Non-genotyped Animals Research Articles

Comparative Analysis of Genomic and Pedigree-Based Approaches for Genetic Evaluation of Morphological Traits in Pura Raza Española Horses

Background: The single-step best linear unbiased predictor (ssGBLUP) has emerged as a reference method for genomic selection in recent years due to its advantages over traditional approaches. Although its application in horses remains limited, ssGBLUP has demonstrated the potential to improve the reliability of estimated breeding values in livestock species. This study aimed to assess the impact of incorporating genomic data using single-step restricted maximum likelihood (ssGREML) on reliability (R2) in the Pura Raza Española (PRE) horse breed, compared to traditional pedigree-based REML. Methods: The analysis involved 14 morphological traits from 7152 animals, including 2916 genotyped individuals. Genetic parameters were estimated using a multivariate model. Results: Results showed that heritability estimates were similar between the two approaches, ranging from 0.08 to 0.76. However, a significant increase in reliability (R2) was observed for ssGREML compared to REML across all morphological traits, with overall gains ranging from 1.56% to 13.30% depending on the trait evaluated. R2 ranged from 6.93% to 22.70% in genotyped animals, significantly lower in non-genotyped animals (0.82% to 12.37%). Interestingly, individuals with low R2 values in REML demonstrated the largest R2 gains in ssGREML. Additionally, this improvement was much greater (5.96% to 19.25%) when only considering stallions with less than 40 controlled foals. Conclusions: Hereby, we demonstrated that the application of genomic selection can contribute to improving the reliability of mating decisions in a large horse breeding program such as the PRE breed.

Inclusion of imputed genotypes from non-genotyped dairy cattle in a Thai multibreed genomic-polygenic evaluation.

This study assessed the impact of incorporating imputed SNP information from non-genotyped animals on genomic-polygenic evaluations in a Thai multibreed dairy population under various levels of imputation accuracy. Data encompassed pedigree and phenotypic records for 305-day milk yield (MY), 305-day fat (Fat), and age at first calving (AFC) from 12,859 first-lactation cows, and genotypic records of various densities from 4,364 animals. A set of 64 animals genotyped with GeneSeek Genomic Profiler 80K and with four or more genotyped progenies was defined as target animals to simulate imputation scenarios for non-genotyped individuals. Actual and imputed genotypes were utilized to construct three SNP sets. All SNP Sets contained actual and imputed SNP markers from genotyped animals. SNP Set 1 contained no SNPs from target animals, whereas SNP Set 2 incorporated imputed SNPs from target animals, and SNP Set 3 added actual SNPs from target animals. Genomic-polygenic evaluations were conducted using a 3-trait single-step model that included contemporary group, calving age, and heterozygosity as fixed effects and animal additive genetic and residual as random effects. The imputation accuracy was similar across non-genotyped animals irrespective of the number of genotyped progenies (average: 40.55%; range: 34.68% to 53.82%). Estimates of additive genetic and environmental variances and covariances for MY and AFC varied across SNP sets. SNP Sets 1 and 2 had slightly higher additive genetic and lower environmental variances and covariances than SNP Set 3. Heritabilities and additive genetic, environmental, and phenotypic correlations between MY, Fat, and AFC were similar across all SNP Sets. Spearman rank correlations between genomic-polygenic EBVs from SNP Sets 2 and 3 were high for all traits (0.9990±0.0003). Utilization of phenotypic and pedigree data from imputed non-genotyped animals enhanced the efficiency and cost-effectiveness of the genetic improvement program in the Thai multibreed dairy cattle population.

Short Communication: Reduced GBLUP equations to core animals in the algorithm for proven and young (APY)

The number of animal genotypes is rapidly increasing, and a major challenge for animal models is inverting the genomic relationship matrix (G). Matrix G has a limited dimensionality, and the algorithm for proven and young (APY) makes inverting a large G possible via the inverse of a block diagonal of G with a size equivalent to the dimensionality of G. APY divides genotyped animals into core and non-core groups, and breeding values of non-core animals are conditioned on the breeding values of core animals. Therefore, there is the possibility of opting out equations for non-core animals from the model. A methodology was presented for a reduced APY genomic BLUP (GBLUP) to equations for core animals. Using a small example dataset, the method was validated by the equality of the full and the reduced model analysis results. Absorption of fixed effect equations into random effect equations was successful in reducing the number of equations to solve and producing the same random effect solutions. Extending the method to APY single-step GBLUP (ssGBLUP) was not computationally justifiable. Other reduction techniques exist for ssGBLUP (regardless of APY or non-APY) that work by reducing the number of equations for non-genotyped animals. The number of equations can further be reduced by data pruning.

