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A total of 16  OsS40  genes of  Oryza sativa  were identified in our previous work, but their functions remain unclear. In this study, 13  OsS40  members were knocked out using the CRISPR/cas9 gene-editing technology. After screening phenotype characterization of CRISPR/Cas9 mutants compared to WT, five  oss40s  mutants exhibited a stay-green phenotype at 30 days after heading. Moreover, increased grain size and grain weight occurred in the  oss40-1, oss40-12 , and  oss40-14  lines, while declined grain weight appeared in the  oss40-7  and  oss40-13  mutants. The transcript levels of several senescence-associated genes (SAGs), chlorophyll degradation-related genes (CDGs), as well as WRKY members were differentially decreased in the five stay-green  oss40s  mutants compared to WT. Five  oss40  mutants also exhibited a stay-green phenotype when the detached leaves were incubated under darkness for 4 days.  OsSWEET4  and  OsSWEET1b  were significantly upregulated, while  OsSWEET1a  and  OsSWEET13  were significantly downregulated in both  oss40-7  and  oss40-14  compared to WT. Furthermore, these five  OsS40  displayed strong transcriptional activation activity and were located in the nucleus. Most of the  OsS40  genes were downregulated in the  oss40-1, oss40-7 , and  oss40-12  mutants, but upregulated in the  oss40-13  and  oss40-14  mutants, indicating coordinated regulation among  OsS40  members. These results suggest that  OsS40-1, OsS40-7, OsS40-12, OsS40-13 , and  OsS40-14  are senescence-associated genes, involved in the senescence and carbon allocation network by modulating other  OsS40  members,  SWEET  member genes, and senescence-related gene expression.

Five OsS40 Family Members Are Identified as Senescence-Related Genes in Rice by Reverse Genetics Approach