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The availability of nitrogen (N) in ocean surface waters affects rates of photosynthesis and marine ecosystem structure. In spite of low dissolved inorganic N concentrations, export production in oligotrophic waters is comparable to more nutrient replete regions. Prior observations raise the possibility that di-nitrogen (N2) fixation supplies a significant fraction of N supporting export production in the Gulf of Mexico. In this study, geochemical tools were used to quantify the relative and absolute importance of both subsurface nitrate and N2 fixation as sources of new N fueling export production in the oligotrophic Gulf of Mexico in May 2017 and May 2018. Comparing the isotopic composition (“δ15N”) of nitrate with the δ15N of sinking particulate N collected during five sediment trap deployments each lasting two to four days indicates that N2 fixation is typically not detected and that the majority (≥80%) of export production is supported by subsurface nitrate. Moreover, no gradients in upper ocean dissolved organic N and suspended particulate N concentration and/or δ15N were found that would indicate significant N2 fixation fluxes accumulated in these pools, consistent with low Trichodesmium spp. abundance. Finally, comparing the δ15N of sinking particulate N captured within vs. below the euphotic zone indicates that during late spring regenerated N is low in δ15N compared to sinking N.

In contrast to its productive coastal margins, the open-ocean Gulf of Mexico (GoM) is notable for highly stratified surface waters with extremely low nutrient and chlorophyll concentrations. Field campaigns in 2017 and 2018 identified low rates of turbulent mixing, which combined with oligotrophic nutrient conditions, give very low estimates for diffusive flux of nitrate into the euphotic zone (< 1 µmol N m−2 d−1). Estimates of local N2-fixation are similarly low. In comparison, measured export rates of sinking particulate organic nitrogen (PON) from the euphotic zone are 2 – 3 orders of magnitude higher (i.e. 462 – 1144 µmol N m−2 d−1). We reconcile these disparate findings with regional scale dynamics inferred independently from remote-sensing products and a regional biogeochemical model and find that laterally-sourced organic matter is sufficient to support >90% of open-ocean nitrogen export in the GoM. Results show that lateral transport needs to be closely considered in studies of biogeochemical balances, particularly for basins enclosed by productive coasts., The middle of the Gulf of Mexico is stratified and highly oligotrophic, yet there are anomalously high fluxes of sinking particulate matter from the euphotic zone. Here the authors show that lateral advection of organic matter supports nitrogen export in the Gulf of Mexico’s open ocean.

Biomass and composition of the phytoplankton community were investigated in the deep-water Gulf of Mexico (GoM) at the edges of Loop Current anticyclonic eddies during May 2017 and May 2018. Using flow cytometry, high-performance liquid chromatography pigments and microscopy, we found euphotic zone integrated chlorophyll a of ~10 mg m−2 and autotrophic carbon ranging from 463 to 1268 mg m−2, dominated by picoplankton (

The highly stratified, oligotrophic regions of the oceans are predominantly nitrogen limited in the surface ocean and light limited at the deep chlorophyll maximum (DCM). Hence, determining light and nitrogen co-limitation patterns for diverse phytoplankton taxa is crucial to understanding marine primary production throughout the euphotic zone. During two cruises in the deep-water Gulf of Mexico, we measured primary productivity (H13CO3−), nitrate uptake (15NO3−), and ammonium uptake (15NH4+) throughout the water column. Primary productivity declined with depth from the mixed-layer to the DCM, averaging 27.1 mmol C m−2 d−1. The fraction of growth supported by NO3− was consistently low, with upper euphotic zone values ranging from 0.01 to 0.14 and lower euphotic zone values ranging from 0.03 to 0.44. Nitrate uptake showed strong diel patterns (maximum during the day), while ammonium uptake exhibited no diel variability. To parameterize taxon-specific phytoplankton nutrient and light utilization, we used a data assimilation approach (Bayesian Markov Chain Monte Carlo) including primary productivity, nutrient uptake, and taxon-specific growth rate measurements. Parameters derived from this analysis define distinct niches for five phytoplankton taxa (Prochlorococcus, Synechococcus, diatoms, dinoflagellates, and prymnesiophytes) and may be useful for constraining biogeochemical models of oligotrophic open-ocean systems.

Dinitrogen (N-2) fixation is an important source of biologically reactive nitrogen (N) to the global ocean. The magnitude of this flux, however, remains uncertain, in part because N-2 fixation rates have been estimated following divergent protocols and because associated levels of uncertainty are seldom reported-confounding comparison and extrapolation of rate measurements. A growing number of reports of relatively low but potentially significant rates of N-2 fixation in regions such as oxygen minimum zones, the mesopelagic water column of the tropical and subtropical oceans, and polar waters further highlights the need for standardized methodological protocols for measurements of N-2 fixation rates and for calculations of detection limits and propagated error terms. To this end, we examine current protocols of the N-15(2) tracer method used for estimating diazotrophic rates, present results of experiments testing the validity of specific practices, and describe established metrics for reporting detection limits. We put forth a set of recommendations for best practices to estimate N-2 fixation rates using N-15(2) tracer, with the goal of fostering transparency in reporting sources of uncertainty in estimates, and to render N-2 fixation rate estimates intercomparable among studies.

