Controls on nitrite oxidation in the upper Southern Ocean: insights from winter kinetics experiments in the Indian sector

Across the Southern Ocean in winter, nitrification is the dominant mixed-layer nitrogen cycle process, with some of the nitrate produced therefrom persisting to fuel productivity during the subsequent growing season. Because this nitrate constitutes a regenerated rather than a new nutrient source to phytoplankton, it will not support the net removal of atmospheric CO2. To better understand the controls on Southern Ocean nitrification, we conducted nitrite oxidation kinetics experiments in surface waters across the western Indian sector in winter. While all experiments (seven in total) yielded a Michaelis-Menten relationship with substrate concentration, the nitrite oxidation rates only increased substantially once the nitrite concentration exceeded 115±2.3 to 245±18nM, suggesting that nitrite-oxidizing bacteria (NOB) require a minimum (i.e., "threshold") nitrite concentration to produce nitrate. The half-saturation constant for nitrite oxidation ranged from 134±8 to 403±24nM, indicating a relatively high affinity of Southern Ocean NOB for nitrite, in contrast to results from culture experiments. Despite the high affinity of NOB for nitrite, its concentration rarely declines below 150nM in the Southern Ocean's mixed layer, regardless of season. In the upper mixed layer, we measured ammonium oxidation rates that were two- to seven-fold higher than the coincident rates of nitrite oxidation, indicating that nitrite oxidation is the rate-limiting step for nitrification in the winter Southern Ocean. The decoupling of ammonium and nitrite oxidation, combined with a possible nitrite concentration threshold for NOB, may explain the non-zero nitrite that persists throughout the Southern Ocean's mixed layer year-round. Additionally, nitrite oxidation may be limited by dissolved iron, the availability of which is low across the upper Southern Ocean. Our findings have implications for understanding the controls on nitrification and ammonium and nitrite distributions, both in the Southern Ocean and elsewhere.