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Because nitrogen availability limits primary production over much of the global ocean, understanding the controls on the marine nitrogen inventory and supply to the surface ocean is essential for understanding biological productivity and exchange of greenhouse gases with the atmosphere. Quantifying the ocean’s inputs, outputs, and internal cycling of nitrogen requires a variety of tools and approaches, including measurements of the nitrogen isotope ratio in organic and inorganic nitrogen species. The marine nitrogen cycle, which shapes nitrogen availability and speciation in the ocean, is linked to the elemental cycles of carbon, phosphorus, and trace elements. For example, the majority of nitrogen cycle oxidation and reduction reactions are mediated by enzymes that require trace metals for catalysis. Recent observations made through global-scale programs such as GEOTRACES have greatly expanded our knowledge of the marine nitrogen cycle. Though much work remains to be done, here we outline key advances in understanding the marine nitrogen cycle that have been achieved through these analyses, such as the distributions and rates of dinitrogen fixation, terrestrial nitrogen inputs, and nitrogen loss processes. 

Major coastal upwelling systems are among the most productive marine ecosystems in the world. They contribute disproportionately to the cycling of carbon and nutrients in the ocean and influence marine biogeochemistry beyond their productive regions. Characterized by intense microbial respiration (both aerobic and anaerobic), major coastal upwelling systems are also hotspots for the production and outgassing of potent greenhouse gases (GHG) such as CO2, N2O, and CH4. Quantifying and understanding these roles in the context of a changing climate is therefore a subject of great interest. Here we provide a short synthesis of the current knowledge of the contributions of major coastal upwelling systems to the cycling of GHG. Despite variations within and among different systems, low-latitude coastal upwelling systems typically act as a net carbon source to the atmosphere, while those at higher latitudes function as weak sinks or remain neutral regarding atmospheric CO2. These systems also significantly contribute to oceanic N2O and CH4 emissions, although the extent of their contribution to the latter remains poorly constrained. We also overview recent and future changes to upwelling systems in the context of a warmer climate and discuss uncertainties and implications for GHG production. Although rapid coastal warming is anticipated in all major coastal upwelling systems, the future changes in upwelling-favorable winds and their implications within the context of increased stratification are uncertain. Finally, we examine the major challenges that impede our ability to accurately predict how major coastal upwelling systems will respond to future climate change, and present recommendations for future research to better capture ongoing changes and disentangle natural and forced variability. 

Abstract Dissolved organic phosphorus (DOP) concentration distributions in the global surface ocean inform our understanding of marine biogeochemical processes such as nitrogen fixation and primary production. The spatial distribution of DOP concentrations in the surface ocean reflect production by primary producers and consumption as an organic nutrient by phytoplankton including diazotrophs and other microbes, as well as other loss processes such as photolysis. Compared to dissolved organic carbon and nitrogen, however, relatively few marine DOP concentration measurements have been made, largely due to the lack of automated analysis techniques. Here we present a database of marine DOP concentration measurements (DOPv2021) that includes new (n = 730) and previously published (n = 3140) observations made over the last ~30 years (1990–2021), including 1751 observations in the upper 50 m. This dataset encompasses observations from all major ocean basins including the poorly represented Indian, South Pacific, and Southern Oceans and provides insight into spatial distributions of DOP in the ocean. It is also valuable for researchers who work on marine primary production and nitrogen fixation. 

Abstract. Across the Southern Ocean in winter, nitrification is the dominantmixed-layer nitrogen cycle process, with some of the nitrate producedtherefrom persisting to fuel productivity during the subsequent growingseason. Because this nitrate constitutes a regenerated rather than a newnutrient source to phytoplankton, it will not support the net removal ofatmospheric CO2. To better understand the controls on Southern Oceannitrification, we conducted nitrite oxidation kinetics experiments insurface waters across the western Indian sector in winter. While allexperiments (seven in total) yielded a Michaelis–Menten relationship withsubstrate concentration, the nitrite oxidation rates only increasedsubstantially once the nitrite concentration exceeded 115±2.3 to245±18 nM, suggesting that nitrite-oxidizing bacteria (NOB) require aminimum (i.e., “threshold”) nitrite concentration to produce nitrate. Thehalf-saturation constant for nitrite oxidation ranged from 134±8 to403±24 nM, indicating a relatively high affinity of Southern OceanNOB for nitrite, in contrast to results from culture experiments. Despitethe high affinity of NOB for nitrite, its concentration rarely declinesbelow 150 nM in the Southern Ocean's mixed layer, regardless of season. Inthe 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 fornitrification in the winter Southern Ocean. The decoupling of ammonium andnitrite oxidation, combined with a possible nitrite concentration thresholdfor NOB, may explain the non-zero nitrite that persists throughout theSouthern Ocean's mixed layer year-round. Additionally, nitrite oxidation maybe limited by dissolved iron, the availability of which is low across theupper Southern Ocean. Our findings have implications for understanding thecontrols on nitrification and ammonium and nitrite distributions, both inthe Southern Ocean and elsewhere. 

