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Background: In breeding programs, recording large-scale feed intake (FI) data routinely at the individual level is costly and difficult compared with other production traits. An alternative approach could be to record FI at the group level since animals such as pigs are normally housed in groups and fed by a shared feeder. However, to date there have been few investigations about the difference between group- and individual-level FI recorded in different environments. We hypothesized that group- and individual-level FI are genetically correlated but different traits. This study, based on the experiment undertaken in purebred DanBred Landrace (L) boars, was set out to estimate the genetic variances and correlations between group- and individual-level FI using a bivariate random regression model, and to examine to what extent prediction accuracy can be improved by adding information of individual-level FI to group-level FI for animals recorded in groups. For both bivariate and univariate models, single-step genomic best linear unbiased prediction (ssGBLUP) and pedigree-based BLUP (PBLUP) were implemented and compared., Results: The variance components from group-level records and from individual-level records were similar. Heritabilities estimated from group-level FI were lower than those from individual-level FI over the test period. The estimated genetic correlations between group- and individual-level FI based on each test day were on average equal to 0.32 (SD = 0.07), and the estimated genetic correlation for the whole test period was equal to 0.23. Our results demonstrate that by adding information from individual-level FI records to group-level FI records, prediction accuracy increased by 0.018 and 0.032 compared with using group-level FI records only (bivariate vs. univariate model) for PBLUP and ssGBLUP, respectively., Conclusions: Based on the current dataset, our findings support the hypothesis that group- and individual-level FI are different traits. Thus, the differences in FI traits under these two feeding systems need to be taken into consideration in pig breeding programs. Overall, adding information from individual records can improve prediction accuracy for animals with group records.

Records on groups of individuals could be valuable for predicting breeding values when a trait is difficult or costly to measure on single individuals, such as feed intake and egg production. Adding genomic information has shown improvement in the accuracy of genetic evaluation of quantitative traits with individual records. Here, we investigated the value of genomic information for traits with group records. Besides, we investigated the improvement in accuracy of genetic evaluation for group-recorded traits when including information on a correlated trait with individual records. The study was based on a simulated pig population, including three scenarios of group structure and size. The results showed that both the genomic information and a correlated trait increased the accuracy of estimated breeding values (EBVs) for traits with group records. The accuracies of EBV obtained from group records with a size 24 were much lower than those with a size 12. Random assignment of animals to pens led to lower accuracy due to the weaker relationship between individuals within each group. It suggests that group records are valuable for genetic evaluation of a trait that is difficult to record on individuals, and the accuracy of genetic evaluation can be considerably increased using genomic information. Moreover, the genetic evaluation for a trait with group records can be greatly improved using a bivariate model, including correlated traits that are recorded individually. For efficient use of group records in genetic evaluation, relatively small group size and close relationships between individuals within one group are recommended.

Background: Crossbreeding is widely used in pig production because of the benefits of heterosis effects and breed complementarity. Commonly, sire lines are bred for traits such as feed efficiency, growth and meat content, whereas maternal lines are also bred for reproduction and longevity traits, and the resulting three-way crossbred pigs are used for production of meat. The most important genetic basis for heterosis is dominance effects, e.g. removal of inbreeding depression. The aims of this study were to (1) present a modification of a previously developed model with additive, dominance and inbreeding depression genetic effects for analysis of data from a purebred sire line and three-way crossbred pigs; (2) based on this model, present equations for additive genetic variances, additive genetic covariance, and estimated breeding values (EBV) with associated accuracies for purebred and crossbred performances; (3) use the model to analyse four production traits, i.e. ultra-sound recorded backfat thickness (BF), conformation score (CONF), average daily gain (ADG), and feed conversion ratio (FCR), recorded on Danbred Duroc and Danbred Duroc-Landrace-Yorkshire crossbred pigs reared in the same environment; and (4) obtain estimates of genetic parameters, additive genetic correlations between purebred and crossbred performances, and EBV with associated accuracies for purebred and crossbred performances for this data set., Results: Additive genetic correlations (with associated standard errors) between purebred and crossbred performances were equal to 0.96 (0.07), 0.83 (0.16), 0.75 (0.17), and 0.87 (0.18) for BF, CONF, ADG, and FCR, respectively. For BF, ADG, and FCR, the additive genetic variance was smaller for purebred performance than for crossbred performance, but for CONF the reverse was observed. EBV on Duroc boars were more accurate for purebred performance than for crossbred performance for BF, CONF and FCR, but not for ADG., Conclusions: Methodological developments led to equations for genetic (co)variances and EBV with associated accuracies for purebred and crossbred performances in a three-way crossbreeding system. As illustrated by the data analysis, these equations may be useful for implementation of genomic selection in this system.