The present of Italian Mediterranean buffalo: precision breeding based on multi-omics data

Genetic evaluation in the Italian Mediterranean Buffalo (IMB) traditionally relied on the BLUP method (best linear unbiased predictor), a mixed model system incorporating both random and fixed effects simultaneously. However, recent advancements in genome sequencing technologies have opened up the opportunity to incorporate genomic information into genetic evaluations. The ssGBLUP (single-step best linear unbiased predictor) has become the method par excellence. It replaces the traditional relationship matrix with one that combines pedigree and genomic relationships, allowing for the estimation of genetic values for non-genotyped animals. The findings of this study highlight how genomic selection enhances the precision of breeding values, facilitates greater genetic advancement and reduces the generation interval, ultimately enabling a rapid return on investment.

Heterogeneity in convergence behaviour of the single-step SNP-BLUP model across different effects and animal groups

BackgroundThe single-step model is becoming increasingly popular for national genetic evaluations of dairy cattle due to the benefits that it offers such as joint breeding value estimation for genotyped and ungenotyped animals. However, the complexity of the model due to a large number of correlated effects can lead to significant computational challenges, especially in terms of accuracy and efficiency of the preconditioned conjugate gradient method used for the estimation. The aim of this study was to investigate the effect of pedigree depth on the model's overall convergence rate as well as on the convergence of different components of the model, in the context of the single-step single nucleotide polymorphism best linear unbiased prediction (SNP-BLUP) model.ResultsThe results demonstrate that the dataset with a truncated pedigree converged twice as fast as the full dataset. Still, both datasets showed very high Pearson correlations between predicted breeding values. In addition, by comparing the top 50 bulls between the two datasets we found a high correlation between their rankings. We also analysed the specific convergence patterns underlying different animal groups and model effects, which revealed heterogeneity in convergence behaviour. Effects of SNPs converged the fastest while those of genetic groups converged the slowest, which reflects the difference in information content available in the dataset for those effects. Pre-selection criteria for the SNP set based on minor allele frequency had no impact on either the rate or pattern of their convergence. Among different groups of individuals, genotyped animals with phenotype data converged the fastest, while non-genotyped animals without own records required the largest number of iterations.ConclusionsWe conclude that pedigree structure markedly impacts the convergence rate of the optimisation which is more efficient for the truncated than for the full dataset.

Implementing genomic selection in the IMB: challenges and opportunities

Single-step genomic best linear unbiased predictor (ssGBLUP) is a method for jointly estimating breeding values (BV) for genotyped and non-genotyped animals. Genomic information in the Italian Mediterranean Buffalo (IMB) is now available. Its inclusion in the genetic evaluation system could increase both the accuracy and genetic progress of the traits of interest of the breed. The study aimed to test the feasibility of ssGBLUP and show the first results of implementing a genomic evaluation for production and type traits in the IMB. Phenotypic information on production (270-day milk, mozzarella yield (MY), protein and fat kg and %, respectively) and morphology: feet and legs (FL) and mammary system (MS) were used for this study. Production records included 743,904 lactations from 276,451 buffalo cows born from 1984 to 2019. Morphological traits were from 91,966 buffalo cows from 2004 to 2022. Regarding the genotypes, 2,017 buffalo cows and 133 bulls were used. Data were analyzed fitting two multi-trait animal models, a 6-trait model for production data and a 2-trait model for morphology data. According to the relationship matrix used, two models were fitted: (i) the BLUP with the numerator relationship matrix (A) and (ii) the ssGBLUP where A and the genomic relationship matrix (G) are blended into H. BVs were estimated with BLUP and ssGBLUP models. The cutoff year used to create the partial data set was 2012. The correlation, accuracy, dispersion, and bias statistics were calculated (LR method). Both bulls (N=49) and cows (N=1288) were used for validations. On average, the correlation between EBVs from partial and whole datasets estimated with BLUP and ssGBLUP increased from 6 to 49% and from 14 to 17% for production and type traits, respectively. Among the traits analyzed, the most affected by the change were protein/fat content, MY, and AM. The accuracy increase for these traits was above 20% when using the ssGBLUP. All LR statistics also improved for non-genotyped females. These results showed that implementing ssGBLUP in the breeding program can generate more accurate predictions for essential traits in dairy IMB than traditional BLUP.

