Information on crop pedigrees can be used to help maximise genetic gain in crop breeding and allow efficient management of genetic resources. We present a pedigree resource of 2,657 wheat (Triticum aestivum L.) genotypes originating from 38 countries, representing more than a century of breeding and variety development. Visualisation of the pedigree enables illustration of the key developments in United Kingdom wheat breeding, highlights the wide genetic background of the UK wheat gene pool, and facilitates tracing the origin of beneficial alleles. A relatively high correlation between pedigree- and marker-based kinship coefficients was found, which validated the pedigree and enabled identification of errors in the pedigree or marker data. Using simulations with a combination of pedigree and genotype data, we found evidence for significant effects of selection by breeders. Within crosses, genotypes are often more closely related than expected by simulations to one of the parents, which indicates selection for favourable alleles during the breeding process. Selection across the pedigree was demonstrated on a subset of the pedigree in which 110 genotyped varieties released before the year 2000 were used to simulate the distribution of marker alleles of 45 genotyped varieties released after the year 2000, in the absence of selection. Allelic diversity in the 45 varieties was found to deviate significantly from the simulated distributions at a number of loci, indicating regions under selection over this period. The identification of one of these regions as coinciding with a strong yield component quantitative trait locus (QTL) highlights both the potential of the remaining loci as wheat breeding targets for further investigation, as well as the utility of this pedigree-based methodology to identify important breeding targets in other crops. Further evidence for selection was found as greater linkage disequilibrium (LD) for observed versus simulated genotypes within all chromosomes. This difference was greater at shorter genetic distances, indicating that breeder selections have conserved beneficial linkage blocks. Collectively, this work highlights the benefits of generating detailed pedigree resources for crop species. The wheat pedigree database developed here represents a valuable community resource and will be updated as new varieties are released at https://www.niab.com/pages/id/501/UK_Wheat_varieties_Pedigree. Breeding activities undertaken in the world’s most important crop species have resulted in large increases in yield potential over the last century. Bread wheat is a key crop for both human and animal nutrition worldwide. To help inform future breeding and research activities, we have developed a pedigree resource of over 2,600 bread wheat accessions, originating from 38 countries and representing more than a century of breeding and variety development. Pedigree-based relationships between lines are largely confirmed by genetic marker data. By combining the genetic and pedigree data sets, we are able to identify genetic signatures of selection across the pedigree, identifying genomic regions selected for via modern breeding activities. The resource developed here will serve as an evolving platform to inform and manage wheat genetic diversity in breeding programmes around the world and highlights the utility of developing and exploiting similar resources in other crop species.
Background: Selection has dramatically shaped genetic and phenotypic variation in bread wheat. We can assess the genomic basis of historical phenotypic changes, and the potential for future improvement, using experimental populations that attempt to undo selection through the randomizing effects of recombination. Results: We bred the NIAB Diverse MAGIC multi-parent population comprising over 500 recombinant inbred lines, descended from sixteen historical UK bread wheat varieties released between 1935 and 2004. We sequence the founders’ genes and promoters by capture, and the MAGIC population by low-coverage whole-genome sequencing. We impute 1.1 M high-quality SNPs that are over 99% concordant with array genotypes. Imputation accuracy only marginally improves when including the founders’ genomes as a haplotype reference panel. Despite capturing 73% of global wheat genetic polymorphism, 83% of genes cluster into no more than three haplotypes. We phenotype 47 agronomic traits over 2 years and map 136 genome-wide significant associations, concentrated at 42 genetic loci with large and often pleiotropic effects. Around half of these overlap known quantitative trait loci. Most traits exhibit extensive polygenicity, as revealed by multi-locus shrinkage modelling. Conclusions: Our results are consistent with a gene pool of low haplotypic diversity, containing few novel loci of large effect. Most past, and projected future, phenotypic changes arising from existing variation involve fine-scale shuffling of a few haplotypes to recombine dozens of polygenic alleles of small effect. Moreover, extensive pleiotropy means selection on one trait will have unintended consequences, exemplified by the negative trade-off between yield and protein content, unless selection and recombination can break unfavorable trait-trait associations.