Background: Records on groups of individuals rather than on single individuals could be valuable for predicting breeding values (BV) of the traits that are difficult or costly to measure individually, such as feed intake in pigs or beef cattle. Here, we present a model, which handles group records from varying group sizes and involves multiple fixed and random effects, for estimating variance components and predicting BV. Moreover, using simulation, we investigated the efficiency of group records for predicting BV in situations with various group sizes and structures, and factors that affect the trait., Results: The results show that the presented model for group records worked well and that variances estimated from group records with varying group sizes were consistent with those estimated from individual records, but with larger standard errors. Ignoring litter and pen effects had very little or no influence on the accuracy of estimated BV (EBV) obtained from group records. However, ignoring litter effects resulted in biased estimates of additive genetic variance and EBV. The presence of litter and pen effects on phenotypes decreased the accuracy of EBV although the prediction model fitted both effects. Having more littermates in the same pen led to a higher accuracy of EBV. The decay of EBV accuracy with increasing group size was more marked for scenarios with litter and pen effects than without. When litters of six individuals were divided into two pens, accuracies of EBV obtained from group records with a size up to 12 (average 9.6) and up to 24 (average 19.2) were 66.6 and 57.6% of those estimated from individual records in the scenario with litter and pen effects on phenotypes. These percentages reached 77.0 and 68.4% in the scenario without litter and pen effects on phenotypes., Conclusions: Our results indicate that the model works appropriately for the analysis of group records from varying group sizes. Using group records for genetic evaluation of traits such as feed intake in pig is feasible and the efficiency of the resulting estimates depends on the size and structure of the groups and on the magnitude of the variances for litter and pen effects.

This study investigated the efficiency of genomic prediction with adding the markers identified by genome-wide association study (GWAS) using a data set of imputed high-density (HD) markers from 54K markers in Chinese Holsteins. Among 3,056 Chinese Holsteins with imputed HD data, 2,401 individuals born before October 1, 2009, were used for GWAS and a reference population for genomic prediction, and the 220 younger cows were used as a validation population. In total, 1,403, 1,536, and 1,383 significant single nucleotide polymorphisms (SNP; false discovery rate at 0.05) associated with conformation final score, mammary system, and feet and legs were identified, respectively. About 2 to 3% genetic variance of 3 traits was explained by these significant SNP. Only a very small proportion of significant SNP identified by GWAS was included in the 54K marker panel. Three new marker sets (54K+) were herein produced by adding significant SNP obtained by linear mixed model for each trait into the 54K marker panel. Genomic breeding values were predicted using a Bayesian variable selection (BVS) model. The accuracies of genomic breeding value by BVS based on the 54K+ data were 2.0 to 5.2% higher than those based on the 54K data. The imputed HD markers yielded 1.4% higher accuracy on average (BVS) than the 54K data. Both the 54K+ and HD data generated lower bias of genomic prediction, and the 54K+ data yielded the lowest bias in all situations. Our results show that the imputed HD data were not very useful for improving the accuracy of genomic prediction and that adding the significant markers derived from the imputed HD marker panel could improve the accuracy of genomic prediction and decrease the bias of genomic prediction., (Copyright © 2018 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

The objective of this study was to evaluate the improvement of the accuracy of estimated breeding values for ability to recycle after calving by using information of genomic markers and phenotypic information of correlated traits. The traits in this study were the interval from calving to first insemination (CFI), based on artificial insemination data, and the interval from calving to first high activity (CFHA), recorded from activity tags, which could better measure ability to recycle after caving. The phenotypic data set included 1,472,313 records from 820,218 cows for CFI, and 36,504 records from 25,733 cows for CFHA. The genomic information was available for 3,159 progeny-tested sires, which were genotyped using Illumina Bovine SNP50 BeadChip (Illumina, San Diego, CA). Heritability estimates were 0.06 for the interval from calving to first insemination and 0.14 for the interval from calving to first high activity, and the genetic correlation between both traits was strong (0.87). Breeding values were obtained using 4 models: conventional single-trait BLUP; conventional multitrait BLUP with pedigree-based relationship matrix; single-trait single-step genomic BLUP; and multitrait single-step genomic BLUP model with joint relationship matrix combining pedigree and genomic information. The results showed that reliabilities of estimated breeding values (EBV) from single-step genomic BLUP models were about 40% higher than those from conventional BLUP models for both traits. Furthermore, using a multitrait model doubled the reliability of breeding values for CFHA, whereas no gain was observed for CFI. The best model was the multitrait single-step genomic BLUP, which resulted in a reliability of EBV 0.19 for CFHA and 0.14 for CFI. The results indicate that even though a relatively small number of records for CFHA were available, with genomic information and using multitrait model, the reliability of EBV for CFHA is acceptable. Thus, it is feasible to include CFHA in Nordic Holstein breeding evaluations to improve fertility performance., (The Authors. Published by the Federation of Animal Science Societies 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/).)

Background: A quantitative trait is controlled both by major variants with large genetic effects and by minor variants with small effects. Genome-wide association studies (GWAS) are an efficient approach to identify quantitative trait loci (QTL), and genomic selection (GS) with high-density single nucleotide polymorphisms (SNPs) can achieve higher accuracy of estimated breeding values than conventional best linear unbiased prediction (BLUP). GWAS and GS address different aspects of quantitative traits, but, as statistical models, they are quite similar in their description of the genetic mechanisms that underlie quantitative traits., Methods: Here, we propose a stepwise linear regression mixed model (StepLMM) to unify GWAS and GS in a single statistical model. First, the variance components of the genomic-BLUP (GBLUP) model are estimated. Then, in the SNP selection step, the linear mixed model (LMM) for GWAS is equivalently transformed into a simple linear regression to improve computation speed, and the most significant SNP is selected and included into the evaluation model. In the SNP dropping step, the SNPs in the evaluation model are tested according to the standard errors of their estimated effects. If non-significant SNPs are present, the least significant one is dropped from the model and variance components are re-estimated. We used extended Bayesian information criteria (eBIC) to evaluate the model optimization, i.e. the model with the smallest eBIC is the final one and includes only significant SNPs., Results: We simulated scenarios with different heritabilities with 100 QTL. StepLMM estimated heritability accurately and mapped QTL precisely. Genomic prediction accuracy was much higher with StepLMM than with GBLUP. The comparison of StepLMM with other GWAS and GS methods based on a dataset from the 16th QTLMAS Workshop showed that StepLMM had medium mapping power, the lowest rate of false positives for QTL mapping, and the highest accuracy for genomic prediction., Conclusions: StepLMM is a combination of GWAS and GBLUP. GWAS and GBLUP are beneficial to each other in a single statistical model, GWAS improves genomic prediction accuracy, while GBLUP increases mapping precision and decreases the rate of false positives of GWAS. StepLMM has a high performance in both GWAS and GS and is feasible for agricultural breeding programs and human genetic studies.

