Nitrification and ammonium dynamics in Taihu Lake, China: seasonal competition for ammonium between nitrifiers and cyanobacteria

Taihu Lake is hypereutrophic and experiences seasonal, cyanobacterial harmful algal blooms. These Microcystis blooms produce microcystin, a potent liver toxin, and are linked to anthropogenic nitrogen (N) and phosphorus (P) loads to lakes. Microcystis spp. cannot fix atmospheric N and must compete with ammonia-oxidizing and other organisms for ammonium (NH4+). We measured NH4+ regeneration and potential uptake rates and total nitrification using stable-isotope techniques. Nitrification studies included abundance of the functional gene for NH4+ 4 oxidation, amoA, for ammonia-oxidizing archaea (AOA) and bacteria (AOB). Potential NH4+ uptake rates ranged from 0.02 to 6.80 mu mol L-1 h(-1) in the light and from 0.05 to 3.33 mu mol L-1 h(-1) in the dark, and NH4+ 4 regeneration rates ranged from 0.03 to 2.37 mu mol L-1 h(-1). Nitrification rates exceeded previously reported rates in most freshwater systems. Total nitrification often exceeded 200 nmol L-1 d(-1) and was > 1000 nmol L-1 d(-1) at one station near a river discharge. AOA amoA gene copies were more abundant than AOB gene copies (p < 0.005) at all times; however, only abundance of AOB amoA (not AOA) was correlated with nitrification rates for all stations and all seasons (p < 0.005). Nitrification rates in Taihu Lake varied seasonally; at most stations, rates were highest in March, lower in June, and lowest in July, corresponding with cyanobacterial bloom progression, suggesting that nitrifiers were poor competitors for NH4+ during the bloom. Regeneration results suggested that cyanobacteria relied extensively on regenerated NH4+ to sustain the bloom. Internal NH4+ regeneration exceeded external N loading to the lake by a factor of 2 but was ultimately fueled by external N loads. Our results thus support the growing literature calling for watershed N loading reductions in concert with existing management of P loads.