignored (first model) the correlations of regular EBV between SE and NE for sires with >or=100 (>or=300) daughters were 0.85 (0.87). When heat stress was considered (second model), the correlation increased by up to 0.01. The correlations of heat stress EBV between NE and SE for sires with >or=100 (>or=300, >or=700) daughters were 0.58 (0.72, 0.81). Evaluations for heat tolerance were similar in cooler and hotter regions for high-reliability sires. Heat stress as modeled explains only a small amount of regional differences, partly because test-day records depict only snapshots of heat stress.

Emphasis by dairy producers on various yield and fitness traits when culling cows was documented for US Holstein calvings since 1982. Least squares differences between cows retained for additional parities and those culled were estimated for milk, fat, and protein yields; somatic cell score (SCS); days open (DO); dystocia score (DS), final score (FS), and 14 type traits. Compared with cows culled during first lactation, superiority for first-parity milk yield was 569 to 1,175 kg for cows with 2 lactations, 642 to 1,283 kg for cows with > or =2 lactations, 710 to 1,350 kg for cows with 3 lactations, and 663 to 1,331 kg for cows with > or =4 lactations. Cows retained for > or =2 lactations had first-parity SCS that were 0.34 to 0.62 lower (more favorable) than those of cows culled during first lactation; first-parity SCS for cows retained for 3 or > or =4 lactations were even more favorable than those of cows with 1 or 2 lactations. The negative genetic relationship between yield and fertility contributed to increased DO as selection for higher milk yield persisted across time despite considerable preference for early conception when culling cows. In 1982, cows retained in the herd for 2, 3, and > or =4 lactations conceived earlier during first lactation (19, 17, and 23 fewer DO, respectively) than those culled during first lactation; those differences had increased to 34, 41, and 52 fewer DO by 2000. Although DS has a negative relationship with survival, first-parity DS were only slightly lower (by 0.10 to 0.14) for survivors than for cows culled during first lactation. Cows retained for > or =2 lactations had greater first-parity FS by 1.4 to 1.9 points than those culled during first lactation. On a standardized basis, the most intense selection during first lactation was for milk and protein yields with less for fat (74 to 86% of that for milk), DO (18 to 74%), FS (22 to 38%), SCS (19 to 37%), and DS (7 to 15%). Producers continued to emphasize the same traits when culling during second and third lactations. Trait priority by producers during culling could aid in setting trait emphasis when selecting bulls for progeny test and could also be useful in developing software for index-based culling guides.

Data included 392,800 records for cows born between 1995 and 1997. Traits analyzed were milk, fat, and protein yields, somatic cell score, days open (DO), 18 linear type traits, final score, and several measures of longevity. Productive life (PL) was defined as the total number of days in milk up to 84 mo of age with a restriction of 305, 500, or 999 d per lactation (PL(305), PL(500), or PL(999), respectively). Herd life was defined as the total number of days from the first calving date to the last (culling) date. A multiple-trait sire model including the effects of registration status, herd-year, age group, month of calving and stage of lactation, sire, and residual was used for parameter estimation. The average duration of the first lactation was 365 d for survivors and 386 d for culled cows. Lactation lengths for the survivors in the next 3 parities all exceeded 330 d. Heritability estimates of between 0.08 and 0.10 were obtained for all definitions of longevity. As maximum recordable PL was increased from 305 to 999 d per lactation, the genetic correlations with milk production increased (from -0.11 to +0.14) and with DO decreased (-0.62 to -0.27). Formulas for an indirect prediction of PL from correlated traits were developed. As maximum PL per lactation was increased, little change in the weights used to predict the various measures of PL, with the exception of DO was found. As the currently used value of PL(305) does not properly account for the longer lactation lengths that are routinely occurring with today's cows, PL with longer lactations may be preferable in routine evaluation.

