Genomic selection strategies in dairy cattle: Strong positive interaction between use of genotypic information and intensive use of young bulls on genetic gain

We tested the following hypotheses: (i) breeding schemes with genomic selection are superior to breeding schemes without genomic selection regarding annual genetic gain of the aggregate genotype (ΔG(AG) ), annual genetic gain of the functional traits and rate of inbreeding per generation (ΔF), (ii) a positive interaction exists between the use of genotypic information and a short generation interval on ΔG(AG) and (iii) the inclusion of an indicator trait in the selection index will only result in a negligible increase in ΔG(AG) if genotypic information about the breeding goal trait is known. We examined four breeding schemes with or without genomic selection and with or without intensive use of young bulls using pseudo-genomic stochastic simulations. The breeding goal consisted of a milk production trait and a functional trait. The two breeding schemes with genomic selection resulted in higher ΔG(AG) , greater contributions of the functional trait to ΔG(AG) and lower ΔF than the two breeding schemes without genomic selection. Thus, the use of genotypic information may lead to more sustainable breeding schemes. In addition, a short generation interval increases the effect of using genotypic information on ΔG(AG) . Hence, a breeding scheme with genomic selection and with intensive use of young bulls (a turbo scheme) seems to offer the greatest potential. The third hypothesis was disproved as inclusion of genomically enhanced breeding values (GEBV) for an indicator trait in the selection index increased ΔG(AG) in the turbo scheme. Moreover, it increased the contribution of the functional trait to ΔG(AG) , and it decreased ΔF. Thus, indicator traits may still be profitable to use even when GEBV for the breeding goal traits are available.     

Assessing the accuracy of prediction for milk fatty acids by using a small reference population of tropical Holstein cows

Fatty acids (FA) have been related to effects on human health, sensory quality and shelf life of dairy products, cow's health and methane emission. However, despite their importance, they are not regularly measured in all dairy herds yet, which can affect the accuracy of estimated breeding values (EBV) for these traits. In this case, an alternative is to use genomic selection. Thus, the aim was to assess the use of genomic information in the genetic evaluation for milk traits in a tropical Holstein population. Monthly records (n = 36,457) of milk FA percentage, daily milk yield and quality traits from 4,203 cows as well as the genotypes of 755 of these cows for 57,368 single nucleotide polymorphisms (SNP) were used. Polygenic and genomic–polygenic models were applied for EBV prediction, and both models were compared through the EBV accuracy calculated from the prediction error and Spearman's correlation among EBV rankings. Prediction accuracy was assessed by using cross‐validation. In this case, the accuracy was the correlation between the genomic breeding values (GEBV) obtained as the sum of SNP effects and the EBV obtained in the polygenic model in each validation group. For all traits, the use of the genomic–polygenic model did not alter the animals' ranking, with correlations higher than 0.87. Nevertheless, through this model, the accuracy increased from 1.5% to 6.8% compared to the polygenic model. The correlations between GEBV and EBV varied from 0.52 to 0.68. Therefore, the use of a small group of genotyped cows in the genetic evaluation can increase the accuracy of EBV for milk FA and other traditional milk traits.     

Impact of fitting dominance and additive effects on accuracy of genomic prediction of breeding values in layers

Most genomic prediction studies fit only additive effects in models to estimate genomic breeding values (GEBV). However, if dominance genetic effects are an important source of variation for complex traits, accounting for them may improve the accuracy of GEBV. We investigated the effect of fitting dominance and additive effects on the accuracy of GEBV for eight egg production and quality traits in a purebred line of brown layers using pedigree or genomic information (42K single‐nucleotide polymorphism (SNP) panel). Phenotypes were corrected for the effect of hatch date. Additive and dominance genetic variances were estimated using genomic‐based [genomic best linear unbiased prediction (GBLUP)‐REML and BayesC] and pedigree‐based (PBLUP‐REML) methods. Breeding values were predicted using a model that included both additive and dominance effects and a model that included only additive effects. The reference population consisted of approximately 1800 animals hatched between 2004 and 2009, while approximately 300 young animals hatched in 2010 were used for validation. Accuracy of prediction was computed as the correlation between phenotypes and estimated breeding values of the validation animals divided by the square root of the estimate of heritability in the whole population. The proportion of dominance variance to total phenotypic variance ranged from 0.03 to 0.22 with PBLUP‐REML across traits, from 0 to 0.03 with GBLUP‐REML and from 0.01 to 0.05 with BayesC. Accuracies of GEBV ranged from 0.28 to 0.60 across traits. Inclusion of dominance effects did not improve the accuracy of GEBV, and differences in their accuracies between genomic‐based methods were small (0.01–0.05), with GBLUP‐REML yielding higher prediction accuracies than BayesC for egg production, egg colour and yolk weight, while BayesC yielded higher accuracies than GBLUP‐REML for the other traits. In conclusion, fitting dominance effects did not impact accuracy of genomic prediction of breeding values in this population.     

