Phosphorus limitation and diel control of nitrogen-fixing cyanobacteria in the Baltic Sea

摘要

Up to half of the annual new nitrogen inputs into the Baltic Sea originate from blooms of N_2-fixing cyanobacteria. Estimates of the magnitude of this new nitrogen vary, partially because relatively few studies have investigated short-term changes in N_2-fixation rates in response to environmental changes in situ, including phosphorus availability, one of the major factors limiting N_2 fixation in the system. We examined cyanobacterial N_2 fixation in response to phosphorus amendments over the diel cycle during 2002 and 2003 in the Baltic Sea, when both Nodularia spumigena and Aphanizomenon sp. formed blooms. Phosphorus stimulated N_2 fixation in the open-sea areas in the Northern Baltic Proper and Gulf of Finland during both years. In microcosm experiments, both chlorophyll a concentration and N_2 fixation were positively related to time (R~2 = 0.79 and 0.54, respectively) for at least 4.5 d after the P amendment. N_2 fixation was enhanced up to 3-fold within 4.5 d by a single P pulse. N_2 fixation continued in the dark at 16 to 61 % of maximum rates during the day, and there were no consistent changes in nitrogenase enzyme abundance in response to darkness. Immunoblotting showed that N_2 fixation is not regulated in response to darkness by size modifications of the Fe and MoFe proteins in N. spumigena or of the Fe protein of Aphanizomenon sp. Capability to fix N_2 at high rates at night allows these cyanobacteria to maximize their utilization of periodic P pulses for subsequent growth.