OsBSK3 and OsBSK2 regulate grain size and leaf angle via MAPK signaling pathway in rice

Grain size and leaf angle are key agronomic traits that determine the final yield. OsBSKs (BRASSINOSTEROID-SIGNALING KINASES) and OsMAPKs (MITOGEN ACTIVATED PROTEIN KINASE) are known to play essential roles in plant growth, development, and stress responses. However, the potential crosstalk between these pathways and their specific roles in regulating grain size and leaf angle remain largely unexplored in rice. Here, we characterized that OsBSKs regulate grain size and leaf angle in rice, and among these, OsBSK2 and OsBSK3 may play more critical roles. The grain size and leaf angle in osbsk3 and osbsk2 mutants are significantly smaller, whereas the OsBSK3-overexpressing lines (OsBSK3-OEs) exhibit considerably larger grain size and leaf angle compared to the others. Furthermore, both OsBSK3 and OsBSK2 interact with OsMKKK10, indirectly activating OsMAPK6 in plant cells. Notably, mutations in MAPK cascade components, such as smg2-1 (an osmkkk10 mutant), smg1-1 (an osmkk4 mutant), and dsg1 (an osmapk6 mutant), resulted in significantly reduced leaf angles. Moreover, these mutations were able to rescue the increased grain size and leaf angle observed in OsBSK3 overexpression lines. Additionally, we also identified OsWRKY53 as a potential downstream target of the OsBSKs-OsMKKK10-OsMKK4-OsMAPK6 cascade in the regulation of grain size and leaf angle. Taken together, the above results not only highlight the essential and specific roles of OsBSK3 and OsBSK2 in regulating rice grain size and leaf angle, but also reveal the mechanism which OsBSK3/OsBSK2 mediate MAPK cascade to regulate rice grain size and leaf angle.