Theoretical efficiency of multiple-trait quantitative trait loci-assisted selection

The effectiveness of five selection methods for genetic improvement of net merit comprising trait 1 of low heritability (h² = 0.1) and trait 2 of high heritability (h² = 0.4) was examined: (i) two‐trait quantitative trait loci (QTL)‐assisted selection; (ii) partial QTL‐assisted selection based on trait 1; (iii) partial QTL‐assisted selection based on trait 2; (iv) QTL‐only selection; and (v) conventional selection index without QTL information. These selection methods were compared under 72 scenarios with different combinations of the relative economic weights, the genetic correlations between traits, the ratio of QTL variance to total genetic variance of the trait, and the ratio of genetic variances between traits. The results suggest that the detection of QTL for multiple‐trait QTL‐assisted selection is more important when the index traits are negatively correlated than when they are positively correlated. In contrast to literature reports that single‐trait marker‐assisted selection (MAS) is the most efficient for low heritability traits, this study found that the identified QTL of the low heritability trait contributed negligibly to total response in net merit. This is because multiple‐trait QTL‐assisted selection is designed to maximize total net merit rather than the genetic response of the individual index trait as in the case of single‐trait MAS. Therefore, it is not economical to identify the QTL of the low heritability traits for the improvement of total net merit. The efficient, cost‐effective selection strategy is to identify the QTL of the moderate or high heritability traits of the QTL‐assisted selection index to facilitate total economic returns. Detection of the QTL of the low h² traits for the QTL‐assisted index selection is justified when the low h² traits have high negative genetic correlation with the other index traits and/or when both economic weights and genetic variances of the low h² traits are larger as compared to the other index traits of higher h². This study deals with theoretical efficiency of QTL‐assisted selection, but the same principle applies to SNP‐based genomic selection when the proportion of the genetic variance ‘explained by the identified QTLs’ in this study is replaced by ‘explained by SNPs’.     

Comparison of heritabilities of dairy traits in Australian Holstein‐Friesian cattle from genomic and pedigree data and implications for genomic evaluations

The reliability of genomic evaluations depends on the proportion of genetic variation explained by the DNA markers. In this study, we have estimated the proportion of variance in daughter trait deviations (DTDs) of dairy bulls explained by 45 993 genome wide single‐nucleotide poly‐ morphism (SNP) markers for 29 traits in Australian Holstein‐Friesian dairy cattle. We compare these proportions to the proportion of variance in DTDs explained by the additive relationship matrix derived from the pedigree, as well as the sum of variance explained by both pedigree and marker information when these were fitted simultaneously. The propor‐ tion of genetic variance in DTDs relative to the total genetic variance (the total genetic variance explained by the genomic relationships and pedigree relationships when both were fitted simultaneously) varied from 32% for fertility to approximately 80% for milk yield traits. When fitting genomic and pedigree relationships simultaneously, the variance unexplained (i.e. the residual variance) in DTDs of the total variance for most traits was reduced compared to fitting either individually, suggesting that there is not complete overlap between the effects. The proportion of genetic variance accounted by the genomic relationships can be used to modify the blending equations used to calculate genomic estimated breeding value (GEBV) from direct genomic breeding value (DGV) and parent average. Our results, from a validation population of young dairy bulls with DTD, suggest that this modification can improve the reliability of GEBV by up to 5%.     

