OsFTIP1-Mediated Regulation of Florigen Transport in Rice Is Negatively Regulated by the Ubiquitin-Like Domain Kinase OsUbDK gamma 4

OsFTIP1, whose abundance is regulated by OsUbDK gamma 4 through the 26S proteasome pathway, affects rice flowering under long days (LDs) via modulating transport of the LD-specific florigen RFT1. Flowering time is a critical agronomic trait that determines successful seed production and adaptation of crop plants. Photoperiodic control of this process in flowering plants is mediated by the long-distance mobile signal called florigen partly encoded by FLOWERING LOCUS T (FT) in Arabidopsis thaliana and its orthologs in other plant species. Despite the progress in understanding FT transport in the dicot model Arabidopsis, the mechanisms of florigen transport in monocots, which provide most of the biomass in agriculture, are unknown. Here, we show that rice FT-INTERACTING PROTEIN1 (OsFTIP1), a member of the family of multiple C2 domain and transmembrane region proteins (MCTPs) and the closest ortholog of Arabidopsis FTIP1, is required for export of RICE FLOWERING LOCUS T 1 (RFT1) from companion cells to sieve elements. This affects RFT1 movement to the shoot apical meristem and its regulation of rice flowering time under long days. We further reveal that a ubiquitin-like domain kinase gamma 4, OsUbDK gamma 4, interacts with OsFTIP1 and modulates its degradation in leaves through the 26S proteasome, which in turn affects RFT1 transport to the shoot apical meristem. Thus, dynamic modulation of OsFTIP1 abundance in leaves by a negative regulator OsUbDK gamma 4 is integral to the role of OsFTIP1 in mediating RFT1 transport in rice and provides key evidence for a conserved role of FTIP1-like MCTPs in mediating florigen transport in flowering plants.