While there is a consensus that the cytochrome b6f complex (cytb6f) in algae and plants is involved in the regulatory mechanism of oxygenic photosynthesis known as light-induced state transitions (STs), no such consensus exists for cyanobacteria. Here, we provide the first direct functional evidence for cytb6f using single-point mutation data. We introduced a PetD-Phe124Ala substitution in the cyanobacterium Synechocystis sp. PCC 6803 to test the key predictions of the hydrophobic-mismatch (HMM) model for cytb6f-driven STs in all oxygenic photosynthetic species. These predictions concern the role of the Phe/Tyr124fg-loop-PetD and the extent and kinetic characteristics of STs. The effects of PetD-F124A mutation on STs were monitored using 77K and Pulse-Amplitude-Modulated (PAM) fluorescence. For comparison, we employed a phycobilisome (PBS)-less Synechocystis mutant and wild-type (WT) strain, as well as the stn7 mutant and WT of Arabidopsis plant. The PetD-F124A mutation reduced the extent of STs and selectively affected the two-exponential kinetics components of the transitions. Under State 1 conditions, the mutant exhibited ~60% less energetic decoupling of PBS from photosystem I (PSI) compared to the WT. It is explainable by the HMM model with the inability of the PetD-F124A mutant, during the induction phase of the State 2→State 1 transition to adopt the cytb6f conformation with minimal hydrophobic thickness. PAM-derived parameters indicated that PSII electron transport function is not inhibited, and no detectable effect on cyclic electron transport around PSI was observed under low-light conditions. Circular dichroism and differential scanning calorimetry confirmed that both the PSI trimer/monomer ratio and the structural integrity of the PBSs are preserved in the mutant. The compensatory response to the mutation includes decreased PSI content and an increase in PBS rod size. In conclusion, (1) cytb6f is involved in cyanobacterial STs; (2) evidence is provided supporting the HMM model; (3) the electron transfer and signal transduction functions of cytb6f are separated into distinct domains; and (4) the signaling pathway regulating STs and pigment-protein composition in Synechocystis involves PetD-Phe124.

Department of Agriculture Forest and Forest Products Sciences Plant Metabolic Physiology Kyushu University Fukuoka 812 8581 Japan

HCEMM BRC Pharmacodynamic Drug Interaction Research Group Hungarian Centre of Excellence for Molecular Medicine H 6728 Szeged Hungary

Institute of Biophysics and Biomedical Engineering Bulgarian Academy of Sciences 1113 Sofia Bulgaria

Institute of Photonics and Electronics of the Czech Academy of Sciences 18200 Prague Czech Republic

Institute of Plant Biology HUN REN Biological Research Centre H 6726 Szeged Hungary

Synthetic and Systems Biology Unit Institute of Biochemistry HUN REN Biological Research Centre H 6726 Szeged Hungary

Temasek Life Sciences Laboratory National University of Singapore Singapore 117604 Singapore

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