Comparison of genomic and traditional BLUP-estimated breeding value accuracy and selection response under alternative trait and genomic parameters

Accuracy of prediction of estimated breeding values based on genome-wide markers (GEBV) and selection based on GEBV as compared with traditional Best Linear Unbiased Prediction (BLUP) was examined for a number of alternatives, including low heritability, number of generations of training, marker density, initial distributions, and effective population size (Ne). Results show that the more the generations of data in which both genotypes and phenotypes were collected, termed training generations (TG), the better the accuracy and persistency of accuracy based on GEBV. GEBV excelled for traits of low heritability regardless of initial equilibrium conditions, as opposed to traditional marker-assisted selection, which is not useful for traits of low heritability. Effective population size is critical for populations starting in Hardy-Weinberg equilibrium but not for populations started from mutation-drift equilibrium. In comparison with traditional BLUP, GEBV can exceed the accuracy of BLUP provided enough TG are included. Unfortunately selection rapidly reduces the accuracy of GEBV. In all cases examined, classic BLUP selection exceeds what was possible for GEBV selection. Even still, GEBV could have an advantage over traditional BLUP in cases such as sex-limited traits, traits that are expensive to measure, or can only be measured on relatives. A combined approach, utilizing a mixed model with a second random effect to account for quantitative trait loci in linkage equilibrium (the polygenic effect) was suggested as a way to capitalize on both methodologies.     

Comparison between genomic predictions using daughter yield deviation and conventional estimated breeding value as response variables

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.     

Estimates of genetic variance and variance of predicted genetic merits using pedigree or genomic relationship matrices in six Brown Swiss cattle populations for different traits

The amount of variance captured in genetic estimations may depend on whether a pedigree‐based or genomic relationship matrix is used. The purpose of this study was to investigate the genetic variance as well as the variance of predicted genetic merits (PGM) using pedigree‐based or genomic relationship matrices in Brown Swiss cattle. We examined a range of traits in six populations amounting to 173 population‐trait combinations. A main aim was to determine how using different relationship matrices affect variance estimation. We calculated ratios between different types of estimates and analysed the impact of trait heritability and population size. The genetic variances estimated by REML using a genomic relationship matrix were always smaller than the variances that were similarly estimated using a pedigree‐based relationship matrix. The variances from the genomic relationship matrix became closer to estimates from a pedigree relationship matrix as heritability and population size increased. In contrast, variances of predicted genetic merits obtained using a genomic relationship matrix were mostly larger than variances of genetic merit predicted using pedigree‐based relationship matrix. The ratio of the genomic to pedigree‐based PGM variances decreased as heritability and population size rose. The increased variance among predicted genetic merits is important for animal breeding because this is one of the factors influencing genetic progress.     

Comparison between Reduced-Representation Genome Sequencing and SNP Chip for Genomic Selection in Yellow-feathered Broiler

This study aimed to compare the accuracy of the genomic estimated breeding value (GEBV) using reduced-representation genome sequencing technology and SNP chip technology to implement genomic selection. A total of 395 individuals (212♂+ 183♀, from 8 half-sib families) were randomly selected from F₂ generation of AH broiler resource population, and genotyped with 10×specific-locus amplified fragment sequencing (SLAF-seq) and Illumina Chicken 60K SNP BeadChip. Genomic best linear unbiased prediction (GBLUP) and BayesCπ were used to compare the accuracy of genomic estimated breeding values (GEBV) for 6 traits: body weight at the 6^th week, body weight at the 12^th week, average daily gain (ADG), average daily feed intake (ADFI), feed conversion ratio (FCR) and residual feed intake (RFI). A 5-fold cross validation procedure was used to verify the accuracies of GEBV between prediction models and between genotyping platforms. The results showed that there was no significant difference between accuracies of GEBV predicted by GBLUP and BayesCπ using the same genotyping platform(P>0.05). The superiority of the two genotyping platforms was different for different traits. For body weight at the 6^th week, the accuracy of GEBV was higher using chip SNPs (P<0.05). On the contrary, the accuracy was higher using SLAF-seq for residual feed intake (P<0.05). Comprehensive comparison of the means of GEBV for 6 traits, the difference between the two genotyping platforms was less than 0.01, therefore, both high throughput sequencing and chip SNPs can be used for genomic selection in yellow-feathered broiler.     

