“…For example, two drought‐tolerant genes ZmVPP1 and ZmTIP1 discovered by GWAS, ZmVPP1 improved drought tolerance of Arabidopsis and maize by increased lateral root number and root biomass (Wang et al, 2016), and ZmTIP1 enhanced drought tolerance of maize through regulating the length of root hairs (Zhang et al, 2020). Several GWAS experiments have been performed for root traits at different growth stages and under different water treatments in maize, including root traits under hydroponic or soil conditions at seedling stage (Liu et al, 2021; Moussa et al, 2021; Pace et al, 2015a), under normal field conditions at maturation stage (Wang et al, 2021; Wu et al, 2022; Zheng et al, 2020), and under both drought stress and normal conditions at reproductive stage (Schneider et al, 2020; Zaidi et al, 2016). Although GWAS of maize root traits under drought stress or WW conditions have been attempted, few causal genes have been identified and elucidated for root system related to traits with maize drought tolerance.…”