Macroalgal blooms favor heterotrophic diazotrophic bacteria in nitrogen-rich and phosphorus-limited coastal surface waters in the Yellow Sea

Macroalgal blooms may lead to dramatic changes in physicochemical variables and biogeochemical cycling in affected waters. However, little is known about the effects of macroalgal blooms on marine bacteria, especially those functioning in nutrient cycles. We measured environmental factors and investigated bacterial diazotrophs in two niches, surface waters that were covered (CC) and non-covered (CF) with massive macroalgal canopies of Ulva prolifera, in the Yellow Sea in the summer of 2011 using real-time PCR and clone library analysis of nifH genes. We found that heterotrophic diazotrophs (Gammaproteobacteria) dominated the communities and were mostly represented by Vibrio-related phylotypes in both CC and CF. Desulfovibrio-related phylotypes were only detected in CC. There were significant differences in community composition in these two environments (p < 0.001) and a much higher abundance of nifH in CC (4.55 × 10⁶ copies l^-1) than in CF (2.49 × 10⁶ copies l^-1). The nifH copy number was inversely related to concentrations of ammonium and dissolved inorganic nitrogen and to the stoichiometric ratios of N:P and N:Si. This indicates that macroalgal blooms significantly affect diazotrophic abundance and community composition and that vibrios and Desulfovibrio-related heterotrophic diazotrophs adapt well to the (N-rich but P-limited) environment during blooming. Potential ecological and microbiological mechanisms behind this scenario are discussed.