Genomic Selection Prediction Models Research Articles

Genome-Wide Association and Prediction of Traits Related to Salt Tolerance in Autotetraploid Alfalfa (Medicago sativa L.).

Soil salinity is a growing problem in world production agriculture. Continued improvement in crop salt tolerance will require the implementation of innovative breeding strategies such as marker-assisted selection (MAS) and genomic selection (GS). Genetic analyses for yield and vigor traits under salt stress in alfalfa breeding populations with three different phenotypic datasets was assessed. Genotype-by-sequencing (GBS) developed markers with allele dosage and phenotypic data were analyzed by genome-wide association studies (GWAS) and GS using different models. GWAS identified 27 single nucleotide polymorphism (SNP) markers associated with salt tolerance. Mapping SNPs markers against the Medicago truncatula reference genome revealed several putative candidate genes based on their roles in response to salt stress. Additionally, eight GS models were used to estimate breeding values of the training population under salt stress. Highest prediction accuracies and root mean square errors were used to determine the best prediction model. The machine learning methods (support vector machine and random forest) performance best with the prediction accuracy of 0.793 for yield. The marker loci and candidate genes identified, along with optimized GS prediction models, were shown to be useful in improvement of alfalfa with enhanced salt tolerance. DNA markers and the outcome of the GS will be made available to the alfalfa breeding community in efforts to accelerate genetic gains, in the development of biotic stress tolerant and more productive modern-day alfalfa cultivars.

Genotype × Environment Studies on Resistance to Late Leaf Spot and Rust in Genomic Selection Training Population of Peanut (Arachis hypogaea L.).

Foliar fungal diseases especially late leaf spot (LLS) and rust are the important production constraints across the peanut growing regions of the world. A set of 340 diverse peanut genotypes that includes accessions from gene bank of International Crops Research Institute for the Semi-Arid Tropics (ICRISAT), elite breeding lines from the breeding program, and popular cultivars were screened for LLS and rust resistance and yield traits across three locations in India under natural and artificial disease epiphytotic conditions. The study revealed significant variation among the genotypes for LLS and rust resistance at different environments. Combined analysis of variance revealed significant environment (E) and genotype × environment (G×E) interactions for both the diseases indicating differential response of genotypes in different environments. The present study reported 31 genotypes as resistant to LLS and 66 to rust across the locations at 90 DAS with maturity duration 103 to 128 days. Twenty-eight genotypes showed resistance to both the diseases across the locations, of which 19 derived from A. cardenasii, five from A. hypogaea, and four from A. villosa. Site regression and Genotype by Genotype x Environment (GGE) biplot analysis identified eight genotypes as stable for LLS, 24 for rust and 14 for pod yield under disease pressure across the environments. Best performing environment specific genotypes were also identified. Nine genotypes resistant to LLS and rust showed 77% to 120% increase in pod yield over control under disease pressure with acceptable pod and kernel features that can be used as potential parents in LLS and rust resistance breeding. Pod yield increase as a consequence of resistance offered to foliar fungal diseases suggests the possibility of considering ‘foliar fungal disease resistance’ as a must-have trait in all the peanut cultivars that will be released for cultivation in rainfed ecologies in Asia and Africa. The phenotypic data of the present study will be used for designing genomic selection prediction models in peanut.

Genomic Prediction of Autotetraploids; Influence of Relationship Matrices, Allele Dosage, and Continuous Genotyping Calls in Phenotype Prediction.

Estimation of allele dosage, using genomic data, in autopolyploids is challenging and current methods often result in the misclassification of genotypes. Some progress has been made when using SNP arrays, but the major challenge is when using next generation sequencing data. Here we compare the use of read depth as continuous parameterization with ploidy parameterizations in the context of genomic selection (GS). Additionally, different sources of information to build relationship matrices were compared. A real breeding population of the autotetraploid species blueberry (Vaccinium corybosum), composed of 1,847 individuals was phenotyped for eight yield and fruit quality traits over two years. Continuous genotypic based models performed as well as the best models. This approach also reduces the computational time and avoids problems associated with misclassification of genotypic classes when assigning dosage in polyploid species. This approach could be very valuable for species with higher ploidy levels or for emerging crops where ploidy is not well understood. To our knowledge, this work constitutes the first study of genomic selection in blueberry. Accuracies are encouraging for application of GS for blueberry breeding. GS could reduce the time for cultivar release by three years, increasing the genetic gain per cycle by 86% on average when compared to phenotypic selection, and 32% when compared with pedigree-based selection. Finally, the genotypic and phenotypic data used in this study are made available for comparative analysis of dosage calling and genomic selection prediction models in the context of autopolyploids.

