Genome based QTL and meta-QTL analyses of thousand-kernel weight in tetraploid wheat

Wheat domestication and subsequent improvement formed a wide phenotypic variation in Grain Weight (GW) between the domesticated wheat species and their wild progenitors. GW continues to be an important goal of many wheat-breeding programs and yet, although studies found many quantitative trait loci (QTLs) for GW, not many genes that underlaythese loci were identified. Here we performed QTL analysis for GW using a Recombinant Inbred Line (RIL) population based of a cross between wild emmer wheat accession 9Zavitan9 and durum wheat variety 9Svevo9. Using the recent Zavitan genome assembly, we anchored the identified QTLs to the reference sequence and added the positions of previously published QTLs for GW in tetraploid wheat. This genome based meta-QTL analysis enabled us to identify a locus on chromosome 6A with a positive effect on GW that was contributed by wild wheat in a few studies. This locus was validated using an introgression line that contains the 6A GW QTL from Zavitan in the background of Svevo with higher grain weight. Using the reference sequence and genes associated with GW from rice, we were able to identify a wheat ortholog in the 6A QTL region to rice gene, OsGRF4. The coding sequence of this gene, TtGRF4-A, showed four SNPs between Zavitan and Svevo. Molecular marker developed for the first SNP showed that the Zavitan allele of TtGRF4-A is rare in a core collection of wild emmer and absent in domesticated emmer genepool. We suggest that TtGRF4-A is a candidate underlying the 6A GW QTL and breeding with its natural Zavitan allele may have the potential to increase wheat yields.