We investigated interactions between photosynthesis and nitrogen fixation in the non-heterocystous marine cyanobacterium Trichodesmium IMS101 at the single-cell level by two-dimensional (imaging) microscopic measurements of chlorophyll fluorescence kinetics. Nitrogen fixation was closely associated with the appearance of cells with high basic fluorescence yield (F 0 ), termed bright cells. In cultures aerated with normal air, both nitrogen fixation and bright cells appeared in the middle of the light phase. In cultures aerated with 5% oxygen, both processes occurred at a low level throughout most of the day. Under 50% oxygen, nitrogen fixation commenced at the beginning of the light phase but declined soon afterwards. Rapid reversible switches between fluorescence levels were observed, which indicated that the elevated F 0 of the bright cells originates from reversible uncoupling of the photosystem II (PSII) antenna from the PSII reaction center. Two physiologically distinct types of bright cells were observed. Type I had about double F 0 compared to the normal F 0 in the dark phase and a PSII activity, measured as variable fluorescence (F v 5 F m 2 F 0 ), similar to normal non-diazotrophic cells. Correlation of type I cells with nitrogen fixation, oxygen concentration, and light suggests that this physiological state is connected to an up-regulation of the Mehler reaction, resulting in oxygen consumption despite functional PSII. Type II cells had more than three times the normal F 0 and hardly any PSII activity measurable by variable fluorescence. They did not occur under low-oxygen concentrations, but appeared under high-oxygen levels outside the diazotrophic period, suggesting that this state represents a reaction to oxidative stress not necessarily connected to nitrogen fixation. In addition to the two high-fluorescence states, cells were observed to reversibly enter a low-fluorescence state. This occurred mainly after a cell went through its bright phase and may represent a fluorescence-quenching recovery phase.Biological fixation of atmospheric nitrogen is performed by certain cyanobacteria when bioavailable forms of nitrogen (nitrate and ammonia) are limited. Nitrogen fixation is catalyzed by an essentially anaerobic enzyme, nitrogenase, which is irreversibly inhibited in vitro when exposed to molecular oxygen (for review, see Postgate, 1998). Diazotrophic cyanobacteria are the only nitrogen-fixing organisms that produce molecular oxygen as a by-product of photosynthesis and must prevent nitrogenase from being damaged by oxygenic photosynthesis (for review, see Gallon, 1992Gallon, , 2001Bergman et al., 1997;Berman-Frank et al., 2003). Most diazotrophic cyanobacteria achieve this by separating photosynthesis and nitrogen fixation either spatially, by differentiating highly specialized cells called heterocysts, or temporally, by fixing nitrogen at night (usually found in unicellular diazotrophic cyanobacteria). In contrast, filamentous non-heterocystous marine cyanobacteria of the genus Trichodesmium execut...