The present study investigated the improvement of prediction reliabilities for 3 production traits in Brazilian Holsteins that had no genotype information by adding information from Nordic and French Holstein bulls that had genotypes. The estimated across-country genetic correlations (ranging from 0.604 to 0.726) indicated that an important genotype by environment interaction exists between Brazilian and Nordic (or Nordic and French) populations. Prediction reliabilities for Brazilian genotyped bulls were greatly increased by including data of Nordic and French bulls, and a 2-trait single-step genomic BLUP performed much better than the corresponding pedigree-based BLUP. However, only a minor improvement in prediction reliabilities was observed in nongenotyped Brazilian cows. The results indicate that although there is a large genotype by environment interaction, inclusion of a foreign reference population can improve accuracy of genetic evaluation for the Brazilian Holstein population. However, a Brazilian reference population is necessary to obtain a more accurate genomic evaluation., (Copyright © 2016 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

Small reference populations limit the accuracy of genomic prediction in numerically small breeds, such like Danish Jersey. The objective of this study was to investigate two approaches to improve genomic prediction by increasing size of reference population in Danish Jersey. The first approach was to include North American Jersey bulls in Danish Jersey reference population. The second was to genotype cows and use them as reference animals. The validation of genomic prediction was carried out on bulls and cows, respectively. In validation on bulls, about 300 Danish bulls (depending on traits) born in 2005 and later were used as validation data, and the reference populations were: (1) about 1050 Danish bulls, (2) about 1050 Danish bulls and about 1150 US bulls. In validation on cows, about 3000 Danish cows from 87 young half-sib families were used as validation data, and the reference populations were: (1) about 1250 Danish bulls, (2) about 1250 Danish bulls and about 1150 US bulls, (3) about 1250 Danish bulls and about 4800 cows, (4) about 1250 Danish bulls, 1150 US bulls and 4800 Danish cows. Genomic best linear unbiased prediction model was used to predict breeding values. De-regressed proofs were used as response variables. In the validation on bulls for eight traits, the joint DK-US bull reference population led to higher reliability of genomic prediction than the DK bull reference population for six traits, but not for fertility and longevity. Averaged over the eight traits, the gain was 3 percentage points. In the validation on cows for six traits (fertility and longevity were not available), the gain from inclusion of US bull in reference population was 6.6 percentage points in average over the six traits, and the gain from inclusion of cows was 8.2 percentage points. However, the gains from cows and US bulls were not accumulative. The total gain of including both US bulls and Danish cows was 10.5 percentage points. The results indicate that sharing reference data and including cows in reference population are efficient approaches to increase reliability of genomic prediction. Therefore, genomic selection is promising for numerically small population.

Background: Commercial pig producers generally use a terminal crossbreeding system with three breeds. Many pig breeding organisations have started to use genomic selection for which genetic evaluation is often done by applying single-step methods for which the pedigree-based additive genetic relationship matrix is replaced by a combined relationship matrix based on both marker genotypes and pedigree. Genomic selection is implemented for purebreds, but it also offers opportunities for incorporating information from crossbreds and selecting for crossbred performance. However, models for genetic evaluation for the three-way crossbreeding system have not been developed., Results: Four-variate models for three-way terminal crossbreeding are presented in which the first three variables contain the records for the three pure breeds and the fourth variable contains the records for the three-way crossbreds. For purebred animals, the models provide breeding values for both purebred and crossbred performances. Heterogeneity of genetic architecture between breeds and genotype by environment interactions are modelled through genetic correlations between these breeding values. Specification of the additive genetic relationships is essential for these models and can be defined either within populations or across populations. Based on these two types of additive genetic relationships, both pedigree-based, marker-based and combined relationships based on both pedigree and marker information are presented. All these models for three-way crossbreeding can be formulated using Kronecker matrix products and therefore fitted using Henderson's mixed model equations and standard animal breeding software., Conclusions: Models for genetic evaluation in the three-way crossbreeding system are presented. They provide estimated breeding values for both purebred and crossbred performances, and can use pedigree-based or marker-based relationships, or combined relationships based on both pedigree and marker information. This provides a framework that allows information from three-way crossbred animals to be incorporated into a genetic evaluation system.

