Abstract
Understanding the mechanistic basis of balanced excitation energy distribution between photosystem II and photosystem I (PSII and PSI) requires detailed investigation of the thylakoid light-harvesting system composed of energetically connected LHCII trimers. The exact mechanisms controlling the excitation energy distribution remain elusive, but reversible phosphorylation is known to be one important component. Here, we addressed the role of grana margins in regulation of excitation energy distribution, as these thylakoid domains host all the complexes of photosynthetic light reactions with dynamic response to environmental cues. First, the effect of detergents for the thylakoid membrane connectivity is explained. We show that a specific interaction between the separate LHCII trimers as well as between the LHCII trimers and the PSII and PSI-LHCI complexes is a prerequisite for energetically connected and functional thylakoid membrane. Second, we demonstrate that the optimization of light reactions under changing light conditions takes place in energetically connected LHCII lake and is attained by lateral rearrangements of the PSII-LHCII and PSI-LHCI-LHCII complexes depending especially on the phosphorylation status of the LHCII protein isoform LHCB2.