Genetic architecture affecting maize agronomic traits identified by variance heterogeneity association mapping

Conventional genome-wide association studies (GWAS) focused on the phenotypic mean differences (mGWAS) but often ignored genetic variants influencing differences in the variance between genotypes. In this study, we performed variance heterogeneity GWAS (vGWAS) analysis for 13 previously measured agronomic traits in a maize population. We discovered a total of 129 significant SNPs. We demonstrated that the genetic loci influencing mean differences and variance heterogeneity formed distinct groups, suggesting that breeders were able to independently select for phenotype mean and variance values. Moreover, vGWAS served as a tractable approach to effectively identify 214 epistatic interaction pairs. In addition, we documented four agronomic traits with decreasing phenotype variance during modern maize breeding history and identified the potential genetic variants influencing this process. In summary, we discovered additional non-additive effects contributing to missing heritability and valuable genetic variants used for breeding varieties with desired phenotypic variance.