Phosphorylation of the transcription factor OsNAC29 by OsMAPK3 activates diterpenoid genes to promote rice immunity

Well-conserved mitogen-activated protein kinase (MAPK) cascades are essential for orchestrating of a wide range of cellular processes in plants, including defense responses against pathogen attack. NAC transcription factors play important roles in plant immunity, but their targets and how they are regulated remain largely unknown. Here, we identified the transcription factor OsNAC29 as a key component of a MAPK signaling pathway involved in rice (Oryza sativa) disease resistance. OsNAC29 binds directly to CACGTG motifs in the promoters of OsTPS28 and OsCYP71Z2, which are crucial for the biosynthesis of the phytoalexin 5,10-diketo-casbene and consequently rice blast resistance. OsNAC29 positively regulates rice blast resistance by promoting the expression of of OsTPS28 and OsCYP71Z2, and the function of OsNAC29 is genetically dependent on OsCYP71Z2 and OsTPS28. Furthermore, OsNAC29 interacts with OsRACK1A and OsMAPK3/6 to form an immune complex; OsMAPK3 phosphorylates OsNAC29 at Thr304 to prevent its proteasome-mediated degradation and promote its function against rice blast fungus. Phosphorylation of OsNAC29 at Thr304 is induced upon Magnaporthe oryzae infection and chitin treatment. Our data demonstrate the positive role of the OsMAPK3-OsNAC29-OsTPS28/OsCYP71Z2 module in rice blast resistance, providing insights into the molecular regulatory network and fine-tuning of NAC transcription factors in rice immunity.