A single-step method allows genetic evaluation using information of phenotypes, pedigree, and markers from genotyped and nongenotyped individuals simultaneously. This paper compared genomic predictions obtained from a single-step BLUP (SSBLUP) method, a genomic BLUP (GBLUP) method, a selection index blending (SELIND) method, and a traditional pedigree-based method (BLUP) for total number of piglets born (TNB), litter size at d 5 after birth (LS5), and mortality rate before d 5 (Mort; including stillbirth) in Danish Landrace and Yorkshire pigs. Data sets of 778,095 litters from 309,362 Landrace sows and 472,001 litters from 190,760 Yorkshire sows were used for the analysis. There were 332,795 Landrace and 207,255 Yorkshire animals in the pedigree data, among which 3,445 Landrace pigs (1,366 boars and 2,079 sows) and 3,372 Yorkshire pigs (1,241 boars and 2,131 sows) were genotyped with the Illumina PorcineSNP60 BeadChip. The results showed that the 3 methods with marker information (SSBLUP, GBLUP, and SELIND) produced more accurate predictions for genotyped animals than the pedigree-based method. For genotyped animals, the average of reliabilities for all traits in both breeds using traditional BLUP was 0.091, which increased to 0.171 w+hen using GBLUP and to 0.179 when using SELIND and further increased to 0.209 when using SSBLUP. Furthermore, the average reliability of EBV for nongenotyped animals was increased from 0.091 for traditional BLUP to 0.105 for the SSBLUP. The results indicate that the SSBLUP is a good approach to practical genomic prediction of litter size and piglet mortality in Danish Landrace and Yorkshire populations.

This study investigated the effect of including Nordic Holsteins in the reference population on the imputation accuracy and prediction accuracy for Chinese Holsteins. The data used in this study include 85 Chinese Holstein bulls genotyped with both 54K chip and 777K (HD) chip, 2862 Chinese cows genotyped with 54K chip, 510 Nordic Holstein bulls genotyped with HD chip, and 4398 Nordic Holstein bulls genotyped with 54K chip and with deregressed proofs for five milk production traits. Based on these data, the accuracy of imputation from 54K to HD marker data and the accuracy of genomic predictions in Chinese Holstein were assessed. The allele correct rate increased around 2.7 and 1.7% in imputation from the 54K to the HD marker data for Chinese Holstein bulls and cows, respectively, when the Nordic HD-genotyped bulls were included in the reference data for imputation. However, the prediction accuracy was improved slightly when using the marker data imputed based on the combined HD reference data, compared with using the marker data imputed based on the Chinese HD reference data only. On the other hand, when using the combined reference population including 4398 Nordic Holstein bulls, the accuracy of genomic predictions increased 6.5 percentage points together with a reduction of prediction bias. The HD markers did not outperform the 54K markers in genomic prediction based on the present data. The results indicate that for Chinese Holsteins, it is necessary to genotype more individuals with 54K chip to increase reference population rather than increasing marker density., (© 2014 Blackwell Verlag GmbH.)

Background: Growth and carcass traits are very important traits for broiler chickens. However, carcass traits can only be measured postmortem. Genomic selection may be a powerful tool for such traits because of its accurate prediction of breeding values of animals without own phenotypic information. This study investigated the efficiency of genomic prediction in Chinese triple-yellow chickens. As a new line, Chinese triple-yellow chicken was developed by cross-breeding and had a small effective population. Two growth traits and three carcass traits were analyzed: body weight at 6 weeks, body weight at 12 weeks, eviscerating percentage, breast muscle percentage and leg muscle percentage., Results: Genomic prediction was assessed using a 4-fold cross-validation procedure for two validation scenarios. In the first scenario, each test data set comprised two half-sib families (family sample) and the rest represented the reference data. In the second scenario, the whole data were randomly divided into four subsets (random sample). In each fold of validation, one subset was used as the test data and the others as the reference data in each single validation. Genomic breeding values were predicted using a genomic best linear unbiased prediction model, a Bayesian least absolute shrinkage and selection operator model, and a Bayesian mixture model with four distributions. The accuracy of genomic estimated breeding value (GEBV) was measured as the correlation between GEBV and the corrected phenotypic value. Using the three models, the correlations ranged from 0.448 to 0.468 for the two growth traits and from 0.176 to 0.255 for the three carcass traits in the family sample scenario, and were between 0.487 and 0.536 for growth traits and between 0.312 and 0.430 for carcass traits in the random sample scenario. The differences in the prediction accuracies between the three models were very small; the Bayesian mixture model was slightly more accurate. According to the results from the random sample scenario, the accuracy of GEBV was 0.197 higher than the conventional pedigree index, averaged over the five traits., Conclusions: The results indicated that genomic selection could greatly improve the accuracy of selection in chickens, compared with conventional selection. Genomic selection for growth and carcass traits in broiler chickens is promising.

This study investigated genomic predictions across Nordic Holstein and Nordic Red using various genomic relationship matrices. Different sources of information, such as consistencies of linkage disequilibrium (LD) phase and marker effects, were used to construct the genomic relationship matrices (G-matrices) across these two breeds. Single-trait genomic best linear unbiased prediction (GBLUP) model and two-trait GBLUP model were used for single-breed and two-breed genomic predictions. The data included 5215 Nordic Holstein bulls and 4361 Nordic Red bulls, which was composed of three populations: Danish Red, Swedish Red and Finnish Ayrshire. The bulls were genotyped with 50 000 SNP chip. Using the two-breed predictions with a joint Nordic Holstein and Nordic Red reference population, accuracies increased slightly for all traits in Nordic Red, but only for some traits in Nordic Holstein. Among the three subpopulations of Nordic Red, accuracies increased more for Danish Red than for Swedish Red and Finnish Ayrshire. This is because closer genetic relationships exist between Danish Red and Nordic Holstein. Among Danish Red, individuals with higher genomic relationship coefficients with Nordic Holstein showed more increased accuracies in the two-breed predictions. Weighting the two-breed G-matrices by LD phase consistencies, marker effects or both did not further improve accuracies of the two-breed predictions., (© 2014 Blackwell Verlag GmbH.)