149 Marker Effect P-Value for Large Genotype Populations with the Algorithm for Proven and Young

Abstract The single-step method (ssGBLUP) allows for simultaneous evaluations of populations composed of genotyped and non-genotyped animals. As a GBLUP-based method, the output of ssGBLUP is breeding values. However, interest might rely on the estimation of SNP effects and on how genome segments are associated with the phenotype of interest. When that is the case, SNP effects can be obtained from a linear transformation of breeding values, and P-values can be used as a measure of estimation certainty. P-values are derived from the prediction error variance of SNP effects, which relies on the prediction error (co)variance matrix of breeding values obtained from the inverse of the left-hand side of the mixed model equations (LHS). However, inverting the LHS becomes unfeasible with large, genotyped populations. Therefore, alternative strategies to overcome this problem and obtain p-values with increasing genotyped populations might be evaluated. In this study, we aimed to estimate marker effect p-values when the algorithm for proven and young (APY) was used to approximate the predictor error (co)variance for animals in a large, genotyped population. We estimate P-values of SNP marker effects for PWG in an Angus cattle population. The dataset was composed of around 844K phenotyped animals, 450K genotypes, and 1.8M pedigree records. Analyses were split into two sets. In the first set, a reduced genotype data of 50K was used, so p-values obtained from a direct-inverse of LHS using the regular G-1 (Exact_G) or APY G-1 (Exact_GAPY) were compared with APY G-1 and an approximation of the prediction error variance that did not require the inversion of the LHS (Approx_GAPY). In the second set of analyses, P-values with Approx_GAPY were estimated with the full genomic set of 450K (Approx_GAPY_450K), and computational requirements were recorded. All analyses were performed in three replicates. In the first set of analyses, genome-wide association with Exact_G uncovered two significant regions in chromosomes 7 and 20. Along all replicates, the same regions were also identified with Exact_GAPY and Approx_GAPY, indicating that similar resolutions are obtained with exact and approximated p-values. Results from Approx_GAPY_450K showed that, with a complete genome set, besides the two regions in chromosomes 7 and 20, two new regions in chromosomes 6 and 14 were identified. On average, the entire procedure for obtaining P-values with 450K genotyped animals had an elapsed wall clock time of 24h with a maximum memory usage of 87.6 GB. Altogether, our results suggest that with APY and the approximation of the prediction error variance, current computational boundaries for obtaining marker effect P-values for large genotyped populations should be lifted.

Validation with single-step SNPBLUP shows that evaluations can continue using a single mean of genotyped individuals, even with multiple breeds

BackgroundIn genomic prediction, it is common to centre the genotypes of single nucleotide polymorphisms based on the allele frequencies in the current population, rather than those in the base generation. The mean breeding value of non-genotyped animals is conditional on the mean performance of genotyped relatives, but can be corrected by fitting the mean performance of genotyped individuals as a fixed regression. The associated covariate vector has been referred to as a ‘J-factor’, which if fitted as a fixed effect can improve the accuracy and dispersion bias of sire genomic estimated breeding values (GEBV). To date, this has only been performed on populations with a single breed. Here, we investigated whether there was any benefit in fitting a separate J-factor for each breed in a three-way crossbred population, and in using pedigree-based expected or genome-based estimated breed fractions to define the J-factors.ResultsFor body weight at 7 days, dispersion bias decreased when fitting multiple J-factors, but only with a low proportion of genotyped individuals with selective genotyping. On average, the mean regression coefficients of validation records on those of GEBV increased with one J-factor compared to none, and further increased with multiple J-factors. However, for body weight at 35 days this was not observed. The accuracy of GEBV remained unchanged regardless of the J-factor method used. Differences between the J-factor methods were limited with correlations approaching 1 for the estimated covariate vector, the estimated coefficients of the regression on the J-factors, and the GEBV.ConclusionsBased on our results and in the particular design analysed here, i.e. all the animals with phenotype are of the same type of crossbreds, fitting a single J-factor should be sufficient, to reduce dispersion bias. Fitting multiple J-factors may reduce dispersion bias further but this depends on the trait and genotyping rate. For the crossbred population analysed, fitting multiple J-factors has no adverse consequences and if this is done, it does not matter if the breed fractions used are based on the pedigree-expectation or the genomic estimates. Finally, when GEBV are estimated from crossbred data, any observed bias can potentially be reduced by including a straightforward regression on actual breed proportions.

Single-step genomic evaluation for growth traits in a Mexican Braunvieh cattle population.