基于简化基因组测序技术和基因芯片技术比较研究黄羽肉鸡基因组选择

旨在比较简化基因组测序技术和基因芯片技术实施基因组选择的基因组估计育种值（GEBV）准确性。本研究在AH肉鸡资源群体F₂代中随机选取395个个体（其中公鸡212只,母鸡183只,来自8个半同胞家系）,同时采用10×SLAF测序技术和Illumina Chicken 60K SNP芯片进行基因标记分型。采用基因组最佳无偏估计法（GBLUP）和BayesCπ对6周体重、12周体重、日均增重、日均采食量、饲料转化率和剩余采食量等6个性状进行GEBV准确性比较研究,并采用5折交叉验证法验证。结果表明,采用同一基因标记分型平台,两种育种值估计方法所得GEBV准确性差异不显著（P>0.05）;不同的性状对基因标记分型平台的选择存在差异,对于6周体重,使用基因芯片可获得更高的GEBV准确性（P<0.05）,对于剩余采食量,则使用简化基因组测序可获得更高的GEBV准确性（P<0.05）。综合6个性状GEBV均值比较,两个基因标记分型平台之间差异不到0.01,高通量测序技术和基因芯片技术都可以用于黄羽肉鸡基因组选择。     

Development of genomic predictions for Angus cattle in Brazil incorporating genotypes from related American sires

Genomic prediction has become the new standard for genetic improvement programs, and currently, there is a desire to implement this technology for the evaluation of Angus cattle in Brazil. Thus, the main objective of this study was to assess the feasibility of evaluating young Brazilian Angus (BA) bulls and heifers for 12 routinely recorded traits using single-step genomic BLUP (ssGBLUP) with and without genotypes from American Angus (AA) sires. The second objective was to obtain estimates of effective population size (N_e) and linkage disequilibrium (LD) in the Brazilian Angus population. The dataset contained phenotypic information for up to 277,661 animals belonging to the Promebo breeding program, pedigree for 362,900, of which 1,386 were genotyped for 50k, 77k, and 150k single nucleotide polymorphism (SNP) panels. After imputation and quality control, 61,666 SNPs were available for the analyses. In addition, genotypes from 332 American Angus (AA) sires widely used in Brazil were retrieved from the AA Association database to be used for genomic predictions. Bivariate animal models were used to estimate variance components, traditional EBV, and genomic EBV (GEBV). Validation was carried out with the linear regression method (LR) using young-genotyped animals born between 2013 and 2015 without phenotypes in the reduced dataset and with records in the complete dataset. Validation animals were further split into progeny of BA and AA sires to evaluate if their progenies would benefit by including genotypes from AA sires. The N_e was 254 based on pedigree and 197 based on LD, and the average LD (±SD) and distance between adjacent single nucleotide polymorphisms (SNPs) across all chromosomes were 0.27 (±0.27) and 40743.68 bp, respectively. Prediction accuracies with ssGBLUP outperformed BLUP for all traits, improving accuracies by, on average, 16% for BA young bulls and heifers. The GEBV prediction accuracies ranged from 0.37 (total maternal for weaning weight and tick count) to 0.54 (yearling precocity) across all traits, and dispersion (LR coefficients) fluctuated between 0.92 and 1.06. Inclusion of genotyped sires from the AA improved GEBV accuracies by 2%, on average, compared to using only the BA reference population. Our study indicated that genomic information could help us to improve GEBV accuracies and hence genetic progress in the Brazilian Angus population. The inclusion of genotypes from American Angus sires heavily used in Brazil just marginally increased the GEBV accuracies for selection candidates.     

Use of genomic recursions and algorithm for proven and young animals for single‐step genomic BLUP analyses – a simulation study

