β
Figure 2. SO Signaling. Blue arrows represent EX1 signaling events. Prior to SO stimulation, EX1 is present in the grana margins in complex with several other proteins, but SO accumulation causes EX1 to disassociate as a result of oxidative modification. EX1 can then 1) translocate to the nucleus (dashed arrow) to promote SORG expression in concert with WRKY transcription factors; or 2) undergo proteolysis by FtsH2. Proteolysis also promotes SORG expression through an as-yet-unknown mechanism. Orange arrows represent carotenoid signaling. SO generated in the grana core oxidizes β-carotene, yielding β-cyclocitral (β-CC). This signaling molecule promotes increased expression of MBS1 and SCL14, which in turn upregulate genes for stress adaptation. β-CC or some other unknown signal promotes expression of the OXI1 kinase, which mediates cell death in response to SO from the grana core. SAFEGAURD1 (SAFE1) also acts independently of EX1 to suppress responses to SO caused by damage to the grana margins. This figure was created with Biorender, and was inspired in part by figures from (Woodson, 2019; Wang et al., 2020; Li et al., 2023).