Genomic Analysis and Prediction within a US Public Collaborative Winter Wheat Regional Testing Nursery.

The development of inexpensive, whole-genome profiling enables a transition to allele-based breeding using genomic prediction models. These models consider alleles shared between lines to predict phenotypes and select new lines based on estimated breeding values. This approach can leverage highly unbalanced datasets that are common to breeding programs. The Southern Regional Performance Nursery (SRPN) is a public nursery established by the USDA-ARS in 1931 to characterize performance and quality of near-release wheat ( L.) varieties from breeding programs in the US Central Plains. New entries are submitted annually and can be re-entered only once. The trial is grown at >30 locations each year and lines are evaluated for grain yield, disease resistance, and agronomic traits. Overall genetic gain is measured across years by including common check cultivars for comparison. We have generated whole-genome profiles via genotyping-by-sequencing (GBS) for 939 SPRN entries dating back to 1992 to explore the potential use of the nursery as a genomic selection (GS) training population (TP). The GS prediction models across years (average = 0.33) outperformed year-to-year phenotypic correlation for yield ( = 0.27) for a majority of the years evaluated, suggesting that genomic selection has the potential to outperform low heritability selection on yield in these highly variable environments. We also examined the predictability of programs using both program-specific and whole-set TPs. Generally, the predictability of a program was similar with both approaches. These results suggest that wheat breeding programs can collaboratively leverage the immense datasets that are generated from regional testing networks.

Genomic selection prediction models comparing sequence capture and SNP array genotyping methods

The successful application of genomic selection (GS) approaches is dependent on genetic makers derived from high-throughput and low-cost genotyping methods. Recent GS studies in trees have predominantly relied on SNP arrays as the source of genotyping, though this technology has a high entry cost. The recent development of alternative genotyping platforms, tailored to specific species and with low entry cost, has become possible due to advances in next-generation sequencing and genome complexity reduction methods such as sequence capture. However, the performance of these new platforms in GS models has not yet been evaluated, or compared to models developed from SNP arrays. Here, we evaluate the impact of these genotyping technologies on the development of GS prediction models for a Eucalyptus breeding population composed of 739 trees phenotyped for 13 wood quality and growth traits. Genotyping data obtained with both methods were compared for linkage disequilibrium, minor allele frequency, and missing data. Phenotypic prediction methods RR-BLUP and BayesB were employed, while predictive ability using cross validation was used to evaluate the performance of GS models derived from the different genotyping platforms. Differences in linkage disequilibrium patterns, minor allele frequency, missing data, and marker distribution were detected between sequence capture and SNP arrays. However, RR-BLUP and BayesB GS models resulted in similar predictive abilities. These results demonstrate that both genotyping methods are equivalent for genomic prediction of the traits evaluated. Sequence capture offers an alternative for species where SNP arrays are not available, or for when the initial development cost is too high.

Prediction accuracies for growth and wood attributes of interior spruce in space using genotyping-by-sequencing.

