Description: Nitrogen fixation, the biological reduction of dinitrogen gas (N2) to ammonium (NH4+), is quantitatively the most important external source of new nitrogen (N) to the open ocean. Classically, the ecological niche of oceanic N2 fixers (diazotrophs) is ascribed to tropical oligotrophic surface waters, often depleted in fixed N, with a diazotrophic community dominated by cyanobacteria. Although this applies for large areas of the ocean, biogeochemical models and phylogenetic studies suggest that the oceanic diazotrophic niche may be much broader than previously considered, resulting in major implications for the global N-budget. Here, we report on the composition, distribution and abundance of nifH, the functional gene marker for N2 fixation. Our results show the presence of eight clades of diazotrophs in the oxygen minimum zone (OMZ) off Peru. Although proteobacterial clades dominated overall, two clusters affiliated to spirochaeta and archaea were identified. N2 fixation was detected within OMZ waters and was stimulated by the addition of organic carbon sources supporting the view that non-phototrophic diazotrophs were actively fixing dinitrogen. The observed co-occurrence of key functional genes for N2 fixation, nitrification, anammox and denitrification suggests that a close spatial coupling of N-input and N-loss processes exists in the OMZ off Peru. The wide distribution of diazotrophs throughout the water column adds to the emerging view that the habitat of marine diazotrophs can be extended to low oxygen/high nitrate areas. Furthermore, our statistical analysis suggests that NO2- and PO43- are the major factors affecting diazotrophic distribution throughout the OMZ. In view of the predicted increase in ocean deoxygenation resulting from global warming, our findings indicate that the importance of OMZs as niches for N2 fixation may increase in the futur

Description: Nitrogen fixation, the biological reduction of dinitrogen gas (N2) to ammonium (NH4+), is quantitatively the most important external source of new nitrogen (N) to the open ocean. Classically, the ecological niche of oceanic N2 fixers (diazotrophs) is ascribed to tropical oligotrophic surface waters, often depleted in fixed N, with a diazotrophic community dominated by cyanobacteria. Although this applies for large areas of the ocean, biogeochemical models and phylogenetic studies suggest that the oceanic diazotrophic niche may be much broader than previously considered, resulting in major implications for the global N-budget. Here, we report on the composition, distribution and abundance of nifH, the functional gene marker for N2 fixation. Our results show the presence of eight clades of diazotrophs in the oxygen minimum zone (OMZ) off Peru. Although proteobacterial clades dominated overall, two clusters affiliated to spirochaeta and archaea were identified. N2 fixation was detected within OMZ waters and was stimulated by the addition of organic carbon sources supporting the view that non-phototrophic diazotrophs were actively fixing dinitrogen. The observed co-occurrence of key functional genes for N2 fixation, nitrification, anammox and denitrification suggests that a close spatial coupling of N-input and N-loss processes exists in the OMZ off Peru. The wide distribution of diazotrophs throughout the water column adds to the emerging view that the habitat of marine diazotrophs can be extended to low oxygen/high nitrate areas. Furthermore, our statistical analysis suggests that NO2− and PO43− are the major factors affecting diazotrophic distribution throughout the OMZ. In view of the predicted increase in ocean deoxygenation resulting from global warming, our findings indicate that the importance of OMZs as niches for N2 fixation may increase in the futur

Description: In the oxygen minimum zones (OMZs) of the tropical oceans, sluggish ventilation combined with strong microbial respiration of sinking organic matter results in the depletion of oxygen (O2). When O2 concentrations drop below ~5 µmol/L, organic matter is generally assumed to be respired with nitrate, ultimately leading to the loss of fixed inorganic nitrogen via anammox and denitrification. However, direct measurements of microbial O2 consumption at low O2 levels are - apart from a single experiment conducted in the OMZ off Peru - so far lacking. At the same time, consistently observed active aerobic ammonium and nitrite oxidation at non-detectable O2 concentrations (〈1 µmol/L) in all major OMZs, suggests aerobic microorganisms, likely including heterotrophs, to be well adapted to near-anoxic conditions. Consequently, microaerobic (〈=5 µmol/L) remineralization of organic matter, and thus release of ammonium, in low- O2 environments might be significantly underestimated at present. Here we present extensive measurements of microbial O2 consumption in OMZ waters, combined with highly sensitive O2 (STOX) measurements and meta-omic functional gene analyses. Short-term incubation experiments with labelled O2 (18-18O2) carried out in the Namibian and Peruvian OMZ, revealed persistent aerobic microbial activity at depths with non-detectable concentrations of O2 (〈=50 nmol/L). In accordance, examination of metagenomes and metatranscriptomes from Chilean and Peruvian OMZ waters identified genes encoding for terminal respiratory oxidases with high O2 affinities as well as their expression by diverse microbial communities. Oxygen consumption was particularly enhanced near the upper OMZ boundaries and could mostly (~80%) be assigned to heterotrophic microbial activity. Compared to previously identified anaerobic microbial processes, microaerobic organic matter respiration was the dominant remineralization pathway and source of ammonium (~90%) in the upper Namibian and Peruvian OMZ. Our results reconcile so-far existing mismatches between ammonium sources and sinks in OMZs, and may help to improve biogeochemical modelling of the effects of future ocean de-oxygenation on aerobic and anaerobic organic matter remineralization in these zones.

Description: In the oxygen minimum zones (OMZs) of the tropical oceans, sluggish ventilation combined with strong microbial respiration of sinking organic matter results in the depletion of oxygen (O2). When O2 concentrations drop below ~5 µmol/L, organic matter is generally assumed to be respired with nitrate, ultimately leading to the loss of fixed inorganic nitrogen via anammox and denitrification. However, direct measurements of microbial O2 consumption at low O2 levels are - apart from a single experiment conducted in the OMZ off Peru - so far lacking. At the same time, consistently observed active aerobic ammonium and nitrite oxidation at non-detectable O2 concentrations (〈1 µmol/L) in all major OMZs, suggests aerobic microorganisms, likely including heterotrophs, to be well adapted to near-anoxic conditions. Consequently, microaerobic (〈=5 µmol/L) remineralization of organic matter, and thus release of ammonium, in low- O2 environments might be significantly underestimated at present. Here we present extensive measurements of microbial O2 consumption in OMZ waters, combined with highly sensitive O2 (STOX) measurements and meta-omic functional gene analyses. Short-term incubation experiments with labelled O2 (18-18O2) carried out in the Namibian and Peruvian OMZ, revealed persistent aerobic microbial activity at depths with non-detectable concentrations of O2 (〈=50 nmol/L). In accordance, examination of metagenomes and metatranscriptomes from Chilean and Peruvian OMZ waters identified genes encoding for terminal respiratory oxidases with high O2 affinities as well as their expression by diverse microbial communities. Oxygen consumption was particularly enhanced near the upper OMZ boundaries and could mostly (~80%) be assigned to heterotrophic microbial activity. Compared to previously identified anaerobic microbial processes, microaerobic organic matter respiration was the dominant remineralization pathway and source of ammonium (~90%) in the upper Namibian and Peruvian OMZ. Our results reconcile so-far existing mismatches between ammonium sources and sinks in OMZs, and may help to improve biogeochemical modelling of the effects of future ocean de-oxygenation on aerobic and anaerobic organic matter remineralization in these zones.