Ocean greening off Hawai'i From June to August 2018, the eruption of Kīlauea volcano triggered a diatom-dominated phytoplankton bloom. Wilson et al. set sail to sample the plume, deploying subsea gliders and using satellite monitoring to measure the dynamics of this rare event in the nutrient-poor Pacific (see the Perspective by Ducklow and Plank). They found subsurface chlorophyll maxima not visible by remote sensing, performed transcriptome and N isotope marker analysis, and measured nutrients, partitioning of biomass into different organisms, and primary production. Much of the data are corroborated by physical modeling of the ocean dynamics. The authors conclude that the plume was fed by the lava heating subsurface water and triggering upwelling of deepwater nutrients to the surface rather than by direct injection of micronutrients from lava. Science , this issue p. 1040 ; see also p. 978

Mixed layer (ML) gross (GOP) and net (NOP) oxygen production rates based on in situ mass balances of triple oxygen isotopes (TOI) and O2/Ar are influenced by vertical transport from below, a term traditionally difficult to constrain. Here, we present a new approach to estimate vertical eddy diffusivity (Kz) based on density gradients in the upper thermocline and wind-speed based rates of turbulent shear at the ML depth. As an example, we use this Kz, verified by an independent 7Be-based estimate, in an O2/TOI budget at a site in the oligotrophic South Pacific Gyre (SPG). NOP equaled 0.31 ± 0.16 mmol m-2 d-1 in the ML (~55-65 m depth) and 1.2 ± 0.4 mmol m-2 d-1 (80%) beneath the ML, while GOP equaled 74 ± 27 mmol m-2 d-1 (86%) in the ML and 12 ± 4 mmol m-2 d-1 (14%) below, revealing a vertical gradient in production rates unquantifiable without the Kz estimate.

Constraining the rates and spatial distribution of dinitrogen (N2) fixation fluxes to the ocean informs our understanding of the environmental sensitivities of N2 fixation as well as the timescale over which the fluxes of nitrogen (N) to and from the ocean may respond to each other. Here we quantify rates of N2 fixation as well as its contribution to export production along a zonal transect in the western tropical South Pacific (WTSP) Ocean using N isotope (“δ15N”) budgets. Comparing measurements of water column nitrate + nitrite δ15N with the δ15N of sinking particulate N at a western, central, and eastern station, these δ15N budgets indicate high, modest, and low rates of N2 fixation at the respective stations. The results also imply that N2 fixation supports exceptionally high, i.e. ≥ 50 %, of export production at the western and central stations, which are also proximal to the largest iron sources. These geochemically based rates of N2 fixation are equal to or greater than those previously reported in the tropical North Atlantic, indicating that the WTSP Ocean has the capacity to support globally significant rates of N2 fixation, which may compensate for N removal in the oxygen-deficient zones of the eastern tropical Pacific.

[1] The allochthonous supply of dissolved organic nitrogen (DON) from gyre margins into the interior of the ocean’s oligotrophic subtropical gyres potentially provides an important source of new N to gyre surface waters, thus sustaining export production. This process requires that a fraction of the transported DON be available to euphotic zone photoautotroph communities via mineralization. In this study, we investigated the biological and physical controls on the distribution and fate of DON within global ocean surface waters. Inputs of nitrate to the euphotic zone at upwelling zones fuel net accumulation of a DON pool that appears to resist rapid microbial remineralization, allowing subsequent advective transport into the subtropical gyres. Zonal gradients in DON concentrations across these gyres imply a DON sink in the surface layer. Assessment of the physical dynamics of gyre circulation and winter mixing revealed a pathway for DON removal from the mixed layer via vertical transport to the deep euphotic zone, which establishes the observed zonal gradients. Incubation experiments from the Florida Straits indicated surface-accumulated DON was largely resistant (over a few months) to utilization by the extant surface bacterioplankton community. In contrast, this same material was remineralized three times more rapidly when exposed to upper mesopelagic bacterioplankton. These results suggest the primary fate of surface DON to be removal via vertical mixing and subsequent mineralization below the mixed layer, implying a limited role for direct DON support of gyre export production from the surface layer. DON may contribute to export production at the eastern edges of the subtropical gyres, but only after its mineralization within the deep euphotic zone.

