Regreening mechanisms in cucumber: insights from a CsSIG2 mutation affecting chloroplast development

CsSIG2 is essential for cucumber chloroplast development, and mutations in CsSIG2 reveal mechanisms that restore chloroplast functionality and drive the regreening phenotype in the mutant. Chloroplast development and leaf color are essential traits that significantly influence plant photosynthesis and overall vigor. This study investigates a natural mutation in the cucumber that leads to a virescent leaf-color (Csvl-6) phenotype characterized by an initial yellow color in cotyledons and young leaves, which gradually transition to green as the plant matures. We utilized bulked segregant analysis and genetic linkage mapping to locate the best candidate gene sigma factor 2 (CsSIG2) on chromosome 6, identifying a single nonsynonymous SNP resulting in an arginine to glycine substitution in the CsSIG2 protein. Comparative transcriptome analysis highlighted that this mutation disrupts early chloroplast biogenesis and delays chlorophyll accumulation, but the chloroplasts can recover, leading to greening during later stages of leaf development. Our findings reveal that the recovery phenomenon involves upregulation of chloroplast-encoded genes responsible for thylakoid membrane formation and photosystem function, alongside altered expression of transcription factors linked to chlorophyll metabolism. This study elucidates the genetic and molecular basis of chloroplast development in cucumber, providing valuable insights into the mechanisms underlying leaf greening, which could inform future breeding efforts focused on manipulating leaf color traits for enhanced crop performance.