In plants and animals,male gamete differentiates and ferti-lizes an egg cell,which develops into different types of specialized cells,tissues,and ultimately an individual.Dur-ing this complex developmental process,germline develop-ment is critical,because it is exquisitely regulated to ensure proper inheritance of genetic and epigenetic information to the progenies for the robust fitness to the environment (Li et al.,2020).In higher plants,male germline undergoes DNA methylation reprogramming,which involves de novo me-thylation and alters gene expression (Walker et al.,2018).