The objectives of this study were to assess differences in the heritability of type (conformation) traits between herds that differ in mean final score and completeness of pedigree and performance data and to estimate genetic correlations among these environments. Measurement of subjective characteristics, such as conformation traits, may be more difficult in herds with poor management conditions, and genetic evaluation of sires using data from such herds could lead to inaccurate selection decisions. Furthermore, missing pedigree data is a significant problem in many herds, and a lack of phenotypic data from maternal relatives may reduce the effectiveness of animal model evaluation systems. These hypotheses were examined using type classification scores of 1,728,836 first-parity Holstein cows (from 54,223 sires) that calved from 1993 to 2002 in 24,207 US dairy herds. These data included 480,927 records from progeny test daughters, but only 254,891 (47%) were from dams that had valid sire identification, and only 132,953 (28%) were from dams that had also been classified. Herds were grouped into quartiles by mean classification score, percentage of known maternal grandsires, and percentage of classified dams. Estimated heritability of final score was 0.20 in herds with mean score <74.5, 0.17 in herds with <25% known maternal grandsires, and 0.19 in herds with <18% classified dams. Conversely, estimates were 0.39 in herds with mean score >78.7, 0.35 in herds with 100% known maternal grandsires, and 0.37 in herds with >71% classified dams. Estimated genetic correlations between quartiles ranged from 0.86 to 0.95. Based on this study, it appears that improvements in animal identification and data collection in progeny test herds would lead to greater accuracy and stability of genetic evaluations for conformation traits in US Holstein cattle.

The objectives of this study were to estimate genetic correlations among body condition scores (BCS) from various sources, dairy form, and measures of cow health. Body condition score and dairy form evaluated during routine type appraisal was obtained from the Holstein Association USA, Inc. A second set of BCS was obtained from Dairy Records Managements Systems (DRMS) and was recorded by producers that use PCDART dairy management software. Disease observations were obtained from recorded veterinarian treatments in several dairy herds in the United States. Estimated breeding values for diseases in Denmark were also obtained. Genetic correlations among BCS, dairy form, and cow health traits in the United States were generated with sire models. Models included fixed effects for age, DIM, and contemporary group. Random effects included sire, permanent environment, herd-year season for health traits, and error. Predicted transmitting abilities (PTA) for BCS and dairy form were correlated with estimated breeding values for disease in Denmark. The genetic correlation estimate between BCS from DRMS and BCS from the Holstein Association USA, Inc., was 0.85. The genetic correlation estimate between BCS and a composite of all diseases in the United States was -0.79, and PTA for BCS was favorably correlated with an index of resistance to disease other than mastitis in Denmark (0.27). Dairy form was positively correlated with a composite of all diseases in the United States (0.85) and was unfavorably correlated with an index for resistance to disease other than mastitis in Denmark (-0.29). Adjustment for protein yield PTA had a minimal affect on correlations between PTA for BCS or dairy form and disease in Denmark. Selection for higher body condition or lower dairy form with continued selection for yield may slow deterioration in cow health as a correlated response to selection for increased yield.

The objectives of this study were to estimate genetic correlations among body condition score (BCS), dairy form, milk yield, and days open in US Holsteins and investigate the potential of using BCS or dairy form evaluations as early indicators of days open. Dairy form and BCS obtained from the Holstein Association USA, Inc., were merged with mature equivalents (ME) for milk yields and days open data from AIPL-USDA. Cows were required to be classified between 24 and 60 mo of age, before 335 d in milk (DIM) and have ME milk of at least 4537 kg. A minimum of 20 daughters per sire and 10 cows per herd-classification visit (HV) or herd-year-season of calving (HYS) were required. The final data set included 159,700 records. Heritabilities and correlations among dairy form, BCS, milk yield, and days open were estimated with multiple trait sire models. Fixed effects included age at classification for dairy form and BCS, age at calving for milk yield, HV for dairy form and BCS, HYS for milk yield and days open, DIM within lactation group for dairy form and BCS and lactation group for milk yield and days open. Correlations among dairy form, BCS, and days open were generated with and without a ME milk covariable. Correlations between ME milk and days open were generated with and without covariables for dairy form or BCS. Random effects included sire and error. The genetic correlation estimates of days open with dairy form, BCS, and ME milk were 0.48, -0.30, and 0.38, respectively. The genetic correlation estimate between days open and dairy form was 0.38 after adjustment for ME milk, whereas the genetic correlation between days open and BCS was -0.24 after adjustment for ME milk. Combining dairy form evaluations with days open evaluations for 19 recently proven bulls resulted in an average increase of 0.06 for reliability of days open evaluations. The addition of BCS evaluations did not increase reliability when dairy form observations were available.