The objective was to compare (pedigree-based) BLUP, genomic BLUP (GBLUP), and single-step GBLUP (ssGBLUP) methods for genomic evaluation of growth traits in a Mexican Braunvieh cattle population. Birth (BW), weaning (WW), and yearling weight (YW) data of a Mexican Braunvieh cattle population were analyzed with BLUP, GBLUP, and ssGBLUP methods. These methods are differentiated by the additive genetic relationship matrix included in the model and the animals under evaluation. The predictive ability of the model was evaluated using random partitions of the data in training and testing sets, consistently predicting about 20% of genotyped animals on all occasions. For each partition, the Pearson correlation coefficient between adjusted phenotypes for fixed effects and non-genetic random effects and the estimated breeding values (EBV) were computed. The random contemporary group (CG) effect explained about 50, 45, and 35% of the phenotypic variance in BW, WW, and YW, respectively. For the three methods, the CG effect explained the highest proportion of the phenotypic variances (except for YW-GBLUP). The heritability estimate obtained with GBLUP was the lowest for BW, while the highest heritability was obtained with BLUP. For WW, the highest heritability estimate was obtained with BLUP, the estimates obtained with GBLUP and ssGBLUP were similar. For YW, the heritability estimates obtained with GBLUP and BLUP were similar, and the lowest heritability was obtained with ssGBLUP. Pearson correlation coefficients between adjusted phenotypes for non-genetic effects and EBVs were the highest for BLUP, followed by ssBLUP and GBLUP. The successful implementation of genetic evaluations that include genotyped and non-genotyped animals in our study indicate a promising method for use in genetic improvement programs of Braunvieh cattle. Our findings showed that simultaneous evaluation of genotyped and non-genotyped animals improved prediction accuracy for growth traits even with a limited number of genotyped animals.

A computationally efficient method for approximating reliabilities in large-scale single-step genomic prediction

BackgroundIn this study, computationally efficient methods to approximate the reliabilities of genomic estimated breeding values (GEBV) in a single-step genomic prediction model including a residual polygenic (RPG) effect are described. In order to calculate the reliabilities of the genotyped animals, a single nucleotide polymorphism best linear unbiased prediction (SNPBLUP) or a genomic BLUP (GBLUP), was used, where two alternatives to account for the RPG effect were tested. In the direct approach, the genomic model included the RPG effect, while in the blended method, it did not but an index was used to weight the genomic and pedigree-based BLUP (PBLUP) reliabilities. In order to calculate the single-step GBLUP reliabilities for the breeding values for the non-genotyped animals, a simplified weighted-PBLUP model that included a general mean and additive genetic effects with weights accounting for the non-genomic and genomic information was used. We compared five schemes for the weights. Two datasets, i.e., a small (Data 1) one and a large (Data 2) one were used.ResultsFor the genotyped animals in Data 1, correlations between approximate reliabilities using the blended method and exact reliabilities ranged from 0.993 to 0.996 across three lactations. The slopes observed by regressing the reliabilities of GEBV from the exact method on those from the blended method were 1.0 for all three lactations. For Data 2, the correlations and slopes ranged, respectively, from 0.980 to 0.986 and from 0.91 to 0.96, and for the non-genotyped animals in Data 1, they ranged, respectively, from 0.987 to 0.994 and from 0.987 to 1, which indicate that the approximations were in line with the exact results. The best approach achieved correlations of 0.992 to 0.994 across lactations.ConclusionsOur results demonstrate that the approximated reliabilities calculated using our proposed approach are in good agreement with the exact reliabilities. The blended method for the genotyped animals is computationally more feasible than the direct method when RPG effects are included, particularly for large-scale datasets. The approach can serve as an effective strategy to estimate the reliabilities of GEBV in large-scale single-step genomic predictions.

A Note on the Conditioning of the H-1 Matrix Used in Single-Step GBLUP.

