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Coastal upwelling currents such as the California Current System (CCS) comprise some of the most productive biological systems on the planet. Diatoms dominate these upwelling events in part due to their rapid response to nutrient entrainment. In this region, they may also be limited by the micronutrient iron (Fe), an important trace element primarily involved in photosynthesis and nitrogen assimilation. The mechanisms behind how diatoms physiologically acclimate to the different stages of the upwelling conveyor belt cycle remain largely uncharacterized. Here, we explore their physiological and metatranscriptomic response to the upwelling cycle with respect to the Fe limitation mosaic that exists in the CCS. Subsurface, natural plankton assemblages that would potentially seed surface blooms were examined over wide and narrow shelf regions. The initial biomass and physiological state of the phytoplankton community had a large impact on the overall response to simulated upwelling. Following on‐deck incubations under varying Fe physiological states, our results suggest that diatoms quickly dominated the blooms by "frontloading" nitrogen assimilation genes prior to upwelling. However, diatoms subjected to induced Fe limitation exhibited reductions in carbon and nitrogen uptake and decreasing biomass accumulation. Simultaneously, they exhibited a distinct gene expression response which included increased expression of Fe‐starvation induced proteins and decreased expression of nitrogen assimilation and photosynthesis genes. These findings may have significant implications for upwelling events in future oceans, where changes in ocean conditions are projected to amplify the gradient of Fe limitation in coastal upwelling regions. [ABSTRACT FROM AUTHOR]

The California Current System is a diatom‐dominated region characterized by seasonal coastal upwelling and additional elevated mesoscale activity. Cyclonic mesoscale eddies in the region trap productive coastal waters with their planktonic communities and transport them offshore with limited interaction with surrounding waters, effectively acting as natural mesocosms, where phytoplankton populations undergo ecological succession as eddies age. This study examines diatom community composition within two mesoscale cyclonic eddies that formed in the same region of the California Current System 2 months apart and in the California Current waters surrounding them. The diatom communities were analyzed in the context of shifting environmental gradients and through a lens of community succession to expand our understanding of biophysical interactions in California Current System cyclonic eddies. Diatom communities within each eddy were different from non‐eddy communities and varied in concert with salinity and dissolved iron (Fe) concentrations. The younger, nearshore eddy displayed higher macronutrient and dissolved Fe concentrations, had higher values for diatom Shannon diversity and evenness, and had nutrient ratios indicative of either eventual silicic acid (Si) or Fe limitation or possibly co‐limitation. The older, offshore eddy displayed low macronutrient and dissolved Fe concentrations, was likely nitrate‐limited, and had lower diatom Shannon diversity and evenness indices. Sequences from the genus Rhizosolenia, some of which form vertically migrating mats to bypass nitrate limitation, dominated in the older eddy. This is of potential significance as the prevalence of Rhizosolenia mats could impact estimates of carbon cycling and export in the wider California coastal area. [ABSTRACT FROM AUTHOR]

Atmospheric deposition represents a major input for micronutrient trace elements (TEs) to the surface ocean and is often quantified indirectly through measurements of aerosol TE concentrations. Sea spray aerosol (SSA) dominates aerosol mass concentration over much of the global ocean, but few studies have assessed its contribution to aerosol TE loading, which could result in overestimates of "new" TE inputs. Low‐mineral aerosol concentrations measured during the U.S. GEOTRACES Pacific Meridional Transect (GP15; 152°W, 56°N to 20°S), along with concurrent towfish sampling of surface seawater, provided an opportunity to investigate this aspect of TE biogeochemical cycling. Central Pacific Ocean surface seawater Al, V, Mn, Fe, Co, Ni, Cu, Zn, and Pb concentrations were combined with aerosol Na data to calculate a "recycled" SSA contribution to aerosol TE loading. Only vanadium was calculated to have a SSA contribution averaging >1% along the transect (mean of 1.5%). We derive scaling factors from previous studies on TE enrichments in the sea surface microlayer and in freshly produced SSA to assess the broader potential for SSA contributions to aerosol TE loading. Maximum applied scaling factors suggest that SSA could contribute significantly to the aerosol loading of some elements (notably V, Cu, and Pb), while for others (e.g., Fe and Al), SSA contributions largely remained <1%. Our study highlights that a lack of focused measurements of TEs in SSA limits our ability to quantify this component of marine aerosol loading and the associated potential for overestimating new TE inputs from atmospheric deposition. Plain Language Summary: Sea spray aerosol (SSA) generated by breaking waves and bursting bubbles can dominate the particle population in the marine atmosphere. We investigated whether this mechanism could mobilize significant amounts of trace elements (TEs)—nutrient and pollutant elements present in the surface ocean in very low concentrations. Previous studies have suggested that TEs could become enriched on SSA during its formation, relative to their concentrations in seawater, and this could make the process a more significant source for aerosol concentrations of some elements. But for others, the SSA contribution is always likely to be insignificant. Our calculations based on sea salt and TE aerosol concentrations in the atmosphere and dissolved TEs in the surface ocean suggest that SSA does not contribute significantly to total aerosol TE loading, with the possible exception of vanadium, which has relatively higher concentrations in seawater than the other elements considered. This study highlights that this process remains poorly quantified due to limited direct measurements. Key Points: Sea spray aerosol (SSA) concentration was quantified along a meridional Pacific Ocean transect during low mineral aerosol conditionsSea spray‐derived aerosol trace elements (TEs) are negligible unless significant enrichment occurs during formation; only vanadium is >1%SSA contributions to aerosol TEs of >100% calculated by applying enrichments from earlier studies suggests these are not widely applicable [ABSTRACT FROM AUTHOR]

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]