The purpose of this study was to examine accuracy of genomic selection via single‐step genomic BLUP (ssGBLUP) when the direct inverse of the genomic relationship matrix ( G ) is replaced by an approximation of G ^?1 based on recursions for young genotyped animals conditioned on a subset of proven animals, termed algorithm for proven and young animals (APY). With the efficient implementation, this algorithm has a cubic cost with proven animals and linear with young animals. Ten duplicate data sets mimicking a dairy cattle population were simulated. In a first scenario, genomic information for 20k genotyped bulls, divided in 7k proven and 13k young bulls, was generated for each replicate. In a second scenario, 5k genotyped cows with phenotypes were included in the analysis as young animals. Accuracies (average for the 10 replicates) in regular EBV were 0.72 and 0.34 for proven and young animals, respectively. When genomic information was included, they increased to 0.75 and 0.50. No differences between genomic EBV (GEBV) obtained with the regular G ^?1 and the approximated G ^?1 via the recursive method were observed. In the second scenario, accuracies in GEBV (0.76, 0.51 and 0.59 for proven bulls, young males and young females, respectively) were also higher than those in EBV (0.72, 0.35 and 0.49). Again, no differences between GEBV with regular G ^?1 and with recursions were observed. With the recursive algorithm, the number of iterations to achieve convergence was reduced from 227 to 206 in the first scenario and from 232 to 209 in the second scenario. Cows can be treated as young animals in APY without reducing the accuracy. The proposed algorithm can be implemented to reduce computing costs and to overcome current limitations on the number of genotyped animals in the ssGBLUP method.     

Accuracy of genomic breeding values for residual feed intake in crossbred beef cattle

The benefit of using genomic breeding values (GEBV) in predicting ADG, DMI, and residual feed intake for an admixed population was investigated. Phenotypic data consisting of individual daily feed intake measurements for 721 beef cattle steers tested over 5 yr was available for analysis. The animals used were an admixed population of spring-born steers, progeny of a cross between 3 sire breeds and a composite dam line. Training and validation data sets were defined by randomly splitting the data into training and testing data sets based on sire family so that there was no overlap of sires in the 2 sets. The random split was replicated to obtain 5 separate data sets. Two methods (BayesB and random regression BLUP) were used to estimate marker effects and to define marker panels and ultimately the GEBV. The accuracy of prediction (the correlation between the phenotypes and GEBV) was compared between SNP panels. Accuracy for all traits was low, ranging from 0.223 to 0.479 for marker panels with 200 SNP, and 0.114 to 0.246 for marker panels with 37,959 SNP, depending on the genomic selection method used. This was less than accuracies observed for polygenic EBV accuracies, which ranged from 0.504 to 0.602. The results obtained from this study demonstrate that the utility of genetic markers for genomic prediction of residual feed intake in beef cattle may be suboptimal. Differences in accuracy were observed between sire breeds when the random regression BLUP method was used, which may imply that the correlations obtained by this method were confounded by the ability of the selected SNP to trace breed differences. This may also suggest that prediction equations derived from such an admixed population may be useful only in populations of similar composition. Given the sample size used in this study, there is a need for increased feed intake testing if substantially greater accuracies are to be achieved.     

Adjusting for heterogeneity of variance for carcass traits affects single and multiple trait selections in genetic evaluation of Japanese Black cattle

Heterogeneity of variance among subclasses of an effect is a potential source of bias in genetic evaluation. Degrees of the heterogeneity of variance among farm‐market‐year‐sex (FMYS) subclasses for carcass weight, beef marbling standard number, rib‐eye area, rib thickness and subcutaneous fat thickness were investigated in Japanese Black cattle. Consequences of adjusting for the heterogeneity on the predicted breeding values (PBVs) or on the genetic indexes derived from the PBVs of the five carcass traits were assessed. A total of 57 461 records were collected between 1997 and 2002 from steers and heifers fattened at farms across Japan. These records were grouped into 1591 FMYS subclasses. Bartlett's test showed that the degree of the heterogeneity of variance among the FMYS subclasses was sizeable in all traits (P < 0.0001). By applying a two‐step adjustment procedure it was possible to reduce the standard deviation, the coefficient of variation and the Gini coefficient of the phenotypic variances by 67.5% to 75.0% in the different traits. The applied adjustment caused a substantial re‐ranking of elite dams in the PBV for each trait as well as in the genetic index. This study provided evidence that the applied adjustment reduces the bias in the PBVs due to heterogeneous variances and increases the accuracy of bull‐dam selection.     

