Modulation of evening complex activity enables north-to-south adaptation of soybean

Soybean,a typical short-day crop,is sensitive to photoperiod,which is a major limiting factor defining its north-to-south cultivation range.The long-juvenile(LJ)trait is controlled primarily by the J locus which has been used for decades by soybean breeders to delay flowering and improve grain yield in tropical regions.The J gene encodes an ortholog of the Arabidopsis Evening Complex(EC)component EARLY FLOWERING 3(ELF3).To identify modifiers of J,we conducted a forward genetic screen and isolated a mutant(eoj57)that in combination with j has longer flowering delay compared with j single mutant plants.Map-based cloning and genome re-sequencing identified eoj57(designated as GmLUX2)as an ortholog of the Arabidopsis EC component LUX ARRHYTHMO(LUX).To validate that GmLUX2 is a modifier of J,we used trans-complementation and identified a natural variant allele with a similar phenotype.We also show that GmLUX2 physically interacts with GmELF3a/b and binds DNA,whereas the mutant and natural variant are attenuated in both activities.Transcriptome analysis shows that the GmLUX2-GmELF3a complex co-regulates the expression of several circadian clock-associated genes and directly represses E1 expression.These results provide mechanistic insight into how GmLUX2-GmELF3 controls flowering time via synergistic regulation of gene expression.These novel insights expand our understanding of the regulation of the EC complex,and facilitate the development of soybean varieties adapted for growth at lower latitudes.