Intensification of Harmful Cyanobacterial Blooms in a Eutrophic, Temperate Lake Caused by Nitrogen, Temperature, and CO2.

Warmer temperatures can significantly increase the intensity of cyanobacterial harmful algal blooms (CHABs) in eutrophic freshwater ecosystems. However, few studies have examined the effects of CO2 2 enrichment in tandem with elevated temperature and/or nutrients on cyanobacterial taxa in freshwater ecosystems. Here, we observed changes in the biomass of cyanobacteria, nutrients, pH, and carbonate chemistry over a two-year period in a shallow, eutrophic freshwater lake and performed experiments to examine the effects and co-effects of CO2, 2 , temperature, and nutrient enrichment on cyanobacterial and N2 2-fixing (diazotrophic) communities assessed via high throughput sequencing of the 16S rRNA and nifH genes, respectively. During both years, there were significant CHABs (50-500 mu g cyanobacterial chlorophyll-a L- 1 ) and lake CO2 2 levels were undersaturated (<= 300 mu atm p CO 2 ). NH4+ 4 + significantly increased the net growth rates of cyanobacteria as well as the biomass of the diazotrophic cyanobacterial order Nostocales under elevated and ambient CO2 2 conditions. In a fall experiment, the N2 2 fixation rates of Nostocales were significantly higher when populations were enriched with CO2 2 and P, relative to CO2-enriched 2-enriched populations that were not amended with P. During a summer experiment, N2 2 fixation rates increased significantly under N and CO2 2- enriched conditions relative to N-enriched and ambient CO2 2 conditions. Nostocales dominated the diazotrophic communities of both experiments, achieving the highest relative abundance under CO2-enriched 2-enriched conditions when N was added in the first experiment and when CO2 2 and temperature were elevated in the second experiment, when N2 2 fixation rates also increased significantly. Collectively, this study indicates that N promotes cyanobacterial blooms including those formed by Dolichospermum and that the biomass and N2 2 fixation rates of diazotrophic cyanobacterial taxa may benefit from enhanced CO2 2 levels in eutrophic lakes.