The main aim of this study was to compare accuracies of imputation and genomic predictions based on single and joint reference populations for Norwegian Red (NRF) and a composite breed (DFS) consisting of Danish Red, Finnish Ayrshire, and Swedish Red. The single nucleotide polymorphism (SNP) data for NRF consisted of 2 data sets: one including 25,000 markers (NRF25K) and the other including 50,000 markers (NRF50K). The NRF25K data set had 2,572 bulls, and the NRF50K data set had 1,128 bulls. Four hundred forty-two bulls were genotyped in both data sets (double-genotyped bulls). The DFS data set (DSF50K) included 50,000 markers of 13,472 individuals, of which around 4,700 were progeny-tested bulls. The NRF25K data set was imputed to 50,000 density using the software Beagle. The average error rate for the imputation of NRF25K decreased slightly from 0.023 to 0.021, and the correlation between observed and imputed genotypes changed from 0.935 to 0.936 when comparing the NRF50K reference and the NRF50K-DFS50K joint reference imputations. A genomic BLUP (GBLUP) model and a Bayesian 4-component mixture model were used to predict genomic breeding values for the NRF and DFS bulls based on the single and joint NRF and DFS reference populations. In the multiple population predictions, accuracies of genomic breeding values increased for the 3 production traits (milk, fat, and protein yields) for both NRF and DFS. Accuracies increased by 6 and 1.3 percentage points, on average, for the NRF and DFS bulls, respectively, using the GBLUP model, and by 9.3 and 1.3 percentage points, on average, using the Bayesian 4-component mixture model. However, accuracies for health or reproduction traits did not increase from the multiple population predictions. Among the 3 DFS populations, Swedish Red gained most in accuracies from the multiple population predictions, presumably because Swedish Red has a closer genetic relationship with NRF than Danish Red and Finnish Ayrshire. The Bayesian 4-component mixture model performed better than the GBLUP model for most production traits for both NRF and DFS, whereas no advantage was found for health or reproduction traits. In general, combining NRF and DFS reference populations was useful in genomic predictions for both the NRF and DFS bulls., (Copyright © 2014 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

Background: For a two-breed crossbreeding system, Wei and van der Werf presented a model for genetic evaluation using information from both purebred and crossbred animals. The model provides breeding values for both purebred and crossbred performances. Genomic evaluation incorporates marker genotypes into a genetic evaluation system. Among popular methods are the so-called single-step methods, in which marker genotypes are incorporated into a traditional animal model by using a combined relationship matrix that extends the marker-based relationship matrix to non-genotyped animals. However, a single-step method for genomic evaluation of both purebred and crossbred performances has not been developed yet., Results: An extension of the Wei and van der Werf model that incorporates genomic information is presented. The extension consists of four steps: (1) the Wei van der Werf model is reformulated using two partial relationship matrices for the two breeds; (2) marker-based partial relationship matrices are constructed; (3) marker-based partial relationship matrices are adjusted to be compatible to pedigree-based partial relationship matrices and (4) combined partial relationship matrices are constructed using information from both pedigree and marker genotypes. The extension of the Wei van der Werf model can be implemented using software that allows inverse covariance matrices in sparse format as input., Conclusions: A method for genomic evaluation of both purebred and crossbred performances was developed for a two-breed crossbreeding system. The method allows information from crossbred animals to be incorporated in a coherent manner for such crossbreeding systems.

Breeding animals can be accurately evaluated using appropriate genomic prediction models, based on marker data and phenotype information. In this study, direct genomic values (DGV) were estimated for 16 traits of Nordic Total Merit (NTM) Index in Nordic Red cattle population using three models and two different response variables. The three models were as follows: a linear mixed model (GBLUP), a Bayesian variable selection model similar to BayesA (BayesA*) and a Bayesian least absolute shrinkage and selection operator model (Bayesian Lasso). The response variables were deregressed proofs (DRP) and conventional estimated breeding values (EBV). The reliability of genomic predictions was measured on bulls in the validation data set as the squared correlation between DGV and DRP divided by the reliability of DRP. Using DRP as response variable, the reliabilities of DGV among the 16 traits ranged from 0.151 to 0.569 (average 0.317) for GBLUP, from 0.152 to 0.576 (average 0.318) for BayesA* and from 0.150 to 0.570 (average 0.320) for Bayesian Lasso. Using EBV as response variable, the reliabilities ranged from 0.159 to 0.580 (average 0.322) for GBLUP, from 0.157 to 0.578 (average 0.319) for BayesA* and from 0.159 to 0.582 (average 0.325) for Bayesian Lasso. In summary, Bayesian Lasso performed slightly better than the other two models, and EBV performed slightly better than DRP as response variable, with regard to prediction reliability of DGV. However, these differences were not statistically significant. Moreover, using EBV as response variable would result in problems with the scale of the resulting DGV and potential problem due to double counting., (© 2013 Blackwell Verlag GmbH.)