Genetic correlations among milk, fat, and protein yields; body size composite (BSC); udder composite (UDC); and productive life (PL) in Holsteins were investigated over time. The data set contained 25,280 records of cows born in Wisconsin between 1979 and 1993. The multiple trait random regression (MT-RR) animal model included registration status, herd-year, age group, and stage of lactation as fixed effects; additive genetic effects with random regressions (RR) on year of birth using the first-order Legendre polynomial; and residual effects. Heterogeneous residual variances were considered in the model. Estimates of variance components and genetic correlations among traits from MT-RR were compared with those estimated with a multiple trait interval (MT-I) model, which assumed that every 3-yr interval was a separate trait and included the same effects as in the MT-RR model except for the RR. Genetic correlations estimated with MT-RR and MT-I models over time among all traits were compared with correlations among breeding values predicted with the single trait (ST) model without RR. Correlations among breeding values predicted with MT-RR, ST, and MT models were also calculated. Additive genetic and residual variances for all traits except PL increased over time; those for PL were constant. As a result, heritability estimates had no significant changes during the 15 yr. Genetic correlations of PL with milk, fat, protein, and BSC declined to zero or negative; those with UDC remained positive. Correlations among breeding values predicted with ST, MT, and MT-RR models were relatively high for all traits except PL. Genetic correlations between PL and other traits varied over time, with some correlations changing sign. For accurate indirect prediction of PL from other traits, the genetic correlations among the traits need to be re-estimated periodically.

Phenotypic and genetic performance of US Holstein clones from embryo splitting (ETS) and nuclear transfer (ETN) was documented for yield and fitness traits. Holstein Association USA has registered 2319 ETS and 215 ETN clones. The number of male clones has decreased. Animals selected for cloning were slightly superior genetically to population mean for yield traits. For females, mean superiority of pedigree merit of ETS clones was 186 kg for milk, 9 kg for fat, and 7 kg for protein compared with the population for the same birth year; for ETN clones, superiority to the population was 165, 10, and 8 kg. Advantage in pedigree merit for male clones generally was slightly greater. The small pedigree advantage for female clones of <1 standard deviation above breed mean indicates that selection of animals to clone was not based primarily on yield. Yield deviations were lower for ETS clones than for their full siblings, which indicates a possible impact of the technology on performance. Yields were lower for ETN clones than for their noncloned full siblings, but differences were small and based on small numbers of clones. Milk composition for cloned cows was not different from that for the population. Estimated genetic merit based on daughter yield was more similar for male clone pairs with apparent identical genotype than for clone pairs from the same biotechnology but nonidentical as confirmed by blood typing. For ETS clones with confirmed identical genotypes, identical genetic merit should be assigned.

The objectives of the current study were to investigate the relationship between body condition score (BCS) and dairy form and changes in genetic parameters for BCS and dairy form within and across lactations and age. Body condition score and dairy form were obtained from the Holstein Association USA, Inc. Records were edited to include those cows classified between 24 and 60 mo of age and between 0 and 335 d in milk (DIM). A minimum of 20 daughters per sire and 15 cows per herd-classification visit were required. The dataset consisted of 135,178 records from 119,215 cows. Repeatability, multiple trait, and random regression models were used to analyze the data. All models included fixed effects for herd-classification visit, age within lactations 1, 2, and 3 or higher, and 5th-order polynomials for DIM. Random effects included sire and permanent environment for all models. Random regression models included age at classification nested within sire or DIM and lactation number nested within sire. Genetic variance for both BCS and dairy form was lowest in early lactation and highest in midlactation. Genetic correlations within and across lactations were high. The genetic correlation between DIM 0 in lactation 1 and DIM 305 in lactation 3 was estimated to be 0.77 for BCS and 0.60 for dairy form. The genetic correlation estimate between 30 mo of age at classification and 50 mo of age at classification was 0.94 for both dairy form and BCS. The repeatability models appeared to generate accurate evaluations for BCS or dairy form at all ages and stages of lactation.

