UCYN-A/haptophyte symbioses dominate N2 fixation in the Southern California Current System

Abstract

The availability of fixed nitrogen (N) is an important factor limiting biological productivity in the oceans. In coastal waters, high dissolved inorganic N concentrations were historically thought to inhibit dinitrogen (N₂) fixation, however, recent N₂ fixation measurements and the presence of the N₂-fixing UCYN-A/haptophyte symbiosis in nearshore waters challenge this paradigm. We characterized the contribution of UCYN-A symbioses to nearshore N₂ fixation in the Southern California Current System (SCCS) by measuring bulk community and single-cell N₂ fixation rates, as well as diazotroph community composition and abundance. UCYN-A1 and UCYN-A2 symbioses dominated diazotroph communities throughout the region during upwelling and oceanic seasons. Bulk N₂ fixation was detected in most surface samples, with rates up to 23.0 ± 3.8 nmol N l⁻¹ d⁻¹, and was often detected at the deep chlorophyll maximum in the presence of nitrate (>1 µM). UCYN-A2 symbiosis N₂ fixation rates were higher (151.1 ± 112.7 fmol N cell⁻¹ d⁻¹) than the UCYN-A1 symbiosis (6.6 ± 8.8 fmol N cell⁻¹ d⁻¹). N₂ fixation by the UCYN-A1 symbiosis accounted for a majority of the measured bulk rates at two offshore stations, while the UCYN-A2 symbiosis was an important contributor in three nearshore stations. This report of active UCYN-A symbioses and broad mesoscale distribution patterns establishes UCYN-A symbioses as the dominant diazotrophs in the SCCS, where heterocyst-forming and unicellular cyanobacteria are less prevalent, and provides evidence that the two dominant UCYN-A sublineages are separate ecotypes.