HHL1 and SOQ1 synergistically regulate nonphotochemical quenching in Arabidopsis

Non-photochemical quenching (NPQ) is an important photo-protective mechanism that quickly dissipates excess light energy as heat. NPQ can be induced in a few seconds to several hours; most studies of this process have focused on the rapid induction of NPQ. Recently, a new, slowly induced form of NPQ, called qH, was found during the discovery of the quenching inhibitor SUPPRESSOR OF QUENCHING 1 (SOQ1). However, the specific mechanism of qH remains unclear. Here, we found that HYPERSENSITIVE TO HIGH LIGHT 1 (HHL1)-a damage repair factor of photosystem II (PSII)-interacts with SOQ1. The enhanced NPQ phenotype of the hhl1 mutant is similar to that of the soq1 mutant, which is not related to energy-dependent quenching or other known NPQ components. Furthermore, the hhl1 soq1 double mutant showed higher NPQ than the single mutants, but its pigment content and composition were similar to those of the wild type. Overexpressing HHL1 decreased NPQ in hhl1 to below wild-type levels, whereas NPQ in hhl1 plants overexpressing SOQ1 was lower than that in hhl1 but higher than that in the wild type. Moreover, we found that HHL1 promotes the SOQ1-mediated inhibition of plastidial lipoprotein (LCNP) through its VWA domain. We propose that HHL1 and SOQ1 synergistically regulate NPQ.