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Abstract. We compare primary productivity estimates based on different photosynthetic ‘currencies’ (electrons, O2 and carbon) collected from the dynamic coastal upwelling waters of the California Current. Fast Repetition Rate Fluorometry and O2/N2 measurements were used to collect high-resolution underway estimates of photosynthetic electron transport rates and net community productivity, respectively, alongside on-station 14C uptake experiments to measure gross carbon fixation rates. Our survey captured two upwelling filaments at Cape Blanco and Cape Mendocino with distinct biogeochemical signatures and iron availabilities, enabling us to examine photosynthetic processes along a natural iron gradient. Significant differences in photo-physiology, cell sizes, Si:NO3- draw-down ratios, and molecular markers of Fe-stress indicated that phytoplankton assemblages near Cape Mendocino were Fe-stressed, while those near Cape Blanco were Fe-replete. Upwelling of O2-poor deep water to the surface complicated O2-based net community productivity estimates, but we were able to correct for these vertical mixing effects using continuous [N2O] surface measurements and depth-profiles of ∂[O2]∂[N2O]. Vertical mixing corrections were strongly correlated to sea surface temperature, which serves as an N2O-independent proxy for upwelling. Following vertical mixing corrections, all three productivity estimates reflected trends in Fe-stress physiology, indicating greater productivity near Cape Blanco compared to Cape Mendocino. For all assemblages, carbon fixation varied as a hyperbolic function of electron transport rates, but the derived parameters of this relationship were highly variable and significantly correlated with physiological indicators of Fe-stress (σPSII, FV/FM, Si:NO3- and diatom-specific PSI gene expression), suggesting that iron availability influenced the coupling between photosynthetic electron transport and subsequent carbon fixation. Net community productivity showed strong coherence with daily-integrated photosynthetic electron transport rates across the entire cruise track, with no apparent relationship with Fe-stress. This result suggests that fluorescence-based estimates of gross photochemistry are still a good indicator for bulk primary productivity, even if Fe-limitation influences the stoichiometric relationship between productivity currencies. 

Abstract 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 Eastern boundary upwelling currents are some of the most biologically productive and diverse regions in the world's oceans. Driven by equatorward winds and Ekman transport, surface waters are transported offshore and replaced by cold, nutrient‐rich deep waters that seed extensive phytoplankton blooms. Studying phytoplankton community succession and physiological acclimation during the initial stages of upwelling is critical to building a comprehensive understanding of phytoplankton responses to upwelling in these important regions. Additionally, factors like lateral transport, seed population dynamics and physiological and molecular shifts are conducive to shaping the community assemblage and primary productivity. This study examines how phytoplankton gene expression and resulting physiology change between early and later phases of upwelling. By incorporating metatranscriptomic analyses and stable isotope incubations to measure nutrient uptake kinetics into our assessment of early and later upwelling stages, we observed variability in phytoplankton assemblages and differential gene expression of phytoplankton that were de‐coupled from their physiology. We show that the gene expression response to a fresh upwelling event precedes their physiological response. Ultimately, understanding how phytoplankton change through the course of an upwelling event is critical to assessing their importance to regional biological rate processes, trophic systems and resulting biogeochemistry. 

Abstract 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. 

Abstract 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.