The main purpose of this study was to evaluate whether the population structure in Danish Jersey (DJ) known from the history of the breed also is reflected in its genomic structure. This is done by comparing the linkage disequilibrium and persistence of phase for subgroups of Jersey animals with high proportions of Danish (DNK) or United States (USJ) origin. Furthermore, it is investigated whether a model explicitly incorporating breed origin of animals, inferred either through the known pedigree or from SNP marker data, leads to improved genomic predictions compared with a model ignoring breed origin. The study of the population structure incorporated 1,730 genotyped Jersey animals. In total 39,542 SNP markers were included in the analysis. The 1,079 genotyped bulls with de-regressed proof for udder health were used in the analysis for the predictions of the genomic breeding values. A range of random regressions models that included the breed origin were analyzed and compared with a basic genomic model that assumes a homogeneous breed structure. The main finding in this study is that the importation of germplasm from the USJ population is readily reflected in the genomes of modern DJ animals. First, linkage disequilibrium in the group of admixed DJ animals is lower compared with the groups of the original DNK and USJ animals. Second, persistence of linkage disequilibrium phase is not conserved for longer marker distances between animals with mainly Danish or United States origin. Third, the STRUCTURE analysis could retrieve genomic-based breed proportions in alignment to the pedigree-based breed proportions. However, including this population structure in a random regression prediction model did not clearly improve the reliabilities of the genomic predictions compared with a basic genomic model.

Selection for litter size at d 5 after farrowing (LS5) was introduced in 2004 to increase the number of piglets weaned and to reduce piglet mortality in Danish Landrace and Yorkshire. The objective of this study was to investigate selection responses for LS5, total number born (TNB), and mortality [MORT, defined as (TNB - LS5)/TNB] when selection for increasing LS5 was a part of the breeding goal. Data were collected from nucleus herds recorded from 2004 to 2010, including first litters of 42,807 Landrace sows and 33,225 Yorkshire sows. The data were analyzed using a 3-trait animal model of TNB, MORT, and LS5. Significant (co) variances were estimated between the 3 traits in both populations. The heritabilities of TNB, MORT, and LS5 were 0.10, 0.09, and 0.09 in Landrace and 0.12, 0.10, and 0.10 in Yorkshire. The genetic correlations were 0.28 and 0.22 between TNB and MORT, 0.74 and 0.68 between TNB and LS5, and -0.43 and -0.57 between MORT and LS5 in Landrace and Yorkshire, respectively. The results show that the genetic improvement of LS5 was a combination of increased TNB and reduced MORT. During the observation period, the genetic improvement was 1.7 piglets per litter for LS5, 1.3 piglets per litter for TNB, and 4.7% for MORT in Landrace and 2.2 piglets per litter, 1.9 piglets per litter, and 5.9% in Yorkshire. Phenotypic improvement was 1.4 piglets per litter for LS5, 0.3 piglets per litter for TNB, and 7.9% for MORT in Landrace and 2.1 piglets per litter, 1.3 piglets per litter, and 7.6% in Yorkshire. In addition, genetic gain was evaluated in 3 phenotypic groups of TNB, representing the 25% smallest litters, the 50% medium litters, and the 25% largest litters. In all 3 groups, the genetic and phenotypic gains of TNB and LS5 increased, whereas MORT reduced in both populations.

In order to optimize the use of genomic selection in breeding plans, it is essential to have reliable estimates of the genomic breeding values. This study investigated reliabilities of direct genomic values (DGVs) in the Jersey population estimated by three different methods. The validation methods were (i) fivefold cross-validation and (ii) validation on the most recent 3 years of bulls. The reliability of DGV was assessed using squared correlations between DGV and deregressed proofs (DRPs). In the recent 3-year validation model, estimated reliabilities were also used to assess the reliabilities of DGV. The data set consisted of 1003 Danish Jersey bulls with conventional estimated breeding values (EBVs) for 14 different traits included in the Nordic selection index. The bulls were genotyped for Single-nucleotide polymorphism (SNP) markers using the Illumina 54 K chip. A Bayesian method was used to estimate the SNP marker effects. The corrected squared correlations between DGV and DRP were on average across all traits 0.04 higher than the squared correlation between DRP and the pedigree index. This shows that there is a gain in accuracy due to incorporation of marker information compared with parent index pre-selection only. Averaged across traits, the estimates of reliability of DGVs ranged from 0.20 for validation on the most recent 3 years of bulls and up to 0.42 for expected reliabilities. Reliabilities from the cross-validation were on average 0.24. For the individual traits, the reliability varied from 0.12 (direct birth) to 0.39 (milk). Bulls whose sires were included in the reference group had an average reliability of 0.25, whereas the bulls whose sires were not included in the reference group had an average reliability that was 0.05 lower.

This study investigated the accuracy of direct genomic breeding values (DGV) using a genomic BLUP model, genomic enhanced breeding values (GEBV) using a one-step blending approach, and GEBV using a selection index blending approach for 15 traits of Nordic Red Cattle. The data comprised 6,631 bulls of which 4,408 bulls were genotyped using Illumina Bovine SNP50 BeadChip (Illumina, San Diego, CA). To validate reliability of genomic predictions, about 20% of the youngest genotyped bulls were taken as test data set. Deregressed proofs (DRP) were used as response variables for genomic predictions. Reliabilities of genomic predictions in the validation analyses were measured as squared correlations between DRP and genomic predictions corrected for reliability of DRP, based on the bulls in the test data sets. A set of weighting (scaling) factors was used to construct the combined relationship matrix among genotyped and nongenotyped bulls for one-step blending, and to scale DGV and its expected reliability in the selection index blending. Weighting (scaling) factors had a small influence on reliabilities of GEBV, but a large influence on the variation of GEBV. Based on the validation analyses, averaged over the 15 traits, the reliability of DGV for bulls without daughter records was 11.0 percentage points higher than the reliability of conventional pedigree index. Further gain of 0.9 percentage points was achieved by combining information from conventional pedigree index using the selection index blending, and gain of 1.3 percentage points was achieved by combining information of genotyped and nongenotyped bulls simultaneously applying the one-step blending. These results indicate that genomic selection can greatly improve the accuracy of preselection for young bulls in Nordic Red population, and the one-step blending approach is a good alternative to predict GEBV in practical genetic evaluation program., (Copyright © 2012 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

