The X‐ray structure of photosystem II reveals a novel electron transport pathway between P680, cytochrome b559 and the energy‐quenching cation, ChlZ+

When water oxidation by photosystem II (PSII) is impaired, an oxidized chlorophyll (Chl(Z)(+)) is formed that quenches excitation and may prevent photodamage. Both the identification of this Chl(+) and the mechanism of its oxidation and reduction are controversial. Using the available X-ray structures of PSII we calculated the efficiency of two proposed quenchers, Chl(Z)(+)(D1) and Chl(Z)(+)(D2). Of these two, only Chl(Z)(+)(D1) can quench to the degree observed experimentally. We also identify a chain of closely spaced pigments in the structure from Thermosynechococcus vulcanus that we propose to form a novel electron transport pathway between Chl(Z)(D1), beta-carotene, P680(+) and cytochrome b(559).