BackgroundGenomic selection (GS) in forestry can substantially reduce the length of breeding cycle and increase gain per unit time through early selection and greater selection intensity, particularly for traits of low heritability and late expression. Affordable next-generation sequencing technologies made it possible to genotype large numbers of trees at a reasonable cost.ResultsGenotyping-by-sequencing was used to genotype 1,126 Interior spruce trees representing 25 open-pollinated families planted over three sites in British Columbia, Canada. Four imputation algorithms were compared (mean value (MI), singular value decomposition (SVD), expectation maximization (EM), and a newly derived, family-based k-nearest neighbor (kNN-Fam)). Trees were phenotyped for several yield and wood attributes. Single- and multi-site GS prediction models were developed using the Ridge Regression Best Linear Unbiased Predictor (RR-BLUP) and the Generalized Ridge Regression (GRR) to test different assumption about trait architecture. Finally, using PCA, multi-trait GS prediction models were developed. The EM and kNN-Fam imputation methods were superior for 30 and 60% missing data, respectively. The RR-BLUP GS prediction model produced better accuracies than the GRR indicating that the genetic architecture for these traits is complex. GS prediction accuracies for multi-site were high and better than those of single-sites while multi-site predictability produced the lowest accuracies reflecting type-b genetic correlations and deemed unreliable. The incorporation of genomic information in quantitative genetics analyses produced more realistic heritability estimates as half-sib pedigree tended to inflate the additive genetic variance and subsequently both heritability and gain estimates. Principle component scores as representatives of multi-trait GS prediction models produced surprising results where negatively correlated traits could be concurrently selected for using PCA2 and PCA3.ConclusionsThe application of GS to open-pollinated family testing, the simplest form of tree improvement evaluation methods, was proven to be effective. Prediction accuracies obtained for all traits greatly support the integration of GS in tree breeding. While the within-site GS prediction accuracies were high, the results clearly indicate that single-site GS models ability to predict other sites are unreliable supporting the utilization of multi-site approach. Principle component scores provided an opportunity for the concurrent selection of traits with different phenotypic optima.Electronic supplementary materialThe online version of this article (doi:10.1186/s12864-015-1597-y) contains supplementary material, which is available to authorized users.

Genomic Relationship Matrix for Correcting Pedigree Errors in Breeding Populations: Impact on Genetic Parameters and Genomic Selection Accuracy

ABSTRACTQuantitative genetic analyses aim to estimate genetic parameters and breeding values to select superior parents, families, and individuals. For these estimates a relationship matrix derived from the pedigree typically is used in a mixed model framework. However, breeding is a complex, multistep process and errors in the pedigree are common. Because errors reduce the accuracy of genetic parameter estimates and affect genetic gain, it is important to correct these errors. Here we show that a realized relationship matrix (RRM) derived from single nucleotide polymorphism markers based on the normality of the relationship coefficients can be used to correct pedigree errors. For a loblolly pine (Pinus taeda L.) breeding population, errors in the pedigree were detected and corrected with the RRM. With the corrected pedigree, best linear unbiased predictor (BLUP) models fit the data significantly better for 14 out of 15 traits evaluated, and the predictive ability of the genomic selection models using ridge regression BLUP increased for 13 traits. The corrected pedigree based on the normality of the relationship coefficients improves accuracy of traditional estimations of heritability and breeding values as well as genomic selection predictions. As more breeding programs begin to use genomic selection, we recommend first using the dense panel of markers to correct pedigree errors and then using the improved information to develop genomic selection prediction models.

Accuracy of Genomic Selection Methods in a Standard Data Set of Loblolly Pine (Pinus taedaL.)

Genomic selection can increase genetic gain per generation through early selection. Genomic selection is expected to be particularly valuable for traits that are costly to phenotype and expressed late in the life cycle of long-lived species. Alternative approaches to genomic selection prediction models may perform differently for traits with distinct genetic properties. Here the performance of four different original methods of genomic selection that differ with respect to assumptions regarding distribution of marker effects, including (i) ridge regression–best linear unbiased prediction (RR–BLUP), (ii) Bayes A, (iii) Bayes Cπ, and (iv) Bayesian LASSO are presented. In addition, a modified RR–BLUP (RR–BLUP B) that utilizes a selected subset of markers was evaluated. The accuracy of these methods was compared across 17 traits with distinct heritabilities and genetic architectures, including growth, development, and disease-resistance properties, measured in a Pinus taeda (loblolly pine) training population of 951 individuals genotyped with 4853 SNPs. The predictive ability of the methods was evaluated using a 10-fold, cross-validation approach, and differed only marginally for most method/trait combinations. Interestingly, for fusiform rust disease-resistance traits, Bayes Cπ, Bayes A, and RR–BLUB B had higher predictive ability than RR–BLUP and Bayesian LASSO. Fusiform rust is controlled by few genes of large effect. A limitation of RR–BLUP is the assumption of equal contribution of all markers to the observed variation. However, RR-BLUP B performed equally well as the Bayesian approaches.The genotypic and phenotypic data used in this study are publically available for comparative analysis of genomic selection prediction models.