Background: Size of the reference population and reliability of phenotypes are crucial factors influencing the reliability of genomic predictions. It is therefore useful to combine closely related populations. Increased accuracies of genomic predictions depend on the number of individuals added to the reference population, the reliability of their phenotypes, and the relatedness of the populations that are combined., Methods: This paper assesses the increase in reliability achieved when combining four Holstein reference populations of 4000 bulls each, from European breeding organizations, i.e. UNCEIA (France), VikingGenetics (Denmark, Sweden, Finland), DHV-VIT (Germany) and CRV (The Netherlands, Flanders). Each partner validated its own bulls using their national reference data and the combined data, respectively., Results: Combining the data significantly increased the reliability of genomic predictions for bulls in all four populations. Reliabilities increased by 10%, compared to reliabilities obtained with national reference populations alone, when they were averaged over countries and the traits evaluated. For different traits and countries, the increase in reliability ranged from 2% to 19%., Conclusions: Genomic selection programs benefit greatly from combining data from several closely related populations into a single large reference population.

Background: Genomic selection can be implemented by a multi-step procedure, which requires a response variable and a statistical method. For pure-bred pigs, it was hypothesised that deregressed estimated breeding values (EBV) with the parent average removed as the response variable generate higher reliabilities of genomic breeding values than EBV, and that the normal, thick-tailed and mixture-distribution models yield similar reliabilities., Methods: Reliabilities of genomic breeding values were estimated with EBV and deregressed EBV as response variables and under the three statistical methods, genomic BLUP, Bayesian Lasso and MIXTURE. The methods were examined by splitting data into a reference data set of 1375 genotyped animals that were performance tested before October 2008, and 536 genotyped validation animals that were performance tested after October 2008. The traits examined were daily gain and feed conversion ratio., Results: Using deregressed EBV as the response variable yielded 18 to 39% higher reliabilities of the genomic breeding values than using EBV as the response variable. For daily gain, the increase in reliability due to deregression was significant and approximately 35%, whereas for feed conversion ratio it ranged between 18 and 39% and was significant only when MIXTURE was used. Genomic BLUP, Bayesian Lasso and MIXTURE had similar reliabilities., Conclusions: Deregressed EBV is the preferred response variable, whereas the choice of statistical method is less critical for pure-bred pigs. The increase of 18 to 39% in reliability is worthwhile, since the reliabilities of the genomic breeding values directly affect the returns from genomic selection.

This study investigated the possibility of increasing the reliability of direct genomic values (DGV) by combining reference populations. The data were from 3,735 bulls from Danish, Swedish, and Finnish Red dairy cattle populations. Single nucleotide polymorphism markers were fitted as random variables in a Bayesian model, using published estimated breeding values as response variables. In total, 17 index traits were analyzed. Reliabilities were estimated using a 5-fold cross validation, and calculated as the within-year squared correlation between estimated breeding values and DGV. Marker effects were estimated using reference populations from individual countries, as well as using a combined reference population from all 3 countries. Single-country reference populations gave mean reliabilities across 17 traits of 0.19 to 0.23, whereas the combined reference gave mean reliabilities of 0.26 for all populations. Using marker effects from 1 population to predict the other 2 gave a loss in mean reliability of 0.14 to 0.21 when predicting Swedish or Finnish animals with Danish marker effects, or vice versa. Using Swedish or Finnish marker effects to predict each other only showed a loss in mean reliability of 0.03 to 0.05. A combined Swedish-Finnish reference population led to an average reliability as high as that from the 3-country reference population, but somewhat different for individual traits. The results from this study show that it is possible to increase the reliability of DGV by combining reference populations from related populations., (Copyright © 2011 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

The purpose of this study was to investigate the imputation error and loss of reliability of direct genomic values (DGV) or genomically enhanced breeding values (GEBV) when using genotypes imputed from a 3,000-marker single nucleotide polymorphism (SNP) panel to a 50,000-marker SNP panel. Data consisted of genotypes of 15,966 European Holstein bulls from the combined EuroGenomics reference population. Genotypes with the low-density chip were created by erasing markers from 50,000-marker data. The studies were performed in the Nordic countries (Denmark, Finland, and Sweden) using a BLUP model for prediction of DGV and in France using a genomic marker-assisted selection approach for prediction of GEBV. Imputation in both studies was done using a combination of the DAGPHASE 1.1 and Beagle 2.1.3 software. Traits considered were protein yield, fertility, somatic cell count, and udder depth. Imputation of missing markers and prediction of breeding values were performed using 2 different reference populations in each country: either a national reference population or a combined EuroGenomics reference population. Validation for accuracy of imputation and genomic prediction was done based on national test data. Mean imputation error rates when using national reference animals was 5.5 and 3.9% in the Nordic countries and France, respectively, whereas imputation based on the EuroGenomics reference data set gave mean error rates of 4.0 and 2.1%, respectively. Prediction of GEBV based on genotypes imputed with a national reference data set gave an absolute loss of 0.05 in mean reliability of GEBV in the French study, whereas a loss of 0.03 was obtained for reliability of DGV in the Nordic study. When genotypes were imputed using the EuroGenomics reference, a loss of 0.02 in mean reliability of GEBV was detected in the French study, and a loss of 0.06 was observed for the mean reliability of DGV in the Nordic study. Consequently, the reliability of DGV using the imputed SNP data was 0.38 based on national reference data, and 0.48 based on EuroGenomics reference data in the Nordic validation, and the reliability of GEBV using the imputed SNP data was 0.41 based on national reference data, and 0.44 based on EuroGenomics reference data in the French validation., (Copyright © 2011 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