The single-step genomic BLUP (ssGBLUP) is used worldwide for the simultaneous genetic evaluation of genotyped and non-genotyped animals. It is easily extendible to all BLUP models by replacing the pedigree-based additive genetic relationship matrix (A) with an augmented pedigree-genomic relationship matrix (H). Theoretically, H does not introduce any artificially inflated variance. However, inflated genetic variances have been observed due to the incomparability between the genomic relationship matrix (G) and A used in H. Usually, G is blended and tuned with A22 (the block of A for genotyped animals) to improve its numerical condition and compatibility. If deflation/inflation is still needed, a common approach is weighting G-1-A22-1 in the form of τG-1-ωA22-1, added to A-1 to form H-1. In some situations, this can violate the conditional properties upon which H is built. Different ways of weighting the H-1 components (A-1, G-1, A22-1, and H-1 itself) were studied to avoid/minimise the violations of the conditional properties of H. Data were simulated on ten populations and twenty generations. Responses to weighting different components of H-1 were measured in terms of the regression of phenotypes on the estimated breeding values (the lower the slope, the higher the inflation) and the correlation between phenotypes and the estimated breeding values (predictive ability). Increasing the weight on H-1 increased the inflation. The responses to weighting G-1 were similar to those for H-1. Increasing the weight on A-1 (together with A22-1) was not influential and slightly increased the inflation. Predictive ability is a direct function of the slope of the regression line and followed similar trends. Responses to weighting G-1-A22-1 depend on the inflation/deflation of evaluations from A-1 to H-1 and the compatibility of the two matrices with the heritability used in the model. One possibility is a combination of weighting G-1-A22-1 and weighting H-1. Given recent advances in ssGBLUP, conditioning H-1 might become an interim solution from the past and then not be needed in the future.

Extension of the reduced animal model to single-step methods.

For a few decades, animal models (AMs) in the form of best linear unbiased prediction (BLUP) have been used for the genetic evaluation of animals. An equation system is set in the order of all the effects in the model, including all the animals in the pedigree. Solving these large equation systems has been a challenge. Reduced AM (RAM) was introduced in 1980, which allowed setting up equations for parents instead of all animals. That greatly reduced the number of equations to be solved. The RAM is followed by a back-solving step, in which progenies' breeding values are obtained conditional on parental breeding values. Initially, pedigree information was utilized to model genetic relationships between animals. With the availability of genomic information, genomic BLUP (GBLUP), single-step GBLUP (ssGBLUP), and single-step marker models were developed. Single-step methods utilize pedigree and genomic information for simultaneous genetic evaluation of genotyped and nongenotyped animals. There has been a shortage of studies on RAM development for genetic evaluation models utilizing genomic information. This study extended the concept of RAM from BLUP to the single-step methods. Using example data, three RAMs were described for ssGBLUP. The order of animal equations was reduced from the total number of animals to (1) genotyped animals and nongenotyped parents, (2) genotyped animals and nongenotyped phenotyped animals, and (3) genotyped animals and nongenotyped parents of phenotyped nongenotyped nonparents. Solutions for the remaining animals are obtained following a back-solving step. All the RAMs produced identical results to the full ssGBLUP. Advances in computational hardware have alleviated many computational limitations, but, on the other hand, the size of data is growing rapidly by the number of animals, traits, phenotypes, genotypes, and genotype density. There is an opportunity for a RAM comeback for the single-step methods to reduce the computational demands by reducing the number of equations.

Genome-wide association study for carcass traits in Hanwoo cattle using additional relatives' information of non-genotyped animals.

The aim of this study was to find significant genomic regions associated with carcass traits in Hanwoo cattle and to compare the benefit of using additional information from non-genotyped animals. Imputed whole-genome sequence data were used along with phenotypic data on 13 715 genotyped animals as well as phenotypes of 440 284 non-genotyped animals that were offspring of 454 genotyped sires. For carcass weight, 15 083 SNPs in 33 QTL regions and 313 candidate genes were identified. We found 410 SNPs in 17 QTL regions containing 122 candidate genes for back fat thickness. In total, 656 SNPs in 19 QTLs with 137 candidate genes for eye muscle area and 79 SNPs in 12 QTL regions with 77 candidate genes were identified for marbling score. The most important candidate genes included ZFAT, TG, PLAG1, CHCHD7, and TOX for carcass weight and eye muscle area, NOG for back fat thickness, and EVOVL5 for marbling score. This study showed that the use of phenotypic records on non-genotyped progeny along with imputed whole-genome sequence data increased the power of detecting new significant genomic regions.

On the equivalence between marker effect models and breeding value models and direct genomic values with the Algorithm for Proven and Young

