Phenomic selection in wheat breeding: prediction of the genotype-by-environment interaction in multi-environment breeding trials.

KEY MESSAGE: Phenomic prediction of wheat grain yield and heading date in different multi-environmental trial scenarios is accurate. Modelling the genotype-by-environment interaction effect using phenomic data is a potentially low-cost complement to genomic prediction. The performance of wheat cultivars in multi-environmental trials (MET) is difficult to predict because of the genotype-by-environment interactions (G × E). Phenomic selection is supposed to be efficient for modelling the G × E effect because it accounts for non-additive effects. Here, phenomic data are near-infrared (NIR) spectra obtained from plant material. While phenomic selection has recently been shown to accurately predict wheat grain yield in single environments, its accuracy needs to be investigated for MET. We used four datasets from two winter wheat breeding programs to test and compare the predictive abilities of phenomic and genomic models for grain yield and heading date in different MET scenarios. We also compared different methods to model the G × E using different covariance matrices based on spectra. On average, phenomic and genomic prediction abilities are similar in all different MET scenarios. Better predictive abilities were obtained when G × E effects were modelled with NIR spectra than without them, and it was better to use all the spectra of all genotypes in all environments for modelling the G × E. To facilitate the implementation of phenomic prediction, we tested MET designs where the NIR spectra were measured only on the genotype-environment combinations phenotyped for the target trait. Missing spectra were predicted with a weighted multivariate ridge regression. Intermediate predictive abilities for grain yield were obtained in a sparse testing scenario and for new genotypes, which shows that phenomic selection is an efficient and practicable prediction method for dealing with G × E.

Phenomic selection in wheat breeding: identification and optimisation of factors influencing prediction accuracy and comparison to genomic selection.

KEY MESSAGE: Phenomic selection is a promising alternative or complement to genomic selection in wheat breeding. Models combining spectra from different environments maximise the predictive ability of grain yield and heading date of wheat breeding lines. Phenomic selection (PS) is a recent breeding approach similar to genomic selection (GS) except that genotyping is replaced by near-infrared (NIR) spectroscopy. PS can potentially account for non-additive effects and has the major advantage of being low cost and high throughput. Factors influencing GS predictive abilities have been intensively studied, but little is known about PS. We tested and compared the abilities of PS and GS to predict grain yield and heading date from several datasets of bread wheat lines corresponding to the first or second years of trial evaluation from two breeding companies and one research institute in France. We evaluated several factors affecting PS predictive abilities including the possibility of combining spectra collected in different environments. A simple H-BLUP model predicted both traits with prediction ability from 0.26 to 0.62 and with an efficient computation time. Our results showed that the environments in which lines are grown had a crucial impact on predictive ability based on the spectra acquired and was specific to the trait considered. Models combining NIR spectra from different environments were the best PS models and were at least as accurate as GS in most of the datasets. Furthermore, a GH-BLUP model combining genotyping and NIR spectra was the best model of all (prediction ability from 0.31 to 0.73). We demonstrated also that as for GS, the size and the composition of the training set have a crucial impact on predictive ability. PS could therefore replace or complement GS for efficient wheat breeding programs.

BWGS: A R package for genomic selection and its application to a wheat breeding programme.