Accelerating the domestication of trees using genomic selection: accuracy of prediction models across ages and environments

• Genomic selection is increasingly considered vital to accelerate genetic improvement. However, it is unknown how accurate genomic selection prediction models remain when used across environments and ages. This knowledge is critical for breeders to apply this strategy in genetic improvement. • Here, we evaluated the utility of genomic selection in a Pinus taeda population of c. 800 individuals clonally replicated and grown on four sites, and genotyped for 4825 single-nucleotide polymorphism (SNP) markers. Prediction models were estimated for diameter and height at multiple ages using genomic random regression best linear unbiased predictor (BLUP). • Accuracies of prediction models ranged from 0.65 to 0.75 for diameter, and 0.63 to 0.74 for height. The selection efficiency per unit time was estimated as 53-112% higher using genomic selection compared with phenotypic selection, assuming a reduction of 50% in the breeding cycle. Accuracies remained high across environments as long as they were used within the same breeding zone. However, models generated at early ages did not perform well to predict phenotypes at age 6 yr. • These results demonstrate the feasibility and remarkable gain that can be achieved by incorporating genomic selection in breeding programs, as long as models are used at the relevant selection age and within the breeding zone in which they were estimated.

Effect of BLUP prediction on genomic selection: practical considerations to achieve greater accuracy in genomic selection

Background Prediction of breeding values (BV) using only genotypic information is the final goal of Genomic Selection (GS) [1]. Commonly, BV prediction from traditional BLUP analysis is the input for constructing GS prediction models, and GS predicted BVs are correlated with traditional BLUP BVs to estimate the accuracy of GS models. The use of GS in plant breeding depends on the accuracy of the GS models to predict the BVs. Therefore, better accuracy and less bias in traditional BLUP BVs should improve the final accuracy of GS predictions. Such improvements in GS predictions are not due to GS modeling itself, but rather to the reduced noise in the BLUP BV used as input. Improvements in BLUP BV can be obtained simply by correcting errors in the pedigree [2] or using more complex approaches, such as applying a realized relationship matrix (RRM) in the BLUP prediction as an alternative to the relationship matrix (A) based on expected values derived from the pedigree [3]. Misspecification of effects in BLUP models tends to produce upward bias in the BV estimates, which also impact GS accuracy [4]. In addition, not correcting with the additive-genetic relationship information in the GS prediction model leads to overestimates in accuracies due to inadequate accounting for confounding genetic relationships found in the training population [5]. The inflated accuracy cannot be exploited in future generations and should be guarded against. Our objective was to use real data to study the effect on the GS accuracy from 1) pedigree errors, 2) incorporation of the RRM in the BLUP analysis, 3) misspecification of non-additive effects in the BLUP analysis and 4) the effect of ignoring the additive-genetic relationship in the GS prediction model.

Stability of Genomic Selection prediction models across ages and environments

Background A tree breeding program is characterized by long generation intervals which, over time, result in a much smaller number of breeding cycles when compared to annual crops. Moreover, most economically important traits in a tree-breeding program are quantitatively inherited, display low heritability and are expressed late in the life cycle. Genomic Selection (GS) is expected to be particularly valuable for tree species, leading to shorter generation intervals and improved genetic gain over time. The main factors that affect the accuracy of GS prediction models are the level of linkage disequilibrium (LD) in the training population, the training population size, the heritability of the trait and the number of QTL regulating its variation. However, it is yet largely unknown how stable prediction models are across environments and different ages. This knowledge is critical for tree breeders that wish to use genomic selection in their genetic improvement program. Here, we report the first assessment of the utility of genomic selection in a conifer species. We developed prediction models for growth traits measured at multiple sites, to evaluate the impact of genotype by environment interactions in their accuracy. Training populations were also measured over multiple ages and models were developed to assess their value in predicting breeding values later in the lifecycle.