Abstract. As a key biogeochemical pathway in the marine nitrogen cycle, nitrification (ammonia oxidation and nitrite oxidation) converts the most reduced form of nitrogen – ammonium–ammonia (NH4+–NH3) – into the oxidized species nitrite (NO2-) and nitrate (NO3-). In the ocean, these processes are mainly performed by ammonia-oxidizing archaea (AOA) and bacteria (AOB) and nitrite-oxidizing bacteria (NOB). By transforming nitrogen speciation and providing substrates for nitrogen removal, nitrification affects microbial community structure; marine productivity (including chemoautotrophic carbon fixation); and the production of a powerful greenhouse gas, nitrous oxide (N2O). Nitrification is hypothesized to be regulated by temperature, oxygen, light, substrate concentration, substrate flux, pH and other environmental factors. Although the number of field observations from various oceanic regions has increased considerably over the last few decades, a global synthesis is lacking, and understanding how environmental factors control nitrification remains elusive. Therefore, we have compiled a database of nitrification rates and nitrifier abundance in the global ocean from published literature and unpublished datasets. This database includes 2393 and 1006 measurements of ammonia oxidation and nitrite oxidation rates and 2242 and 631 quantifications of ammonia oxidizers and nitrite oxidizers, respectively. This community effort confirms and enhances our understanding of the spatial distribution of nitrification and nitrifiers and their corresponding drivers such as the important role of substrate concentration in controlling nitrification rates and nitrifier abundance. Some conundrums are also revealed, including the inconsistent observations of light limitation and high rates of nitrite oxidation reported from anoxic waters. This database can be used to constrain the distribution of marine nitrification, to evaluate and improve biogeochemical models of nitrification, and to quantify the impact of nitrification on ecosystem functions like marine productivity and N2O production. This database additionally sets a baseline for comparison with future observations and guides future exploration (e.g., measurements in the poorly sampled regions such as the Indian Ocean and method comparison and/or standardization). The database is publicly available at the Zenodo repository: https://doi.org/10.5281/zenodo.8355912 (Tang et al., 2023). 

Abstract Biological dinitrogen fixation is the major source of new nitrogen to marine systems and thus essential to the ocean’s biological pump. Constraining the distribution and global rate of dinitrogen fixation has proven challenging owing largely to uncertainty surrounding the controls thereon. Existing South Atlantic dinitrogen fixation rate estimates vary five-fold, with models attributing most dinitrogen fixation to the western basin. From hydrographic properties and nitrate isotope ratios, we show that the Angola Gyre in the eastern tropical South Atlantic supports the fixation of 1.4–5.4 Tg N.a⁻¹, 28-108% of the existing (highly uncertain) estimates for the basin. Our observations contradict model diagnoses, revealing a substantial input of newly-fixed nitrogen to the tropical eastern basin and no dinitrogen fixation west of 7.5˚W. We propose that dinitrogen fixation in the South Atlantic occurs in hotspots controlled by the overlapping biogeography of excess phosphorus relative to nitrogen and bioavailable iron from margin sediments. Similar conditions may promote dinitrogen fixation in analogous ocean regions. Our analysis suggests that local iron availability causes the phosphorus-driven coupling of oceanic dinitrogen fixation to nitrogen loss to vary on a regional basis. 

Abstract Ocean circulation supplies the surface ocean with the nutrients that fuel global ocean productivity. However, the mechanisms and rates of water and nutrient transport from the deep ocean to the upper ocean are poorly known. Here, we use the nitrogen isotopic composition of nitrate to place observational constraints on nutrient transport from the Southern Ocean surface into the global pycnocline (roughly the upper 1.2 km), as opposed to directly from the deep ocean. We estimate that 62 ± 5% of the pycnocline nitrate and phosphate originate from the Southern Ocean. Mixing, as opposed to advection, accounts for most of the gross nutrient input to the pycnocline. However, in net, mixing carries nutrients away from the pycnocline. Despite the quantitative dominance of mixing in the gross nutrient transport, the nutrient richness of the pycnocline relies on the large-scale advective flow, through which nutrient-rich water is converted to nutrient-poor surface water that eventually flows to the North Atlantic.