Leaf shape and vertical distribution of soybean affect canopy structure,photosynthetic efficiency,and yield.The exis-tence of different leaf shapes and sizes on the same plant,which is known as heterophylly,has been observed in many flowering plant species.Yet,the genetic characteristics and genetic basis of heterophylly in soybean remain unknown.In this study,leaf characteristics such as leaf length,leaf width,leaf shape index,and heterophylly index were investigated in 283 soybean germ-plasm resources for two consecutive years in Nanjing,Jiangsu Province.A total of 181 related loci were detected by genome-wide association study(GWAS),among which 18 loci could be repeatedly detected in two environments or among multiple traits.Us-ing the loci associated with leaf characteristics,we integrated the GWAS approach with the expression profiling data and gene-based association and functional annotation of orthologs in Arabidopsis to identify candidate genes involved in leaf devel-opment in soybean.The known soybean leaf shape regulatory gene Ln(Glyma.20G116200)was found upstream of locus Chr20:36152820.In addition,two candidate genes(Glyma.19G192700 and Glyma.19G194100)were identified near the related locus Chr19:45155943 on chromosome 19,homologous genes of growth-regulating factor 4(GRF4),and LITTLE ZIPPER 3(ZPR3),respectively.These results lay a solid foundation for expanding our understanding of the genetic mechanism of hetero-phylly in soybean.