This study compared genomic predictions using conventional estimated breeding values (EBV) and daughter yield deviations (DYD) as response variables based on simulated data. Eight scenarios were simulated in regard to heritability (0.05 and 0.30), number of daughters per sire (30, 100, and unequal numbers with an average of 100 per sire) and numbers of genotyped sires (all or half of sires were genotyped). The simulated genome had a length of 1200 cM with 15,000 equally spaced Single-nucleotide polymorphism (SNP) markers and 500 randomly distributed Quantitative trait locus (QTL). In the simulated scenarios, the EBV approach was as effective as or slightly better than the DYD approach at predicting breeding value, dependent on simulated scenarios and statistical models. Applying a Bayesian common prior model (the same prior distribution of marker effect variance) and a linear mixed model (GBLUP), the EBV and DYD approaches provided similar genomic estimated breeding value (GEBV) reliabilities, except for scenarios with unequal numbers of daughters and half of sires without genotype, for which the EBV approach was superior to the DYD approach (by 1.2 and 2.4%). Using a Bayesian mixture prior model (mixture prior distribution of marker effect variance), the EBV approach resulted in slightly higher reliabilities of GEBV than the DYD approach (by 0.3-3.6% with an average of 1.9%), and more obvious in scenarios with low heritability, small or unequal numbers of daughters, and half of sires without genotype. Moreover, the results showed that the correlation between GEBV and conventional parent average (PA) was lower (corresponding to a relatively larger gain by including PA) when using the DYD approach than when using the EBV approach. Consequently, the two approaches led to similar reliability of an index combining GEBV and PA in most scenarios. These results indicate that EBV can be used as an alternative response variable for genomic prediction., (© 2010 Blackwell Verlag GmbH.)

This study investigated the reliability of genomic estimated breeding values (GEBV) in the Danish Holstein population. The data in the analysis included 3,330 bulls with both published conventional EBV and single nucleotide polymorphism (SNP) markers. After data editing, 38,134 SNP markers were available. In the analysis, all SNP were fitted simultaneously as random effects in a Bayesian variable selection model, which allows heterogeneous variances for different SNP markers. The response variables were the official EBV. Direct GEBV were calculated as the sum of individual SNP effects. Initial analyses of 4 index traits were carried out to compare models with different intensities of shrinkage for SNP effects; that is, mixture prior distributions of scaling factors (standard deviation of SNP effects) assuming 5, 10, 20, or 50% of SNP having large effects and the others having very small or no effects, and a single prior distribution common for all SNP. It was found that, in general, the model with a common prior distribution of scaling factors had better predictive ability than any mixture prior models. Therefore, a common prior model was used to estimate SNP effects and breeding values for all 18 index traits. Reliability of GEBV was assessed by squared correlation between GEBV and conventional EBV (r(2)(GEBV, EBV)), and expected reliability was obtained from prediction error variance using a 5-fold cross validation. Squared correlations between GEBV and published EBV (without any adjustment) ranged from 0.252 to 0.700, with an average of 0.418. Expected reliabilities ranged from 0.494 to 0.733, with an average of 0.546. Averaged over 18 traits, r(2)(GEBV, EBV) was 0.13 higher and expected reliability was 0.26 higher than reliability of conventional parent average. The results indicate that genomic selection can greatly improve the accuracy of preselection for young bulls compared with traditional selection based on parent average information.

Crossbreeding is currently increasing in dairy cattle production. Several studies have shown an environment-dependent heterosis [i.e., an interaction between heterosis and environment (H x E)]. An H x E interaction is usually estimated from a few discrete environment levels. The present study proposes a reaction norm model to describe H x E interaction, which can deal with a large number of environment levels using few parameters. In the proposed model, total heterosis consists of an environment-independent part, which is described as a function of heterozygosity, and an environment-dependent part, which is described as a function of heterozygosity and environmental value (e.g., herd-year effect). A Bayesian approach is developed to estimate the environmental covariates, the regression coefficients of the reaction norm, and other parameters of the model simultaneously in both linear and nonlinear reaction norms. In the nonlinear reaction norm model, the H x E is approximated using linear splines. The approach was tested using simulated data, which were generated using an animal model with a reaction norm for heterosis. The simulation study includes 4 scenarios (the combinations of moderate vs. low heritability and moderate vs. low herd-year variation) of H x E interaction in a nonlinear form. In all scenarios, the proposed model predicted total heterosis very well. The correlation between true heterosis and predicted heterosis was 0.98 in the scenarios with low herd-year variation and 0.99 in the scenarios with moderate herd-year variation. This suggests that the proposed model and method could be a good approach to analyze H x E interactions and predict breeding values in situations in which heterosis changes gradually and continuously over an environmental gradient. On the other hand, it was found that a model ignoring H x E interaction did not significantly harm the prediction of breeding value under the simulated scenarios in which the variance for environment-dependent heterosis effects was small (as it generally is), and sires were randomly used over production environments.