Oxygen Deficient Zones (ODZs) of the world’s oceans represent a relatively small fraction of the ocean by volume (<0.05% for suboxic and<5% for hypoxic) yet are receiving increased attention by experimentalists and modelers due to their importance in ocean nutrient cycling and predicted susceptibility to expansion and/or contraction forced by global warming. Conventional methods to study these biogeochemically important regions of the ocean have relied on well-developed but still relatively high cost and labor-intensive shipboard methods that include mass-spectrometric analysis of nitrogen-to-argon ratios (N2/Ar) and nutrient stoichiometry (relative abundance of nitrate, nitrite, and phosphate). Experimental studies of denitrification rates and processes typically involve either
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in-situ orin-vitro incubations using isotopically labeled nutrients. Over the last several years we have been developing a Gas Tension Device (GTD) to study ODZ denitrification including deployment in the largest ODZ, the Eastern Tropical North Pacific (ETNP). The GTD measures total dissolved gas pressure from which dissolved N2concentration is calculated. Data from two cruises passing through the core of the ETNP near 17 °N in late 2020 and 2021 are presented, with additional comparisons at 12 °N for GTDs mounted on a rosette/CTD as well as modified profiling Argo-style floats. Gas tension was measured on the float with an accuracy of< 0.1% and relatively low precision (< 0.12%) when shallow (P< 200 dbar) and high precision (< 0.03%) when deep (P > 300 dbar). We discriminate biologically produced N2(ie., denitrification) from N2in excess of saturation due to physical processes (e.g., mixing) using a new tracer – ‘preformed excess-N2’. We used inert dissolved argon (Ar) to help test the assumption that preformed excess-N2is indeed conservative. We used the shipboard measurements to quantify preformed excess-N2by cross-calibrating the gas tension method to the nutrient-deficit method. At 17 °N preformed excess-N2decreased from approximately 28 to 12 µmol/kg over σ0 =24–27 kg/m3with a resulting precision of ±1 µmol N2/kg; at 12 °N values were similar except in the potential density range of 25.7< σ0< 26.3 where they were lower by 1 µmol N2/kg due likely to being composed of different source waters. We then applied these results to gas tension and O2(< 3 µmol O2/kg) profiles measured by the nearby float to obtain the first autonomous biogenic N2profile in the open ocean with an RMSE of ± 0.78 µM N2, or ± 19%. We also assessed the potential of the method to measure denitrification rates directly from the accumulation of biogenic N2during the float drifts between profiling. The results suggest biogenic N2rates of ±20 nM N2/day could be detected over >16 days (positive rates would indicate denitrification processes whereas negative rates would indicate predominantly dilution by mixing). These new observations demonstrate the potential of the gas tension method to determine biogenic N2accurately and precisely in future studies of ODZs. -
Abstract Concentrations and elemental ratios of suspended particulate organic matter influence many biogeochemical processes in the ocean, including patterns of phytoplankton nutrient limitation and links between carbon, nitrogen and phosphorus cycles. Here we present direct measurements of cellular nutrient content and stoichiometric ratios for discrete phytoplankton populations spanning broad environmental conditions across several ocean basins. Median cellular carbon-to-phosphorus and nitrogen-to-phosphorus ratios were positively correlated with vertical nitrate-to-phosphate flux for all phytoplankton groups and were consistently higher for cyanobacteria than eukaryotes. Light and temperature were inconsistent predictors of stoichiometric ratios. Across nutrient-rich and phosphorus-stressed biomes in the North Atlantic, but not in the nitrogen-stressed tropical North Pacific, we find that a combination of taxonomic composition and environmental acclimation best predict bulk particulate organic matter composition. Our findings demonstrate the central role of plankton biodiversity and plasticity in controlling linkages between ocean nutrient and carbon cycles in some regions.
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Abstract Anaerobic ammonium oxidation (anammox) contributes substantially to ocean nitrogen loss, particularly in anoxic marine zones (AMZs). Ammonium is scarce in AMZs, raising the hypothesis that organic nitrogen compounds may be ammonium sources for anammox. Biochemical measurements suggest that the organic compounds urea and cyanate can support anammox in AMZs. However, it is unclear if anammox bacteria degrade these compounds to ammonium themselves, or rely on other organisms for this process. Genes for urea degradation have not been found in anammox bacteria, and genomic evidence for cyanate use for anammox is limited to a cyanase gene recovered from the sediment bacterium Candidatus Scalindua profunda. Here, analysis of Ca. Scalindua single amplified genomes from the Eastern Tropical North Pacific AMZ revealed genes for urea degradation and transport, as well as for cyanate degradation. Urease and cyanase genes were transcribed, along with anammox genes, in the AMZ core where anammox rates peaked. Homologs of these genes were also detected in meta-omic datasets from major AMZs in the Eastern Tropical South Pacific and Arabian Sea. These results suggest that anammox bacteria from different ocean regions can directly access organic nitrogen substrates. Future studies should assess if and under what environmental conditions these substrates contribute to the ammonium budget for anammox.