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Coastal upwelling of nutrients and metals along eastern boundary currents fuels some of the most biologically productive marine ecosystems. Although iron is a main driver of productivity in many of these regions, iron cycling and acquisition by microbes remain poorly constrained, in part due to the unknown composition of organic ligands that keep bioavailable iron in solution. In this study, we investigated organic ligand composition in discrete water samples collected across the highly productive California Coastal upwelling system. Siderophores were observed in distinct nutrient regimes at concentrations ranging from 1 pM to 18 pM. Near the shallow continental shelf, ferrioxamine B was observed in recently upwelled, high chlorophyll surface waters while synechobactins were identified within nepheloid layers at 60–90 m depth. In offshore waters characterized by intermediate chlorophyll, iron, and nitrate concentrations, we found amphibactins and an unknown siderophore with a molecular formula of C33H58O8N5Fe. Highest concentrations were measured in the photic zone, however, amphibactins were also found in waters as deep as 1500 m. The distribution of siderophores provides evidence for microbial iron deficiency across a range of nutrient regimes and indicates siderophore production and acquisition is an important strategy for biological iron uptake in iron limited coastal systems. Polydisperse humic ligands were also detected throughout the water column and were particularly abundant near the benthic boundary. Our results highlight the fine‐scale spatial heterogeneity of metal ligand composition in an upwelling environment and elucidate distinct sources that include biological production and the degradation of organic matter in suboxic waters. [ABSTRACT FROM AUTHOR]

Summary: Frequent blooms of phytoplankton occur in coastal upwelling zones creating hotspots of biological productivity in the ocean. As cold, nutrient‐rich water is brought up to sunlit layers from depth, phytoplankton are also transported upwards to seed surface blooms that are often dominated by diatoms. The physiological response of phytoplankton to this process, commonly referred to as shift‐up, is characterized by increases in nitrate assimilation and rapid growth rates. To examine the molecular underpinnings behind this phenomenon, metatranscriptomics was applied to a simulated upwelling experiment using natural phytoplankton communities from the California Upwelling Zone. An increase in diatom growth following 5 days of incubation was attributed to the genera Chaetoceros and Pseudo‐nitzschia. Here, we show that certain bloom‐forming diatoms exhibit a distinct transcriptional response that coordinates shift‐up where diatoms exhibited the greatest transcriptional change following upwelling; however, comparison of co‐expressed genes exposed overrepresentation of distinct sets within each of the dominant phytoplankton groups. The analysis revealed that diatoms frontload genes involved in nitrogen assimilation likely in order to outcompete other groups for available nitrogen during upwelling events. We speculate that the evolutionary success of diatoms may be due, in part, to this proactive response to frequently encountered changes in their environment. [ABSTRACT FROM AUTHOR]

Phytoplankton community structure in the eastern Equatorial Pacific (EEP) and the Peru upwelling system was determined using ChemTax analysis of pigment data. Photosynthetic efficiencies (Fv : Fm), concentrations of macronutrients, iron (Fe), and the redox proteins flavodoxin (Flv) and ferredoxin (Fd) were used to assess the physiological status of the algal communities with respect to Fe. Diel periodicity in Fv :Fm ratios was observed in Fe-stressed populations of Prochlorococcus and type 4 haptophytes (Phaeocystis) in the EEP with daytime values (0.05-0.30) displaying a distinct noontime minimum. On the Peruvian continental shelf, upwelling of Fe-rich subsurface waters led to massive diatom blooms (reaching chlorophyll a [Chl a] values >45 µg L-1) with near-maximal photosynthetic efficiencies (Fv: Fm >0.6). Assimilation and luxurious uptake of Fe, scavenging processes, and Ekman transport off the shelf break led to rapid decreases in Fe concentration and dramatic shifts in phytoplankton assemblages. The shelf break region was dominated by cryptophyte populations with relatively high photosynthetic competency (Fv:Fm = 0.4-0.5). In this region, subnanomolar Fe concentrations limited further diatom growth. As Fe concentrations decreased offshore, haptophyte (coccolithophorids and Phaeocystis) populations contributed approximately 40-80% toward the total Chl a concentration. Haptophyte populations were Fe stressed, as determined by high Fly indices (Flv [Flv + Fd]-1) and low photosynthetic efficiencies (Fv:Fm ∼0.3). The Flv index was very high in regions where Fe concentrations were <0.2 nmol L-1, but it was 0 when Fe levels were >0.4 nmol L-1. In the Peru upwelling, Fe concentrations between 0.2 and 0.4 nmol L-1 represented a transitional region that marked the threshold switch in community Fd and Flv expression. [ABSTRACT FROM AUTHOR]