BackgroundSingle-step genomic predictions obtained from a breeding value model require calculating the inverse of the genomic relationship matrix ({mathbf{G}}^{-1}). The Algorithm for Proven and Young (APY) creates a sparse representation of {mathbf{G}}^{-1} with a low computational cost. APY consists of selecting a group of core animals and expressing the breeding values of the remaining animals as a linear combination of those from the core animals plus an error term. The objectives of this study were to: (1) extend APY to marker effects models; (2) derive equations for marker effect estimates when APY is used for breeding value models, and (3) show the implication of selecting a specific group of core animals in terms of a marker effects model.ResultsWe derived a family of marker effects models called APY-SNP-BLUP. It differs from the classic marker effects model in that the row space of the genotype matrix is reduced and an error term is fitted for non-core animals. We derived formulas for marker effect estimates that take this error term in account. The prediction error variance (PEV) of the marker effect estimates depends on the PEV for core animals but not directly on the PEV of the non-core animals. We extended the APY-SNP-BLUP to include a residual polygenic effect and accommodate non-genotyped animals. We show that selecting a specific group of core animals is equivalent to select a subspace of the row space of the genotype matrix. As the number of core animals increases, subspaces corresponding to different sets of core animals tend to overlap, showing that random selection of core animals is algebraically justified.ConclusionsThe APY-(ss)GBLUP models can be expressed in terms of marker effect models. When the number of core animals is equal to the rank of the genotype matrix, APY-SNP-BLUP is identical to the classic marker effects model. If the number of core animals is less than the rank of the genotype matrix, genotypes for non-core animals are imputed as a linear combination of the genotypes of the core animals. For estimating SNP effects, only relationships and estimated breeding values for core animals are needed.

Correcting for base-population differences and unknown parent groups in single-step genomic predictions of Norwegian Red cattle.

Bias and inflation in genomic evaluation with the single-step methods have been reported in several studies. Incompatibility between the base-populations of the pedigree-based and the genomic relationship matrix (G) could be a reason for these biases. Inappropriate ways of accounting for missing parents could be another reason for biases in genetic evaluations with or without genomic information. To handle these problems, we fitted and evaluated a fixed covariate (J) that contains ones for genotyped animals and zeros for unrelated non-genotyped animals, or pedigree-based regression coefficients for related non-genotyped animals. We also evaluated alternative ways of fitting the J covariate together with genetic groups on biases and stability of breeding value estimates, and of including it into G as a random effect. In a whole vs. partial data set comparison, four scenarios were investigated for the partial data: genotypes missing, phenotypes missing, both genotypes and phenotypes missing, and pedigree missing. Fitting J either as fixed or random reduced level-bias and inflation and increased stability of genomic predictions as compared to the basic model where neither J nor genetic groups were fitted. In most models, genomic predictions were largely biased for scenarios with missing genotype and phenotype information. The biases were reduced for models which combined group and J effects. Models with these corrected group covariates performed better than the recently published model where genetic groups were encapsulated and fitted as random via the Quaas and Pollak transformation. In our Norwegian Red cattle data, a model which combined group and J regression coefficients was preferred because it showed least bias and highest stability of genomic predictions across the scenarios.

Imputation of non-genotyped F1 dams to improve genetic gain in swine crossbreeding programs.

This study investigated using imputed genotypes from non-genotyped animals which were not in the pedigree for the purpose of genetic selection and improving genetic gain for economically relevant traits. Simulations were used to mimic a 3-breed crossbreeding system that resembled a modern swine breeding scheme. The simulation consisted of three purebred (PB) breeds A, B, and C each with 25 and 425 mating males and females, respectively. Males from A and females from B were crossed to produce AB females (n = 1,000), which were crossed with males from C to produce crossbreds (CB; n = 10,000). The genome consisted of three chromosomes with 300 quantitative trait loci and ~9,000 markers. Lowly heritable reproductive traits were simulated for A, B, and AB (h2 = 0.2, 0.2, and 0.15, respectively), whereas a moderately heritable carcass trait was simulated for C (h2 = 0.4). Genetic correlations between reproductive traits in A, B, and AB were moderate (rg = 0.65). The goal trait of the breeding program was AB performance. Selection was practiced for four generations where AB and CB animals were first produced in generations 1 and 2, respectively. Non-genotyped AB dams were imputed using FImpute beginning in generation 2. Genotypes of PB and CB were used for imputation. Imputation strategies differed by three factors: 1) AB progeny genotyped per generation (2, 3, 4, or 6), 2) known or unknown mates of AB dams, and 3) genotyping rate of females from breeds A and B (0% or 100%). PB selection candidates from A and B were selected using estimated breeding values for AB performance, whereas candidates from C were selected by phenotype. Response to selection using imputed genotypes of non-genotyped animals was then compared to the scenarios where true AB genotypes (trueGeno) or no AB genotypes/phenotypes (noGeno) were used in genetic evaluations. The simulation was replicated 20 times. The average increase in genotype concordance between unknown and known sire imputation strategies was 0.22. Genotype concordance increased as the number of genotyped CB increased with little additional gain beyond 9 progeny. When mates of AB were known and more than 4 progeny were genotyped per generation, the phenotypic response in AB did not differ (P > 0.05) from trueGeno yet was greater (P < 0.05) than noGeno. Imputed genotypes of non-genotyped animals can be used to increase performance when 4 or more progeny are genotyped and sire pedigrees of CB animals are known.

