The water-water cycle is a major electron sink in Camellia species when CO2 assimilation is restricted.

The water-water cycle (WWC) is thought to dissipate excess excitation energy and balance the ATP/NADPH energy budget under some conditions. However, the importance of the WWC in photosynthetic regulation remains controversy. We observed that three Camellia cultivars exhibited high rates of photosynthetic electron flow under high light when photosynthesis was restricted. We thus tested the hypothesis that the WWC is a major electron sink in the three Camellia cultivars when CO2 assimilation is restricted. Light response curves indicated that the WWC was strongly increased with photorespiration and was positively correlated with extra ATP supplied from other flexible mechanisms excluding linear electron flow, implying that the WWC is an important alternative electron sink to balance ATP/NADPH energy demand for sustaining photorespiration in Camellia cultivars. Interestingly, when photosynthesis was depressed by the decreases in stomatal and mesophyll conductance, the rates of photosynthetic electron flow through photosystem II declined slightly and the rates of WWC was enhanced. Furthermore, the increased electron flow of WWC was positively correlated with the ratio of Rubisco oxygenation to carboxylation, supporting the involvement of alternative electron flow in balancing the ATP/NADPH energy budget. We propose that the WWC is a crucial electron sink to regulate ATP/NADPH energy budget and dissipate excess energy excitation in Camellia species when CO2 assimilation is restricted.