Aims: Selection for optimal root system architecture (RSA) is important to ensure genetic gains in the sustainable production of wheat (Triticum aestivum L.). Here we examine the hypothesis that past wheat breeding has led to changes in RSA and that future breeding efforts can focus directly on RSA to improve adaptation to target environments. Methods: We conducted field trials using diverse wheat varieties, including modern and historic UK varieties and non-UK landraces, tested under contrasting tillage regimes (non-inversion tillage versus conventional ploughing) for two trial years or different seeding rates (standard versus high rate) for one trial year. We used field excavation, washing and measurement of root crowns (‘shovelomics’) to characterise RSA traits, including: numbers of seminal, crown and nodal roots per plant, and crown root growth angle. Results: We found differences among genotypes for all root traits. Modern varieties generally had fewer roots per plant than historic varieties. On average, there were fewer crown roots and root angles were wider under shallow non-inversion tillage compared with conventional ploughing. Crown root numbers per plant also tended to be smaller at a high seeding rate compared with the standard. There were significant genotype-by-year, genotype-by-tillage and genotype-by-seeding-rate interactions for many root traits. Conclusions: Smaller root systems are likely to be a result of past selection that facilitated historical yield increases by reducing below-ground competition within the crop. The effects of crop management practices on RSA depend on genotype, suggesting that future breeding could select for improved RSA traits in resource-efficient farming systems.
Linking high-throughput environmental data (enviromics) to genomic prediction (GP) is a cost-effective strategy for increasing selection intensity under genotype-by-environment interactions (G × E). This study developed a data-driven approach based on Environment–Phenotype Association (EPA) aimed at recycling important G × E information from historical breeding data. EPA was developed in two applications: (1) scanning a secondary source of genetic variation, weighted from the shared reaction-norms of past-evaluated genotypes and (2) pinpointing weights of the similarity among trial-sites (locations), given the historical impact of each envirotyping data variable for a given site. These results were then used as a dimensionality reduction strategy, integrating historical data to feed multi-environment GP models, which led to the development of four new G × E kernels considering genomics, enviromics, and EPA outcomes. The wheat trial data used included 36 locations, 8 years, and three target populations of environments (TPEs) in India. Four prediction scenarios and six kernel models within/across TPEs were tested. Our results suggest that the conventional GBLUP, without enviromic data or when omitting EPA, is inefficient in predicting the performance of wheat lines in future years. Nevertheless, when EPA was introduced as an intermediary learning step to reduce the dimensionality of the G × E kernels while connecting phenotypic and environmental-wide variation, a significant enhancement of G × E prediction accuracy was evident. EPA revealed that the effect of seasonality makes strategies such as “covariable selection” unfeasible because G × E is year-germplasm specific. We propose that the EPA effectively serves as a “reinforcement learner” algorithm capable of uncovering the effect of seasonality over the reaction-norms, with the benefits of better forecasting the similarities between past and future trialing sites. EPA combines the benefits of dimensionality reduction while reducing the uncertainty of genotype-by-year predictions and increasing the resolution of GP for the genotype-specific level.
The extent to which the quality and yield of plant varieties are influenced by the environment is important for their successful uptake by end users particularly as climatic fluctuations are resulting in environments that are highly variable from one growing season to another. The genotype-by-environment interaction (GEI) of milling quality and yield was studied using four winter oat varieties in multi-locational trials over 4 years in the U.K. Significant differences across the 22 environments were found between physical grain quality and composition as well as grain yield, with the environment having a significant effect on all of the traits measured. Grain yield was closely related to grain number m−2 whereas milling quality traits were related to grain size attributes. Considerable genotype by environment interaction was obtained for all grain quality traits and stability analysis revealed that the variety Mascani was the least sensitive to the environment for all milling quality traits measured whereas the variety Balado was the most sensitive. Examination of environmental conditions at specific within-year stages of crop development indicated that both temperature and rainfall during grain development were correlated with grain yield and β-glucan content and with the ease of removing the hull (hullability).
Cereal crop breeders have achieved considerable genetic gain in genetically complex traits, such as grain yield, while maintaining genetic diversity. However, focus on selection for yield has negatively impacted other important traits. To better understand multi-trait selection within a breeding context, and how it might be optimized, we analysed genotypic and phenotypic data from a genetically diverse, 16-founder wheat multi-parent advanced generation inter-cross population. Compared to single-trait models, multi-trait ensemble genomic prediction models increased prediction accuracy for almost 90 % of traits, improving grain yield prediction accuracy by 3–52 %. For complex traits, non-parametric models (Random Forest) also outperformed simplified, additive models (LASSO), increasing grain yield prediction accuracy by 10–36 %. Simulations of recurrent genomic selection then showed that sustained greater forward prediction accuracy optimized long-term genetic gains. Simulations of selection on grain yield found indirect responses in related traits, involving optimized antagonistic trait relationships. We found multi-trait selection indices could effectively optimize undesirable relationships, such as the trade-off between grain yield and protein content, or combine traits of interest, such as yield and weed competitive ability. Simulations of phenotypic selection found that including Random Forest rather than LASSO genetic models, and multi-trait rather than single-trait models as the true genetic model accelerated and extended long-term genetic gain whilst maintaining genetic diversity. These results (i) suggest important roles of pleiotropy and epistasis in the wider context of wheat breeding programmes, and (ii) provide insights into mechanisms for continued genetic gain in a limited genepool and optimization of multiple traits for crop improvement.