Genome-enabled classification of stayability in Nellore cattle under a machine learning framework

Stayability (STAY) is a binary trait with significant value economically. It measures both the cow`s reproductive performance and longevity simultaneously. Thus, STAY is one of the most important female selection criterion in Nellore beef cattle breeding programs. The “success” for STAY is defined as the ability of a cow to stay in the herd up to 76 months of age and to have at least three calve. Despite its importance, STAY has not been investigated under a machine learning (ML) framework, which might allow to intuitively capture linear and nonlinear relationships (e.g., non-additive effects) between a response variable and other predictor variables. In this study, we compared different ML tools using a genome-enabled approach to classify daughters (non-genotyped animals but with STAY records) of genotyped sires. In total, 44,626 STAY records from daughters of 559 bulls genotyped with the 777K SNP panel were available for this study. The genotyped data were subdivided into three SNP sets based on the top-ranked effect on STAY: 1K-, 3K-, and 5K-SNP panels. The following ML algorithms were evaluated: AdaBoost (ADA), Naïve Bayes (NB), Decision Tree (DT), Deep Neural Network (DNN), k-Nearest Neighbors (NN), Multi-Layer Perceptron Neural Network (MLP), and Support Vector Machine (SVM). The analyses were performed using free Scikit-learn for the Python programming language. No relevant improvements in the learning process of the evaluated algorithms were observed when the number of SNPs in the genotype dataset was increased (i.e., 1K-, 3K-, or 5K-SNP panel). In short, NB outperformed the other algorithms considering, for example, the balanced accuracy (0.62 ± 0.01) and sensitivity (0.56 ± 0.02) metrics. In conclusion, the use of the 1K-SNP panel allowed efficient genomic classification and the NB algorithm outperformed the other methods as indicated by various classification metrics. To best of our knowledge, this is the first study using ML and genome-enabled classification of STAY in beef cattle.

Breeding value reliabilities for multiple-trait single-step genomic best linear unbiased predictor

Approximate multistep methods to calculate reliabilities for estimated breeding values in large genetic evaluations were developed for single-trait (ST-R2A) and multitrait (MT-R2A) single-step genomic BLUP (ssGBLUP) models. First, a traditional animal model was used to estimate the amount of nongenomic information for the genotyped animals. Second, this information was used with genomic data in a genomic BLUP model (genomic BLUP/SNP-BLUP) to approximate the total amount of information and ssGBLUP reliabilities for the genotyped animals. Finally, reliabilities for the nongenotyped animals were calculated using a traditional animal model where the increased information due to genomic data for the genotyped animals is accounted for by including pseudo-record counts for the genotyped animals. The approaches were tested using a multiple-trait ssGBLUP model on 2 data sets. The first data set (data 1) was small enough such that exact ssGBLUP model reliabilities could be computed by inversion and compared with the approximation method reliabilities. Data 1 had 46,535 first-, 35,290 second-, and 23,780 third-lactation 305-d milk yield records from 47,124 Finnish Red dairy cows. The pedigree comprised 64,808 animals, of which 19,757 were genotyped. We examined the efficiency of the MT-R2A approximation on a large data set (data 2) derived from the joint Nordic (Danish, Finnish, and Swedish) Holstein dairy cattle data. Data 2 had 17.8 million 305-d milk records from 8.3 million cows and first 3 lactations. The pedigree had 11 million animals of which 274,145 were genotyped on 46,342 SNP markers. For data 1, correlations between the exact ssGBLUP model and the ST-R2A for the genotyped (nongenotyped) animals were 0.995 (0.987), 0.965 (0.984), and 0.950 (0.983) for first, second, and third lactation, respectively. Correspondingly, correlations between exact ssGBLUP reliabilities and MT-R2A for the genotyped (nongenotyped) animals were 0.995 (0.993), 0.992 (0.991), and 0.990 (0.990) for first, second, and third lactation, respectively. The regression coefficients (b1) of ssGBLUP reliability on ST-R2A for the genotyped (nongenotyped) animals ranged from 0.87 (0.94) for first lactation to 0.68 (0.93) for third lactation, whereas for MT-R2A they were between 0.91 (0.99) for first lactation to 0.89 (0.99) for third lactation. Correspondingly, the intercepts varied from 0.11 (0.05) to 0.3 (0.06) for ST-R2A and from 0.06 (0.01) to 0.07 (0.02) for MT-R2A. The computing time for the approximation method was approximately 12% of that required by the direct exact approach. In conclusion, the developed approximate approach allows calculating estimated breeding value reliabilities in the ssGBLUP model even for large data sets.

