“…Genome-wide association study (GWAS) is a powerful tool for gene mapping in plants and animals, and has been widely used for genetic dissection of complex quantitative traits in many major crop species ( Xiao et al , 2017 ; Liu and Yan, 2019 ; Cortes et al , 2021 ). In maize, GWAS has been successfully employed to understand the genetic basis of, and to identify candidate genes related to, flowering time and plant height ( Vanous et al , 2018 ; Li et al , 2020 ; Zheng et al , 2021a ), stalk traits ( Zhang et al , 2018 ; Mazaheri et al , 2019 ), tassel architecture ( Wu et al , 2016 ; Y. Wang et al , 2019 ), nitrogen use efficiency ( Morosini et al , 2017 ; Ertiro et al , 2020 ), low-phosphorus tolerance ( Xu et al , 2018 ; Q. J. Wang et al , 2019 ), cold tolerance ( Yi et al , 2021 ), drought tolerance ( Zaidi et al , 2016 ; Yuan et al , 2019 ; Liu and Qin, 2021 ), grain yield and yield-related traits ( Zhang et al , 2020a ; Yang et al , 2020 ; Ma and Cao, 2021 ), and grain quality ( Zhang et al , 2020b ; Zheng et al , 2021b ; Zhou et al , 2021 ). Moreover, several candidate genes involved in genetic control of root-related traits have been identified in maize by a GWAS approach.…”