Increasing world population, global climate change, de-creased farmland, environmental pollution and ecological deterioration represent unprecedent challenges for crop production to ensure global food security (Hickey et al., 2019;Li et al., 2021a). It is estimated that by the year 2050, 50％more food is needed to feed the increasing population (Bailey-Serres et al., 2019). Thus, it is urgent to boost crop production by using cutting-edge technologies. Genome editing technologies have revolutionized the plant research field and offer great potential in crop improvement (Ma et al., 2015; Li et al., 2021b; Zhan et al., 2021). Of the several ge-nome editing technologies, clustered regularly interspaced palindromic repeats (CRISPR) and its associated Cas protein (CRISPR/Cas), has become the dominant one in the past several years because it is simple, efficient and cost-effective.