Diatoms are prominent eukaryotic phytoplankton despite being limited by the micronutrient iron in vast expanses of the ocean. As iron inputs are often sporadic, diatoms have evolved mechanisms such as the ability to store iron that enable them to bloom when iron is resupplied and then persist when low iron levels are reinstated. Two iron storage mechanisms have been previously described: the protein ferritin and vacuolar storage. To investigate the ecological role of these mechanisms among diatoms, iron addition and removal incubations were conducted using natural phytoplankton communities from varying iron environments. We show that among the predominant diatoms, Pseudo-nitzschia were favored by iron removal and displayed unique ferritin expression consistent with a long-term storage function. Meanwhile, Chaetoceros and Thalassiosira gene expression aligned with vacuolar storage mechanisms. Pseudo-nitzschia also showed exceptionally high iron storage under steady-state high and low iron conditions, as well as following iron resupply to iron-limited cells. We propose that bloom-forming diatoms use different iron storage mechanisms and that ferritin utilization may provide an advantage in areas of prolonged iron limitation with pulsed iron inputs. As iron distributions and availability change, this speculated ferritin-linked advantage may result in shifts in diatom community composition that can alter marine ecosystems and biogeochemical cycles. [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]

Abstract: The metabolic activity and growth of phytoplankton taxa drives their ecological function and contribution to biogeochemical processes. We present the first quantitative, taxon‐resolved silica production rates, growth rates, and silica content estimates for co‐occurring diatoms along two cross‐shelf transects off the California coast using the fluorescent tracer 2‐(4‐pyridyl)‐5‐((4‐(2‐dimethylaminoethylaminocarbamoyl)methoxy)phenyl)oxazole, and confocal microscopy. Taxon contribution to total diatom community silica production was predominantly a function of the surface area (SA) of new frustule that each taxon created as opposed to cell abundance or frustule thickness. The influential role of SA made large diatoms disproportionately important to community silica production over short time scales (< 1 d). In some cases, large taxa that comprised only ∼ 15% of numerical cell abundance accounted for over 50% of total community silica production. Over longer time scales relevant to bloom dynamics, the importance of SA declines and growth rate becomes the dominant influence on contribution to production. The relative importance of SA and growth rate in relation to silica production was modeled as the time needed for a smaller, faster‐growing taxon to create more SA than a larger, slower‐growing taxon. Differences in growth rate between the taxa effected the model outcome more than differences in SA. Shifts in relative silica production among taxa are time restricted by finite resources that limit the duration of a bloom. These patterns offer clues as to how taxa respond to their environment and the consequences for both species succession and the potential diatom contribution to elemental cycling. [ABSTRACT FROM AUTHOR]

Abstract: A 3‐day shipboard incubation experiment was conducted in the California Current System in July 2014 to investigate the cycling of iron (Fe), copper (Cu), nickel (Ni), cadmium (Cd), manganese (Mn), cobalt (Co), lead (Pb), and scandium (Sc) under a range of light and particle conditions. Filtered (< 0.2 μm) and unfiltered treatments were incubated under the following light conditions: Dark, light (“UV”), and light without the ultraviolet (UV) wavelengths (“noUV”). The experiment was sampled for carbon and Fe uptake rates, dissolved trace metal concentrations (Fe, Cu, Ni, Cd, Mn, Co, Pb, Sc), Fe and Cu speciation, size‐fractionated concentrations of Cd and Fe, and diatom community composition from DNA sequencing. Exposure to UV light increased phytoplankton Fe uptake in the first 24 h of the incubation relative to the noUV treatment, suggesting that a fraction of the ambient ligand‐bound Fe was photoreactive. Fe‐binding organic ligand production was observed in the unfiltered light treatments in association with increasing chlorophyll a, and evidence for Cu‐binding ligand production in these treatments was also observed. Biological uptake of Cd and Co was observed along with scavenging of dissolved Pb. Manganese appeared to be rapidly oxidized by Mn‐oxidizing bacteria with concomitant drawdown of dissolved Ni. Scandium displayed similar trends to Fe, reinforcing the limited observations of the physicochemical similarities between these two elements in seawater. Overall, this study highlights distinct impacts of photochemical processes, scavenging, and biological effects on marine trace metal cycling in an environment characterized by seasonal upwelling. [ABSTRACT FROM AUTHOR]

Little is known about marine mineral deposits in the Arctic Ocean, an ocean dominated by continental shelf and basins semi-closed to deep-water circulation. Here, we present data for ferromanganese crusts and nodules collected from the Amerasia Arctic Ocean in 2008, 2009, and 2012 (HLY0805, HLY0905, and HLY1202). We determined mineral and chemical compositions of the crusts and nodules and the onset of their formation. Water column samples from the GEOTRACES program were analyzed for dissolved and particulate scandium concentrations, an element uniquely enriched in these deposits. The Arctic crusts and nodules are characterized by unique mineral and chemical compositions with atypically high growth rates, detrital contents, Fe/Mn ratios, and low Si/Al ratios, compared to deposits found elsewhere. High detritus reflects erosion of submarine outcrops and North America and Siberia cratons, transport by rivers and glaciers to the sea, and distribution by sea ice, brines, and currents. Uniquely high Fe/Mn ratios are attributed to expansive continental shelves, where diagenetic cycling releases Fe to bottom waters, and density flows transport shelf bottom water to the open Arctic Ocean. Low Mn contents reflect the lack of a mid-water oxygen minimum zone that would act as a reservoir for dissolved Mn. The potential host phases and sources for elements with uniquely high contents are discussed with an emphasis on scandium. Scandium sorption onto Fe oxyhydroxides and Sc-rich detritus account for atypically high scandium contents. The opening of Fram Strait in the Miocene and ventilation of the deep basins initiated Fe-Mn crust growth ∼15 Myr ago. [ABSTRACT FROM AUTHOR]