Effect of selection and selective genotyping for creation of reference on bias and accuracy of genomic prediction

Reference populations for genomic selection usually involve selected individuals, which may result in biased prediction of estimated genomic breeding values (GEBV). In a simulation study, bias and accuracy of GEBV were explored for various genetic models with individuals selectively genotyped in a typical nucleus breeding program. We compared the performance of three existing methods, that is, Best Linear Unbiased Prediction of breeding values using pedigree‐based relationships (PBLUP), genomic relationships for genotyped animals only (GBLUP) and a Single‐Step approach (SSGBLUP) using both. For a scenario with no‐selection and random mating (RR), prediction was unbiased. However, lower accuracy and bias were observed for scenarios with selection and random mating (SR) or selection and positive assortative mating (SA). As expected, bias disappeared when all individuals were genotyped and used in GBLUP. SSGBLUP showed higher accuracy compared to GBLUP, and bias of prediction was negligible with SR. However, PBLUP and SSGBLUP still showed bias in SA due to high inbreeding. SSGBLUP and PBLUP were unbiased provided that inbreeding was accounted for in the relationship matrices. Selective genotyping based on extreme phenotypic contrasts increased the prediction accuracy, but prediction was biased when using GBLUP. SSGBLUP could correct the biasedness while gaining higher accuracy than GBLUP. In a typical animal breeding program, where it is too expensive to genotype all animals, it would be appropriate to genotype phenotypically contrasting selection candidates and use a Single‐Step approach to obtain accurate and unbiased prediction of GEBV.     

基于基因组和系谱信息的不同选配方案效果模拟研究

基因组选配（genomic mating,GM）是利用基因组信息进行优化的选种选配,可以有效控制群体近交水平的同时实现最大化的遗传进展。但基因组选配是对群体中所有个体进行选配,这与实际的育种工作有点相悖。本研究模拟了遗传力为0.5的9 000头个体的基础群数据,每个世代根据GEBV选择30头公畜、900头母畜作为种用个体,而后使用基因组选配、同质选配、异质选配、随机交配4种不同的选配方案。其中基因组选配中分别选取遗传进展最大的解、家系间方差最大的解、近交最小的解所对应的交配方案进行选育。每种方案选育5个世代,比较其后代群体的平均GEBV、每世代的遗传进展、近交系数、遗传方差,并重复5次取平均值。结果表明,3种基因组选配方案的ΔG均显著高于随机交配和异质选配（P<0.01）,而且,选取遗传进展最大的基因组选配方案的ΔG比同质选配还高出4.3%。3种基因组选配的方案的ΔF比同质选配低22.2%~94.1%,而且选取近交最小的基因组选配方案ΔF比异质选配低11.8%。同质选配的遗传方差迅速降低,在第5世代显著低于除基因组选配中选择遗传进展最大的方案以外的所有方案（P<0.05）,3种基因组选配方案的遗传方差比同质选配高10.8%~32.2%。这表明基因组选配不仅可以获得比同质选配更高的遗传进展,同时有效的降低了近交水平,并且减缓了遗传方差降低速度,保证了一定的遗传变异。基因组选配作为一种有效的可持续育种方法,在畜禽育种中开展十分有必要。     

The unified approach to the use of genomic and pedigree information in genomic evaluations revisited

Previous proposals for a unified approach for amalgamating information from animals with or without genotypes have combined the numerator relationship matrix A with the genomic relationship G estimated from the markers. These approaches have resulted in biased genomic EBV (GEBV), and methodology was developed to overcome these problems. Firstly, a relationship matrix, G(FG) , based on linkage analysis was derived using the same base population as A, which (i) utilizes the genomic information on the same scale as the pedigree information and (ii) permits the regression coefficients used to propagate the genomic data from the genotyped to ungenotyped individuals to be calculated in the light of the genomic information, rather than ignoring it. Secondly, the elements of G were regressed back towards their expected values in the A matrix to allow for their estimation errors. These developments were combined in a methodology LDLAb and tested on simulated populations where either parents were phenotyped and offspring genotyped or vice versa. The LDLAb method was demonstrated to be a unified approach that maximized accuracy of GEBV compared to previous methodologies and removed the bias in the GEBV. Although LDLAb is computationally much more demanding than MLAC, it demonstrates how to make best use the marker information and also shows the computational problems that need to be solved in the future to make best use of the marker data.     

