MAPKKK28 functions upstream of the MKK1‐MPK1 cascade to regulate abscisic acid responses in rice

The mitogen-activated protein kinase (MAPK) cascade (MAPKKK-MAPKK-MAPK) plays a critical role in biotic and abiotic stress responses and abscisic acid (ABA) signalling. A previous study has shown that the ABA-activated MKK1-MPK1 cascade is essential in regulating ABA response and stress tolerance in rice. However, the specific MAPKKK upstream of the MKK1-MPK1 cascade in ABA signalling remains unknown. Here, we identified that MAPKKK28, a previously uncharacterized member of the rice MEKK family, is involved in regulating ABA responses, including seed germination, root growth, stomatal closure, and the tolerance to oxidative stress and osmotic stress. We found that MAPKKK28 directly interacts with and phosphorylates MKK1. Further analysis indicated that the activation of both MKK1 and MPK1 depends on MAPKKK28 in ABA signalling. Genetic analysis revealed that MAPKKK28 functions upstream of the MKK1-MPK1 cascade to positively regulate ABA responses and enhance tolerance to oxidative and osmotic stress. These results not only reveal a new complete MAPK cascade in plants but also uncover its importance in ABA signalling.