Nickel (Ni) is a micronutrient that plays a role in nitrogen uptake and fixation in the modern ocean may have impacted rates of methanogenesis on geological timescales. Here we present the results of a diagnostic model of global ocean Ni fluxes which addresses key questions about the biogeochemical processes which cycle Ni in the modern oceans. Our approach starts with extrapolating the sparse available observations of Ni data from the GEOTRACES project into a global gridded climatology of ocean Ni concentrations. Three different machine learning techniques were tested, each relying on marine tracers with better observational coverage such as macronutrient concentrations and physical parameters. The ocean transport of this global Ni concentration field is then estimated using the OCIM2 ocean circulation inverse model, revealing regions of net convergence or divergence. These diagnostics are not based on any assumption about Ni biogeochemical cycling, but their spatial patterns can be interpreted as reflecting biogeochemical processes. We find that the spatial pattern of Ni uptake in the surface ocean is similar to phosphate (P) uptake, but not silicate (Si) uptake, suggesting that Ni is not incorporated into diatom frustules. We find that Ni:P ratios at uptake do not decrease with Ni concentrations approaching 2 nM, which challenges the hypothesis of a ~2 nM pool of non-bioavailable Ni in the surface ocean. Finally, the net regeneration of Ni occurs deeper in the ocean than P remineralization, which could be explained by reversible scavenging or the presence of a refractory Ni phase.

We examined the biogeochemical impact of pairs of mesoscale cyclones and anticyclones in spatial proximity (<200 km apart) in the North Pacific Subtropical Gyre. While previous studies have demonstrated that upwelling associated with the intensification of cyclonic eddies supplies nutrients to the euphotic zone, we find that cyclonic eddies in their mature stage sustain plankton growth by increasing the diapycnal flux of nutrients to the lower portion of the euphotic zone. This increased supply results from enhanced vertical gradients in inorganic nutrients due to erosion of the nutricline that accompanied plankton growth during eddy intensification. From a biological standpoint, increased nutrient flux was linked with expansion of eukaryotic phytoplankton biomass and intensification of the deep chlorophyll maximum layer. This perturbation in the plankton community was associated with increased fluxes of biominerals (opal and calcium carbonate) and isotopically enriched nitrogen in particles exported in the cyclone. The time-integrated effects of thermocline uplifts and depressions were predictable deficits and surpluses of inorganic nutrients and dissolved oxygen in the lower euphotic zone. However, the stoichiometry of changes in oxygen and inorganic nutrients differed from that predicted for production and consumption of phytoplankton biomass, consistent with additional biological processes that decouple changes in oxygen and nutrient concentrations. The dynamics revealed by this study may be a common feature of oligotrophic ecosystems, where mesoscale biogeochemical perturbations are buffered by the deep chlorophyll maximum layer, which limits the ecological impact of eddies in the well-lit, near-surface ocean.

In stratified oligotrophic waters, phytoplankton communities forming the deep chlorophyll maximum (DCM) are isolated from atmospheric iron sources above and remineralized iron below. Reduced supply leads to a minimum in dissolved iron (dFe) near 100 m, but it is unclear if iron limits growth at the DCM. Here, we propose that natural iron addition events occur regularly with the passage of mesoscale eddies, which alter the supply of dFe and other nutrients relative to the supply of light, and can be used to test for iron limitation at the DCM. This framework is applied to two eddies sampled in the North Pacific Subtropical Gyre. Observations in an anticyclonic eddy center indicated downwelling of iron-rich surface waters, leading to increased dFe at the DCM but no increase in productivity. In contrast, uplift of isopycnals within a cyclonic eddy center increased supply of both nitrate and dFe to the DCM, and led to dominance of picoeukaryotic phytoplankton. Iron addition experiments did not increase productivity in either eddy, but did enhance leucine incorporation at ambient light in the cyclonic eddy, a potential indicator of iron stress among Prochlorococcus. Rapid cycling of siderophores and low dFe:nitrate uptake ratios also indicate that a portion of the microbial community was stressed by low iron. However, near-complete nitrate drawdown in this eddy, which represents an extreme case in nutrient supply compared to nearby Hawaii Ocean Time-series observations, suggests that recycling of dFe in oligotrophic ecosystems is sufficient to avoid iron limitation in the DCM under typical conditions.