我国荷斯坦青年公牛基因组选择效果分析

本研究基于我国荷斯坦奶牛基因组遗传评估和生产性能测定（DHI）结果,旨在分析我国荷斯坦公牛基因组选择的效果。选择1 686头既有基因组遗传评估成绩又有后裔测定成绩的荷斯坦公牛,利用2019年12月基因组遗传评估结果及其女儿的产奶和体型性状数据,通过R软件与Excel计算公牛基因组评估结果与公牛女儿表型数据间的相关性,对我国荷斯坦青年公牛基因组选择效果进行分析。相关性分析结果表明,荷斯坦公牛的基因组性能指数（GCPI）与后裔测定性能指数（CPI）呈正相关（r_s>0.3）,其中产奶量和体细胞评分的基因组育种值（GEBV）与估计育种值（EBV）呈较强的正相关（0.4 < r_s < 0.8）。对公牛女儿表型数据分析结果表明,女儿产奶量、乳蛋白率、乳脂率与肢蹄评分的表型值与公牛GEBV分组趋势一致,且公牛不同产奶性状GEBV组间的女儿性状表型值大部分达到极显著差异（P<0.01）;北京及上海地区公牛产奶性状、体细胞评分和肢蹄评分的GEBV分组与女儿表型值趋势较其他省市（地区）更一致,且GEBV高组与低组之间差值均高于其他省市（地区）。基于1 686头荷斯坦公牛基因组选择及其女儿表型数据的分析结果表明,我国荷斯坦公牛的基因组遗传评估准确性较好,其中产奶量、乳蛋白率、体细胞评分和肢蹄评分的表型数据更好地反映了基因组选择的效果;北京及上海地区较其他省市（地区）更能反映我国荷斯坦公牛基因组选择的效果。     

全基因组选择及其在奶牛育种中应用进展

全基因组选择是指基于基因组育种值（GEBV）的选择方法,指通过检测覆盖全基因组的分子标记,利用基因组水平的遗传信息对个体进行遗传评估,以期获得更高的育种值估计准确度。由于可显著缩短世代间隔,全基因组选择作为一项育种新技术在奶牛育种中具有广阔的应用前景,目前已经成为各国的研究热点。不同国家的试验结果表明,奶牛育种中基于GEBV的遗传评估可靠性在20%~67%之间,如果代替常规后裔测定体系,可节省92%的育种成本。本文综述了全基因组选择的基本原理及其在各国奶牛育种中的应用现状和所面临的问题。     

Genomic selection in American mink (Neovison vison) using a single-step genomic best linear unbiased prediction model for size and quality traits graded on live mink

Genomic selection relies on single-nucleotide polymorphisms (SNPs), which are often collected using medium-density SNP arrays. In mink, no such array is available; instead, genotyping by sequencing (GBS) can be used to generate marker information. Here, we evaluated the effect of genomic selection for mink using GBS. We compared the estimated breeding values (EBVs) from single-step genomic best linear unbiased prediction (SSGBLUP) models to the EBV from ordinary pedigree-based BLUP models. We analyzed seven size and quality traits from the live grading of brown mink. The phenotype data consisted of ~20,600 records for the seven traits from the mink born between 2013 and 2016. Genotype data included 2,103 mink born between 2010 and 2014, mostly breeding animals. In total, 28,336 SNP markers from 391 scaffolds were available for genomic prediction. The pedigree file included 29,212 mink. The predictive ability was assessed by the correlation (r) between progeny trait deviation (PTD) and EBV, and the regression of PTD on EBV, using 5-fold cross-validation. For each fold, one-fifth of animals born in 2014 formed the validation set. For all traits, the SSGBLUP model resulted in higher accuracies than the BLUP model. The average increase in accuracy was 15% (between 3% for fur clarity and 28% for body weight). For three traits (body weight, silky appearance of the under wool, and guard hair thickness), the difference in r between the two models was significant (P < 0.05). For all traits, the regression slopes of PTD on EBV from SSGBLUP models were closer to 1 than regression slopes from BLUP models, indicating SSGBLUP models resulted in less bias of EBV for selection candidates than the BLUP models. However, the regression coefficients did not differ significantly. In conclusion, the SSGBLUP model is superior to conventional BLUP model in the accurate selection of superior animals, and, thus, it would increase genetic gain in a selective breeding program. In addition, this study shows that GBS data work well in genomic prediction in mink, demonstrating the potential of GBS for genomic selection in livestock species.