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Ulva prolifera , a typical green-tide-forming alga, can accumulate a large biomass in a relatively short time period, suggesting that photosynthesis in this organism, particularly its carbon fixation pathway, must be very efficient. Green algae are known to generally perform C 3  photosynthesis, but recent metabolic labeling and genome sequencing data suggest that they may also perform C 4  photosynthesis, so C 4  photosynthesis might be more wide-spread than previously anticipated. Both C 3  and C 4  photosynthesis genes were found in  U. prolifera  by transcriptome sequencing. We also discovered the key enzymes of C 4  metabolism based on functional analysis, such as pyruvate orthophosphate dikinase (PPDK), phosphoenolpyruvate carboxylase (PEPC), and phosphoenolpyruvate carboxykinase (PCK). To investigate whether the alga operates a C 4 -like pathway, the expression of  rbc L and PPDK and their enzyme activities were measured under various forms and intensities of stress (differing levels of salinity, light intensity, and temperature). The expression of  rbc L and PPDK and their enzyme activities were higher under adverse circumstances. However, under conditions of desiccation, the expression of  rbc L and ribulose-1, 5-biphosphate carboxylase (RuBPCase) activity was lower, whereas that of PPDK was higher. These results suggest that elevated PPDK activity may alter carbon metabolism and lead to a partial operation of C 4 -type carbon metabolism in  U. prolifera , probably contributing to its wide distribution and massive, repeated blooms in the Yellow Sea.

Evidence of Coexistence of C3 and C4 Photosynthetic Pathways in a Green-Tide-Forming Alga, Ulva prolifera