The concentration and 15N/14N ratio of total nitrogen (TN) were measured in precipitation samples collected at Bermuda between January and December 2000. By correcting for nitrate, analyzed previously, the concentration and δ15N of “reduced” N (RN, i.e., ammonium + organic N) were also determined. The TN precipitation flux (∼ 10–19 mmol N m− 2 yr− 1) is twice the NO3− precipitation flux, and the mass-weighted annual average δ15N of TN, − 2.3‰, is higher than the δ15N of NO3− in the same samples (− 4.5‰), indicating that RN has an annual average δ15N of − 0.6‰. While neither the concentration nor the flux of RN (6.8 µM and 5.2 mmol N m− 2 yr− 1, respectively) in precipitation shows statistically significant seasonal variation, the δ15N of RN varies significantly from − 2.7‰ in the cool season to 1.5‰ in the warm season. This seasonality in the δ15N of RN is similar to that of NO3−, implying that RN and NO3− in precipitation may have related sources or, more speculatively, mechanisms of inter-conversion. Additionally, the seasonality of the RN δ15N at Bermuda is similar to that of typical ammonium concentrations in precipitation at Bermuda, both showing maxima in the spring and late summer, raising the possibility that the maxima in the RN δ15N derives from ammonium at those times. Finally, the low δ15N of the TN flux will cause it to have an effect on the δ15N of Sargasso Sea thermocline NO3− that is in the same sense as the effect of N2 fixation, with slightly greater isotopic leverage. If the atmospheric TN flux is not marine-derived, it could explain a substantial fraction of the previously documented upward decrease in NO3− δ15N from deep water into the thermocline of the Sargasso Sea, for example, ∼ 20 to 35‰ of it, assuming a N2 fixation rate of 45 mmol N m-2 yr-1 as estimated by Hansell et al. [Hansell, D.A., Bates, N.R., and Olson, D.B., 2004. Excess nitrate and nitrogen fixation in the North Atlantic Ocean. Mar. Chem., 84:243–265.].

We compare the output of an 18-box geochemical model of the ocean with measurements to investigate the controls on both the mean values and variation of nitrate delta N-15 and delta O-18 in the ocean interior. The delta O-18 of nitrate is our focus because it has been explored less in previous work. Denitrification raises the delta N-15 and delta O-18 of mean ocean nitrate by equal amounts above their input values for N-2 fixation (for delta N-15) and nitrification (for delta O-18), generating parallel gradients in the delta N-15 and delta O-18 of deep ocean nitrate. Partial nitrate assimilation in the photic zone also causes equivalent increases in the delta N-15 and delta O-18 of the residual nitrate that can be transported into the interior. However, the regeneration and nitrification of sinking N can be said to decouple the N and O isotopes of deep ocean nitrate, especially when the sinking N is produced in a low latitude region, where nitrate consumption is effectively complete. The delta N-15 of the regenerated nitrate is equivalent to that originally consumed, whereas the regeneration replaces nitrate previously elevated in delta O-18 due to denitrification or nitrate assimilation with nitrate having the delta O-18 of nitrification. This lowers the delta O-18 of mean ocean nitrate and weakens nitrate delta O-18 gradients in the interior relative to those in delta N-15. This decoupling is characterized and quantified in the box model, and agreement with data shows its clear importance in the real ocean. At the same time, the model appears to generate overly strong gradients in both delta O-18 and delta N-15 within the ocean interior and a mean ocean nitrate delta O-18 that is higher than measured. This may be due to, in the model, too strong an impact of partial nitrate assimilation in the Southern Ocean on the delta N-15 and delta O-18 of preformed nitrate and/or too little cycling of intermediate-depth nitrate through the low latitude photic zone.

[1] N and O isotope analyses of water column nitrate between Bermuda and Puerto Rico document a bolus of low-δ15N nitrate throughout the Sargasso Sea thermocline, which we attribute primarily to the input of recently fixed N. Although previous work suggests southward increases in N2 fixation and ventilation age, no meridional trend in nitrate δ15N is apparent. In the upper 200 m, the algal uptake-driven increase in nitrate δ18O is greater than in δ15N, because of (1) a higher fraction of nitrate from N2 fixation at shallower depths and/or (2) cycling of N between nitrate assimilation and nitrification. A mean depth profile of newly fixed nitrate estimated from the nitrate isotope data is compared with results from an ocean circulation model forced with different Atlantic fields of N2 fixation. The nitrate from N2 fixation is communicated between the model's North and South Atlantic and suggests a whole Atlantic N2 fixation rate between 15 and 24 Tg N a−1. One important caveat is that fixed N in atmospheric deposition may contribute a significant proportion of the low-δ15N N in the Sargasso Sea thermocline, in which case the relatively low rate we estimate for N2 fixation would still be too high.

Some of our suppositions about the marine nitrogen cycle may be wrong. An innovative analysis of nutrients at the ocean's surface reveals a feedback mechanism that might hold the whole cycle in balance.