Chuanyou Li / Institute of Genetics and Developmental Biology, Chinese Academy of Sciences
Stem cell specification and organ patterning in multicellular organisms relies on the precise spatiotemporal control of RNA polymerase II (Pol II)-dependent gene transcription, in which the evolutionarily conserved Mediator co-activator complex plays an essential role. The GRAS family proteins SHR and SCR orchestrate a transcriptional program that determines the fate and asymmetric cell divisions (ACDs) of stem cells generating the root ground tissue. The mechanism by which SHR-SCR relays context-specific regulatory signals to the Pol II general transcription machinery is unknown. Here we report the role of Mediator in controlling the spatiotemporal transcriptional output of SHR-SCR during stem cell ACD and ground tissue patterning. The Mediator subunit MED31 interacted with SCR but not SHR. Reduction of MED31 disrupted the spatiotemporal activation of CYCD6;1, a cell-cycle gene, leading to defective ACD of stem cells generating ground tissue. MED31 was recruited to the promoter of CYCD6;1 in a SCR-dependent manner, and MED31 and SHR competed each other for binding to SCR in a dose-dependent manner. We show that MED31 is involved in the formation of a MED31-SCR-SHR ternary complex through the interface protein SCR. We demonstrate that the relative protein abundance of MED31 and SHR in different cell types regulates the dynamic formation of the MED31-SCR-SHR ternary complex, which provides a tunable switch to strictly control the spatiotemporal transcriptional output.