Control of nitrogen fixation by oxygen in purple nonsulfur bacteria

Some members of the facultatively phototrophic bacteria are able to grow diazotrophically in the presence of oxygen. As in other diazotrophs, the nitrogenase of the phototrophic bacteria is highly sensitive to oxygen; therefore, both the function and the expression of nitrogenase are strictly controlled by oxygen. This review focuses on the different levels of oxygen control in the two most extensively studied facultatively phototrophic bacteria, Rhodospirillum rubrum and Rhodobacter capsulatus. Current data show that oxygen controls nitrogen fixation at least at the levels of (1) transcription of nif genes, (2) the accumulation of the three different nitrogenase polypeptides, (3) the cellular activity of nitrogen fixation. In Rba. capsulatus, activation of the nifH promoter is the least oxygen-sensitive step, and nitrogen fixation is the most oxygen-sensitive step. ADP-Ribosylation of nitrogenase, occurring under conditions of ammonium-dependent inactivation of the enzyme, is not observed when Rba. capsulatus is exposed either suddenly or at a steady state to increased oxygen concentrations. Future research is required to understand the mechanisms of protection of nitrogenase against oxygen damage, and also the mechanisms by which oxygen controls the formation and activity of nitrogenase; this will add significantly to the biologically important question of how cells deal with the presence of toxic oxygen.