We compare the relative contribution of large phytoplankton (>5 and >10 µm) to uptake rates of carbon (C), nitrate (NO3-), and dissolved silicon (Si) and uptake ratios of Si:NO3- and Si:C in Monterey Bay, California (a high-iron region) with three low-iron regions in the eastern tropical Pacific: the Costa Rica upwelling dome, Humboldt current, and Peru upwelling. We also demonstrate the effect of iron enrichment on the above parameters in the latter three regions. In Monterey Bay waters, the µ5-µm size fraction accounted on average for the majority of particulate organic C and was responsible for 92% of total C uptake, 81% of total NO3- uptake, and 98% of total Si uptake, In contrast, in low-iron eastern tropical Pacific waters, the >5-µm size fraction accounted for less than half of the particulate organic C and was responsible for a substantially smaller proportion of total C and NO3- uptake: 27-43% for C and 34-59% for NO3-. Iron enrichment experiments in the eastern tropical Pacific resulted in much higher NO3-, C, and Si uptake rates, although increases were restricted to cells in the >5-µm size fraction. Si:NO3- and Si:C uptake ratios decreased after iron addition at most locations, and decreases were a direct result of lower Si : NO3- and Si : C uptake ratios in the >5-µm size fraction. Our results suggest that iron availability is a major factor regulating primary production, new production, Si uptake, and Si:NO3- and Si:C uptake ratios in the larger phytoplankton size classes in high-nitrate, low-chlorophyll (HNLC) regions in the eastern tropical Pacific. [ABSTRACT FROM AUTHOR]

Presents a study which demonstrated the photochemical reactivity of siderophores produced by marine heterotrophic bacteria and cyanobacteria based on characteristic ferric ion complexes. Role of siderophores on the oceanic biogeochemical cycling of iron and the regulation of marine primary productivity; Materials and methodology of the study; Discussion on the results of the study.

Organic dissolved iron (dFe)-binding ligands were measured by competitive ligand exchange-adsorptive cathodic stripping voltammetry (CLE-ACSV) at multiple analytical windows (side reaction coefficient of salicylaldoxime, and 100) in surface and benthic boundary layer (BBL) samples along the central California coast during spring and summer. The weakest ligands were detected in the BBL at the lowest analytical window with average log in the summer and 10.8 ± 0.2 in the spring. Between 3% and 18% of the dFe complexation in the BBL was accounted for by HS, which were measured separately in samples by ACSV and may indicate a source of dFe-binding ligands from San Francisco Bay. The strongest ligands were found in nearshore spring surface waters at the highest analytical window with average log, and the concentrations of these ligands declined rapidly offshore. The ligand pools in the surface and BBL waters were distinct from each other based on principal components analysis, with variances in the BBL ligand pool explained by sample location, and variance in surface waters explained by water mass. The use of multiple analytical window analysis elucidated several distinct iron-binding ligand pools, each with unique distributions in the central California Current system. [ABSTRACT FROM AUTHOR]

In the northern Gulf of Alaska, mesoscale, anticyclonic eddies have been implicated as a mechanism of the cross-shelf exchange of iron (Fe)-replete coastal waters with Fe-deplete subarctic Alaskan gyre waters. Based on existing hydrography and macronutrient distributions, a Sitka eddy sampled during August 2007 is divided into a surface eddy core, a shallow subsurface eddy core, and a deeper subsurface eddy core. The distributions of aluminum (Al), manganese (Mn), and Fe in the eddy are examined and compared to distributions at shelf stations similar to where the eddy formed as well as basin stations representative of the Fe-limited subarctic Alaskan gyre. Relative to basin stations, dissolved and particulate Al and dissolved Mn were elevated in eddy core waters. Reactive Fe concentrations within the shallow subsurface eddy core were nearly seven times greater than reactive Fe concentrations at similar densities at the basin stations. This shallow subsurface eddy core is likely mixed into the euphotic zone during storm-induced mixing as well as mixing during isopycnal relaxation as an eddy dies out. The flux of reactive Fe to surface waters from the shallow subsurface core of eddies in the Gulf of Alaska is a significant source compared to both remote and local atmospheric dust deposition. It is hypothesized that Fe supply from eddies in the eastern Gulf of Alaska shifts the core of the high-nutrient, lower-than-expected chlorophyll water farther west in conjunction with an "eddy desert" in that same region. [ABSTRACT FROM AUTHOR]

The physicochemical speciation of dissolved iron (Fe) across natural dissolved Fe gradients in the oceanic and shelf domains of the southeastern Bering Sea was examined in surface and subsurface samples using competitive ligand exchange-adsorptive cathodic stripping voltammetry with the added ligand salicylaldoxime. Two ligand classes were measured in all samples, a stronger L1 ligand class and a weaker L2 ligand class. Conditional stability constants for both ligand classes were comparable between surface and subsurface samples, with mean log Kcond FeL1,Fe0 = 11.5 ± 0.3 and mean log KcondFeL2,Fe0 = 10.3 ± 0.3 in surface samples, and mean log KcondFeL1,Fe0 = 11.4 ± 0.2 with a weaker ligand and mean log KcondFeL2,Fe0 of 10.2 ± 0.2 in subsurface samples. The concentrations of dissolved Fe were strongly correlated with ambient stronger L1 ligand concentrations for all samples with dissolved Fe concentrations greater than 0.2 nmol L-1. In samples with dissolved Fe concentrations less than 0.2 nmol L-1, large and variable excesses of L1 ligand concentrations were measured, coincident with observed Fe stress or limitation on the ambient phytoplankton. These observations suggest that the phytoplankton community is readily able to access dissolved Fe from the FeL1 complex, resulting in excess L1 in these waters. The available speciation data from other sources indicate that a significant correlation exists between dissolved Fe and L1 ligand concentrations in samples with intermediate dissolved Fe, and this is a seemingly ubiquitous feature of dissolved Fe cycling in the marine environment. [ABSTRACT FROM AUTHOR]