“…With the currently available high-throughput genotyping technologies, genome-wide association studies (GWAS) have been developed as a powerful approach to characterize the genetic basis of complex traits and locus-locus interactions in plants (Nordborg and Weigel, 2008). More recent studies combining GWAS with metabolomics (mGWAS) demonstrate a powerful forward genetics strategy that can be used to dissect the genetic and biochemical basis of plant metabolism, including that in model species, such as Arabidopsis thaliana, tomato, rice (Oryza sativa), and Medicago truncatula, and other crop species, such as barley, perilla (Perilla ocymoides), and pepper (Capsicum annuum) (Achnine et al, 2005;Yamazaki et al, 2008;Widodo et al, 2009;Wahyuni et al, 2011;Carreno-Quintero et al, 2013;Luo, 2015). Moreover, recent studies have revealed that linkage disequilibrium mapping in combination with linkage mapping improves the ability to identify important loci.…”