We developed an integrated R library called BWGS to enable easy computation of Genomic Estimates of Breeding values (GEBV) for genomic selection. BWGS, for BreedWheat Genomic selection, was developed in the framework of a cooperative private-public partnership project called Breedwheat (https://breedwheat.fr) and relies on existing R-libraries, all freely available from CRAN servers. The two main functions enable to run 1) replicated random cross validations within a training set of genotyped and phenotyped lines and 2) GEBV prediction, for a set of genotyped-only lines. Options are available for 1) missing data imputation, 2) markers and training set selection and 3) genomic prediction with 15 different methods, either parametric or semi-parametric. The usefulness and efficiency of BWGS are illustrated using a population of wheat lines from a real breeding programme. Adjusted yield data from historical trials (highly unbalanced design) were used for testing the options of BWGS. On the whole, 760 candidate lines with adjusted phenotypes and genotypes for 47 839 robust SNP were used. With a simple desktop computer, we obtained results which compared with previously published results on wheat genomic selection. As predicted by the theory, factors that are most influencing predictive ability, for a given trait of moderate heritability, are the size of the training population and a minimum number of markers for capturing every QTL information. Missing data up to 40%, if randomly distributed, do not degrade predictive ability once imputed, and up to 80% randomly distributed missing data are still acceptable once imputed with Expectation-Maximization method of package rrBLUP. It is worth noticing that selecting markers that are most associated to the trait do improve predictive ability, compared with the whole set of markers, but only when marker selection is made on the whole population. When marker selection is made only on the sampled training set, this advantage nearly disappeared, since it was clearly due to overfitting. Few differences are observed between the 15 prediction models with this dataset. Although non-parametric methods that are supposed to capture non-additive effects have slightly better predictive accuracy, differences remain small. Finally, the GEBV from the 15 prediction models are all highly correlated to each other. These results are encouraging for an efficient use of genomic selection in applied breeding programmes and BWGS is a simple and powerful toolbox to apply in breeding programmes or training activities.

Exploiting the Repetitive Fraction of the Wheat Genome for High-Throughput Single-Nucleotide Polymorphism Discovery and Genotyping.

Transposable elements (TEs) account for more than 80% of the wheat genome. Although they represent a major obstacle for genomic studies, TEs are also a source of polymorphism and consequently of molecular markers such as insertion site-based polymorphism (ISBP) markers. Insertion site-based polymorphisms have been found to be a great source of genome-specific single-nucleotide polymorphism (SNPs) in the hexaploid wheat ( L.) genome. Here, we report on the development of a high-throughput SNP discovery approach based on sequence capture of ISBP markers. By applying this approach to the reference sequence of chromosome 3B from hexaploid wheat, we designed 39,077 SNPs that are evenly distributed along the chromosome. We demonstrate that these SNPs can be efficiently scored with the KASPar (Kompetitive allele-specific polymerase chain reaction) genotyping technology. Finally, through genetic diversity and genome-wide association studies, we also demonstrate that ISBP-derived SNPs can be used in marker-assisted breeding programs.

Association study between the gibberellic acid insensitive gene and leaf length in a Lolium perenne L. synthetic variety.

BACKGROUND: Association studies are of great interest to identify genes explaining trait variation since they deal with more than just a few alleles like classical QTL analyses. They are usually performed using collections representing a wide range of variability but which could present a genetic substructure. The aim of this paper is to demonstrate that association studies can be performed using synthetic varieties obtained after several panmictic generations. This demonstration is based on an example of association between the gibberellic acid insensitive gene (GAI) polymorphism and leaf length polymorphism in 'Herbie', a synthetic variety of perennial ryegrass. METHODS: Leaf growth parameters, consisted of leaf length, maximum leaf elongation rate (LERmax) and leaf elongation duration (LED), were evaluated in spring and autumn on 216 plants of Herbie with three replicates. For each plant, a sequence of 370 bp in GAI was analysed for polymorphism. RESULTS: Genetic effect was highly significant for all traits. Broad sense heritabilities were higher for leaf length and LERmax with about 0.7 in each period and 0.5 considering both periods than for LED with about 0.4 in each period and 0.3 considering both periods. GAI was highly polymorphic with an average of 12 bp between two consecutive SNPs and 39 haplotypes in which 9 were more frequent. Linkage disequilibrium declined rapidly with distance with r 2 values lower than 0.2 beyond 150 bp. Sequence polymorphism of GAI explained 8-14% of leaf growth parameter variation. A single SNP explained 4% of the phenotypic variance of leaf length in both periods which represents a difference of 33 mm on an average of 300 mm. CONCLUSIONS: Synthetic varieties in which linkage disequilibrium declines rapidly with distance are suitable for association studies using the "candidate gene" approach. GAI polymorphism was found to be associated with leaf length polymorphism which was more correlated to LERmax than to LED in Herbie. It is a good candidate to explain leaf length variation in other plant material.