The Arabidopsis Mediator complex subunits MED14/SWP and MED16/SFR6/IEN1 differentially regulate defense gene expression in plant immune responses

Pathogen infection in plants triggers large-scale transcriptional changes, both locally and systemically. Emerging evidence suggests that the Arabidopsis Mediator complex plays a crucial role in these transcriptional changes. Mediator is highly conserved in eukaryotes, and its core comprises more than 20 subunits organized into three modules named head, middle and tail. The head and middle modules interact with general transcription factors and RNA polymeraseII, whereas the tail module associates with activators, and signals through the head and middle modules to the basal transcription machinery. In Arabidopsis, three tail module subunits, MED14, MED15 and MED16, have been identified. Both MED15 and MED16 have been implicated in plant immunity, but the role of MED14 has not been established. Here, we report the characterization of an Arabidopsis T-DNA insertion mutant of the MED14 gene. Similarly to the med15 and/or med16 mutations, the med14 mutation significantly suppresses salicylic acid-induced defense responses, alters transcriptional changes induced by the avirulent bacterial pathogen Pseudomonas syringae pv. tomato (Pst) DC3000/avrRpt2, and renders plants susceptible to both Pst DC3000/avrRpt2 and Pst DC3000. The med14 mutation also completely compromises biological induction of systemic acquired resistance (SAR), indicating that the tail module as a whole is essential for SAR. Interestingly, unlike the med16 mutation, which differentially affects expression of several SAR positive and negative regulators, med14 inhibits induction of a large group of defense genes, including both SAR positive and negative regulators, suggesting that individual subunits of the Mediator tail module employ distinct mechanisms to regulate plant immune responses.