The objectives of this study were to estimate the heritability of body condition score (BCS) with data that could be used to generate genetic evaluations for BCS in the US, and to estimate the relationship among BCS, dairy form and selected type traits. Body condition score and linear type trait records were obtained from Holstein Association USA Inc. Because BCS was a new trait for classifiers, scoring distribution and accuracy was not normal. Records from 11 of 29 classifiers were eliminated to generate a data set that should represent BCS data recorded in the future. Edited data included 128,478 records for analysis of first lactation cows and 207,149 records for analysis of all cows. Heritabilities and correlations were estimated with ASREML using sire models. Models included age at calving nested within lactation, 5th order polynomials of DIM, fixed herd-classification visit effects and random sire and error. Genetic correlation estimates were generated between first lactation data that had records from 11 classifiers removed and data with no classifiers removed. Genetic correlation estimates were 0.995 and above between data with and without classifiers removed for scoring distributions, but heritability estimates were higher with the classifiers edited from the data. Heritability estimates for type traits and final score were similar to previously reported estimates. The heritability estimate for BCS was 0.19 for first lactation cows and 0.22 for all cows. The genetic correlation estimate for first lactation cows between BCS and dairy form was -0.73, whereas the genetic correlation estimate between BCS and strength was 0.72. Genetic correlation estimates were nearly identical when cows from all lactations were included in the analyses. Body condition score had a genetic correlation with final score closer to zero (0.08) than correlations of final score with dairy form, stature or strength.

Conformation final scores obtained by the Holstein Association were used in this study. Five subsets were sampled from the original complete data, which contained about 7 million records, and (co)variance components for additive genetic, permanent environmental, and residual effects were estimated with a multiple-trait (MT) model at different ages of cows and with a random regression model. Transmitting abilities were predicted with the random-regression (RR) model and with the repeatability model for the national evaluation, using the entire data set. The RR model included fixed and random regressions on age at classification for additive genetic and permanent environmental effects. Additive genetic and residual variances estimated with the MT model increased with aging. Genetic correlations between age groups decreased with their distance. Heritability and repeatability estimates with the RR model were 0.27 and 0.88 on average, increasing with cow's age. Correlations between transmitting abilities predicted with the RR model at various ages and with the repeatability model were in the range of 0.89 to 0.99. In the 1980s, predicted transmitting abilities (PTA) of sires increased with their daughters' age, but in 1991 and 1995, the PTA decreased. In general, sires whose daughters were evaluated at mature ages tended to have higher PTA. For sires and cows, genetic trends estimated from the repeatability model and from the RR model were similar. The low genetic trend of cows at 60 mo for recent years was due to many grade cows that had only single records and little pedigree information.

The volume and complexity of genetic information is increasing because of new traits and better models. New traits may include reproduction, health, and carcass. More comprehensive models include the test day model in dairy cattle or a growth model in beef cattle. More complex models, which may include nonadditive effects such as inbreeding and dominance, also provide additional information. The amount of information per animal may increase drastically if DNA marker typing becomes routine and quantitative trait loci information is utilized. In many industries, evaluations are run more frequently. They result in faster genetic progress and improved management and marketing opportunities but also in extra costs and information overload. Adopting new technology and making some organizational changes can help realize all the added benefits of the improvements to the genetic evaluation systems at an acceptable cost. Continuous genetic evaluation, in which new records are accepted and breeding values are updated continuously, will relieve time pressures. An online mating system with access to both genetic and marketing information can result in mating recommendations customized for each user. Such a system could utilize inbreeding and dominance information that cannot efficiently be accommodated in the current sire summaries or off-line mating programs. The new systems will require a new organizational approach in which the task of scientists and technicians will not be simply running the evaluations but also providing the research, design, supervision, and maintenance required in the entire system of evaluation, decision making, and distribution.