Single-step genomic evaluation of milk production traits in Canadian Alpine and Saanen dairy goats

Genomic evaluations are routine in most plant and livestock breeding programs but are used infrequently in dairy goat breeding schemes. In this context, the purpose of this study was to investigate the use of the single-step genomic BLUP method for predicting genomic breeding values for milk production traits (milk, protein, and fat yields; protein and fat percentages) in Canadian Alpine and Saanen dairy goats. There were 6,409 and 12,236 Alpine records and 3,434 and 5,008 Saanen records for each trait in first and later lactations, respectively, and a total of 1,707 genotyped animals (833 Alpine and 874 Saanen). Two validation approaches were used, forward validation (i.e., animals born after 2013 with an average estimated breeding value accuracy from the full data set ≥0.50) and forward cross-validation (i.e., subsets of all animals included in the forward validation were used in successive replications). The forward cross-validation approach resulted in similar validation accuracies (0.55 to 0.66 versus 0.54 to 0.61) and biases (-0.01 to -0.07 versus -0.03 to 0.11) to the forward validation when averaged across traits. Additionally, both single and multiple-breed analyses were compared, and similar average accuracies and biases were observed across traits. However, there was a small gain in accuracy from the use of multiple-breed models for the Saanen breed. A small gain in validation accuracy for genomically enhanced estimated breeding values (GEBV) relative to pedigree-based estimated breeding values (EBV) was observed across traits for the Alpine breed, but not for the Saanen breed, possibly due to limitations in the validation design, heritability of the traits evaluated, and size of the training populations. Trait-specific gains in theoretical accuracy of GEBV relative to EBV for the validation animals ranged from 17 to 31% in Alpine and 35 to 55% in Saanen, using the cross-validation approach. The GEBV predicted from the full data set were 12 to 16% more accurate than EBV for genotyped animals, but no gains were observed for nongenotyped animals. The largest gains were found for does without lactation records (35-41%) and bucks without daughter records (46-54%), and consequently, the implementation of genomic selection in the Canadian dairy goat population would be expected to increase selection accuracy for young breeding candidates. Overall, this study represents the first step toward implementation of genomic selection in Canadian dairy goat populations.

Short communication: investigation of the feasibility of genomic selection in Icelandic Cattle.

Icelandic Cattle is a local dairy cattle breed in Iceland. With about 26,000 breeding females, it is by far the largest among the indigenous Nordic cattle breeds. The objective of this study was to investigate the feasibility of genomic selection in Icelandic Cattle. Pedigree-based best linear unbiased prediction (PBLUP) and single-step genomic best linear unbiased prediction (ssGBLUP) were compared. Accuracy, bias, and dispersion of estimated breeding values (EBV) for milk yield (MY), fat yield (FY), protein yield (PY), and somatic cell score (SCS) were estimated in a cross validation-based design. Accuracy (r^) was estimated by the correlation between EBV and corrected phenotype in a validation set. The accuracy (r^) of predictions using ssGBLUP increased by 13, 23, 19, and 20 percentage points for MY, FY, PY, and SCS for genotyped animals, compared with PBLUP. The accuracy of nongenotyped animals was not improved for MY and PY, but increased by 0.9 and 3.5 percentage points for FY and SCS. We used the linear regression (LR) method to quantify relative improvements in accuracy, bias (Δ^), and dispersion (b^) of EBV. Using the LR method, the relative improvements in accuracy of validation from PBLUP to ssGBLUP were 43%, 60%, 50%, and 48% for genotyped animals for MY, FY, PY, and SCS. Single-step GBLUP EBV were less underestimated (Δ^), and less overdispersed (b^) than PBLUP EBV for FY and PY. Pedigree-based BLUP EBV were close to unbiased for MY and SCS. Single-step GBLUP underestimated MY EBV but overestimated SCS EBV. Based on the average accuracy of 0.45 for ssGBLUP EBV obtained in this study, selection intensities according to the breeding scheme of Icelandic Cattle, and assuming a generation interval of 2.0 yr for sires of bulls, sires of dams and dams of bulls, genetic gain in Icelandic Cattle could be increased by about 50% relative to the current breeding scheme.