Genetic evaluations for the length of productive life based on actual DHIA culling data have been available in the US since January 1994. Although substantial genetic variation in productive life exists, the reliability of selection is often low, particularly for recently progeny-tested bulls having daughters that have not yet had an opportunity to be culled. Correlated production and conformation traits, which have higher heritability than productive life and are available earlier in life, may be used to enhance evaluations of productive life for young bulls that have little or no direct culling information available. Genetic correlations between productive life and milk, fat, dairy form, and udder traits ranged from +0.22 to +0.46. The maximum reliability of the indirect prediction of productive life from 16 correlated type and production traits was 0.56, and the maximum reliability from a subset of 10 traits was 0.51. Indirect information about productive life that was derived from type and production traits was combined with actual culling information to increase the total amount of available information for many recently progeny-tested bulls. The procedures described herein for enhancing direct evaluations for the productive life of dairy sires with indirect information about production and type were implemented by the USDA Animal Improvement Programs Laboratory and the Holstein Association USA in July 1994.

The potential of trait-based analysis to detect quantitative trait loci was investigated using blood group polymorphisms as the marker systems and milk and type traits in Holstein cattle as the quantitative traits. Within large half-sib families, animals were ranked on their predicted transmitted abilities or phenotypes, and blood group allele frequencies were compared between the upper and lower 5% tails of the distributions. Genotype frequencies within large families were also examined for evidence of selection. All of the major effects that had previously been detected using linear model analyses were identified by the trait-based analyses of a C blood group effect on rump angle, an L effect on milk yield and composition traits, an S effect on milk fat yield, and a direct effect of the M locus on milk and protein yields. These results provide additional support for the biological validity of these associations and also demonstrate the utility of trait-based analysis for the detection of quantitative trait loci within existing dairy breeding programs. However, just as in the linear model analyses, an analytical strategy should be utilized that allows the identification of the effects that are consistent across environments and genetic backgrounds.

This study addresses the utility of 11 blood groups as selection aids in Holstein breeding schemes and considers issues inherent to the approach of resolving quantitative variation into components that are due to quantitative trait loci. The data consisted of predicted transmitting abilities of 22,614 bulls, first lactation information on 1,924,171 cows, and type scores on 447,800 cows. Linear models were fitted under male half-sib designs, female half-sib designs, and granddaughter designs as well as under the assumption of direct effects of the markers. The evolution of allele frequencies through time was determined, and previous research results were synthesized according to criteria of consistency of biological significance. The inconsistency of results across studies and analytical designs alludes to the importance of the intrinsic nonadditivity of genetic and biological phenomena to quantitative trait locus detection and marker-assisted selection. In our analyses, three associations met the criteria of consistency--a C blood group effect on rump angle, an L effect on milk yield and composition traits, and an S effect on milk fat yield. The M locus appears to be directly associated with effects on milk and protein yields. An enhanced understanding of the biochemical and physiological bases of quantitative genetics should be a long-term objective of this type of genetic analysis.

Differences in breeding values between dominance and additive models were examined theoretically and with field data. Data included 5.2 million records on stature from 3.0 million US Holsteins. The largest full-sib family had 29 animals, and 7% of all animals had at least one full sib. The dominance model, which accounted for dominance covariances, included the following effects: management, age, stage of lactation, permanent environment, animal additive, and parental dominance (one-quarter of dominance variance) as well as a regression coefficient for inbreeding percentage. Two reduced models were also assumed; in the first, the parental dominance effect was removed, and, in the second, the inbreeding regression coefficient was also removed. The correlations between breeding values in the three models were > 0.999, but breeding values of some animals from full-sib families changed > 5 standard deviations of parental dominance. The largest changes were observed for parents with large numbers of full-sib progeny, with limited information from parents, and without individual performance records. On average, the differences were up to four times larger for cows than for bulls and up to five times larger for dams than for sires. The greatest differences in breeding values between the dominance and the additive models were observed for dams with full-sib progeny, female full sibs, and low reliability bulls with full sibs in the extended family. Animals with large amounts of additive information as progeny-tested bulls were influenced little by the inclusion of dominance. Animals with a large proportion of information coming from animals with dominance relationships, such as cows originating via embryo transfer changed the most.

Estimates of variance components were obtained with method R for several additive and dominance models. The data included 301,960 records for first parity and 280,040 records for later parities of Holsteins. The single-record model included effects of management, regression on inbreeding percentage, age at calving, stage of lactation, and additive and dominance effects. The repeatability model included these effects in addition to permanent environment. For the single-record model, estimates were 46% of the total variance for additive variance, 12% of total variance for dominance variance, and -0.06 for the regression coefficient on inbreeding. In the repeatability model, the variance for permanent environment was estimated at 5%; other estimates were similar. When the dominance effect was eliminated, the estimate of the variance for permanent environment increased to 17% for the repeatability model. Elimination of stage of lactation increased regression on inbreeding to 0.09 and the estimate of dominance variance to 17% in the single-record model. The same change increased the estimate of additive variance to 64% for the repeatability model. Elimination of regression on inbreeding or stage of lactation had a small effect on the estimates. The presence or absence of the dominance effect had little influence on additive variance. In the absence of dominance, the estimate of the permanent environment effect included the dominance effect. Estimates of variances with method R are very sensitive to age adjustments. With the adjustments, the estimates of the dominance and additive variances are consistent.

A method was developed to account for heterogeneous phenotypic variance for final type (conformation) score within herd-year-month-classifier subclasses for national genetic evaluations of US Holsteins. Phenotypic variances decreased 3.5-fold as within-subclass means increased from 70 to 89 points, and variances increased slightly over time and with increases in subclass size and proportion of registered animals. A model containing known management characteristics was fitted to phenotypic standard deviations within subclass, and resulting solutions were used to estimate the prior variance for each subclass. Prior estimates and phenotypic variance estimates within subclass were combined using Bayesian methods, resulting in posterior estimates that contained more information than either the prior estimates or the within-subclass statistics. Observations were standardized to a common variance using the mean of the posterior density of the phenotypic standard deviation within subclass. Moderate changes in PTA occurred for low reliability bulls, bulls with many progeny in a single herd, and foreign bulls subjected to positive assortative mating. Significant increases occurred in PTA of superior cows in subclasses with high means. Incorporation of a heterogeneous variance adjustment into national genetic evaluations should increase fairness of selection among cows and young bulls and may increase the rate of genetic gain from female selection.

Advanced reproductive techniques are creating the large numbers of close relatives needed to study gene interactions. Identical triplets, a set of 26 full sisters, a family of 4215 three-quarter sisters (same sire and maternal grandsire), a family of 76,698 half sisters, and 1.6 million granddaughters of Round Oak Rag Apple Elevation now have lactation records. Similarity of closest relatives might be explained by similar nonadditive as well as additive genetic merit. The 23,015 families of full sisters with mean family size of 3 provide nearly as much information about dominance variation as do the 55,779 families of three-quarter sisters with mean family size of 13; the 79 families of clones provide little information by comparison. Hypothetically, REML analysis of all US Holstein data could provide estimates of dominance and additive x additive variance with standard errors approximately 1% of phenotypic variance, but estimates of any higher order interactions would have standard errors greater than 10%. The tilde-hat approximation proved to be incompatible with animal models but was used for sire-maternal grandsire analysis of 765,868 first lactation records. Dominance variance was estimated as 3.5% of phenotypic variance for milk and 3.3% for fat with standard error of 4.2%. With constant data set size, variances are estimated most precisely if family sizes equal 1 plus ratio of within-family to between-family variance. An animal model evaluation including dominance relationships for 581,670 animals was computed, but gene interactions from distant ancestor pairs were ignored. Mating advice and improved additive predictions, especially for clones, could be obtained by including dominance in models.