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Abstract The availability of the micronutrient iron is important in regulating phytoplankton growth across much of the world’s oceans, particularly in the high-nutrient, low-chlorophyll regions. Compared to known mechanisms of iron acquisition and conservation in autotrophic protists (e.g. diatoms), those of dinoflagellates remain unclear, despite their frequent presence in offshore iron-limited waters. Here, we investigate the strategies of an ecologically important mixotrophic dinoflagellate to coping with low iron conditions. Coupled gene expression and physiological responses as a function of iron availability were examined in oceanic and coastal strains of the dinoflagellate Karlodinium. Under iron-replete conditions, grazing was only detected in coastal variants, resulting in faster growth rates compared to when grown autotrophically. Under iron-limited conditions, all isolates exhibited slower growth rates, reduced photosynthetic efficiencies, and lower cellular iron quotas than in iron-replete conditions. However, oceanic isolates exhibited higher relative growth rates compared to coastal isolates under similar low iron concentrations, suggesting they are better adapted to coping under iron limitation. Yet the oceanic isolates did not exhibit the ability to appreciably reduce cell volume or increase iron-use efficiencies compared to the coastal isolates to cope with iron limitation, as often observed in oceanic diatoms. Rather, molecular pathway analysis and corresponding gene expression patterns suggest that oceanic Karlodinium utilizes a high-affinity iron uptake system when iron is low. Our findings reveal cellular mechanisms by which dinoflagellates have adapted to low iron conditions, further shedding light on how they potentially survive in variable iron regions of the world’s oceans. 

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 The continental shelf of the Western Antarctic Peninsula (WAP) is a highly variable system characterized by strong cross-shelf gradients, rapid regional change, and large blooms of phytoplankton, notably diatoms. Rapid environmental changes coincide with shifts in plankton community composition and productivity, food web dynamics, and biogeochemistry. Despite the progress in identifying important environmental factors influencing plankton community composition in the WAP, the molecular basis for their survival in this oceanic region, as well as variations in species abundance, metabolism, and distribution, remains largely unresolved. Across a gradient of physicochemical parameters, we analyzed the metabolic profiles of phytoplankton as assessed through metatranscriptomic sequencing. Distinct phytoplankton communities and metabolisms closely mirrored the strong gradients in oceanographic parameters that existed from coastal to offshore regions. Diatoms were abundant in coastal, southern regions, where colder and fresher waters were conducive to a bloom of the centric diatom,Actinocyclus. Members of this genus invested heavily in growth and energy production; carbohydrate, amino acid, and nucleotide biosynthesis pathways; and coping with oxidative stress, resulting in uniquely expressed metabolic profiles compared to other diatoms. We observed strong molecular evidence for iron limitation in shelf and slope regions of the WAP, where diatoms in these regions employed iron-starvation induced proteins, a geranylgeranyl reductase, aquaporins, and urease, among other strategies, while limiting the use of iron-containing proteins. The metatranscriptomic survey performed here reveals functional differences in diatom communities and provides further insight into the environmental factors influencing the growth of diatoms and their predicted response to changes in ocean conditions. IMPORTANCEIn the Southern Ocean, phytoplankton must cope with harsh environmental conditions such as low light and growth-limiting concentrations of the micronutrient iron. Using metratranscriptomics, we assessed the influence of oceanographic variables on the diversity of the phytoplankton community composition and on the metabolic strategies of diatoms along the Western Antarctic Peninsula, a region undergoing rapid climate change. We found that cross-shelf differences in oceanographic parameters such as temperature and variable nutrient concentrations account for most of the differences in phytoplankton community composition and metabolism. We opportunistically characterized the metabolic underpinnings of a large bloom of the centric diatomActinocyclusin coastal waters of the WAP. Our results indicate that physicochemical differences from onshore to offshore are stronger than between southern and northern regions of the WAP; however, these trends could change in the future, resulting in poleward shifts in functional differences in diatom communities and phytoplankton blooms. 

Biogeochemical cycles constitute Earth’s life support system and distinguish our planet from others in this solar system. Microorganisms are the primary drivers of these cycles. Understanding the controls on marine microbial dynamics and how microbes will respond to environmental change is essential for building and assessing model-based forecasts and generating robust projections of climate change impacts on ocean productivity and biogeochemical cycles. An international community effort has been underway to create a global-scale marine microbial biogeochemistry research program to tackle gaps in this understanding. The BioGeoSCAPES: Ocean Metabolism and Nutrient Cycles on a Changing Planet program will identify and quantify how marine microbes adjust to a changing climate and assess the consequences for global biogeochemical cycles. This article summarizes the ongoing efforts to launch BioGeoSCAPES. 

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 The oceanic biogeochemical cycling of iron is globally important yet difficult to fully understand due to the many chemical processes involved. There is potential to use scandium, which has a similar ionic size and charge density to trivalent iron but lacks redox cycling, as a simpler analog for specific parts of the iron cycle, if we can sufficiently develop our understanding of scandium's reactivity. Here we move closer to this understanding. We look at particle reactivity and solubility through a 24‐hr incubation experiment: 5 nmol/kg of dissolved scandium and/or iron were added to filtered and unfiltered California Current System water. Particulate scandium formed only in the unfiltered treatments, at a quantity unlikely to have been taken up biologically. This is the first direct observation of scavenging of scandium, an attribute shared with iron. Our results also serve as the first test of scandium solubility in seawater: 1.9 nmol/kg of dissolved scandium was stable in the filtered treatment, 50 times more than the highest natural concentrations so far observed. This indicates that, in contrast to iron, scandium's oceanic cycling is unlikely to be influenced by solubility limits. We also compare particulate depth profiles: labile particulate iron was disproportionally higher than that of scandium in shelf‐influenced samples, likely due to iron reductively dissolving in the sediments, which scandium cannot do, and then precipitating in oxic seawater. Due to this combination of behaviors, our results suggest that paired observations of scandium and iron may help distinguish between iron sourced from sediment resuspension and reductive dissolution. 

Photosynthetic carbon (C) fixation by phytoplankton in the Southern Ocean (SO) plays a critical role in regulating air–sea exchange of carbon dioxide and thus global climate. In the SO, photosynthesis (PS) is often constrained by low iron, low temperatures, and low but highly variable light intensities. Recently, proton-pumping rhodopsins (PPRs) were identified in marine phytoplankton, providing an alternate iron-free, light-driven source of cellular energy. These proteins pump protons across cellular membranes through light absorption by the chromophore retinal, and the resulting pH energy gradient can then be used for active membrane transport or for synthesis of adenosine triphosphate. Here, we show that PPR is pervasive in Antarctic phytoplankton, especially in iron-limited regions. In a model SO diatom, we found that it was localized to the vacuolar membrane, making the vacuole a putative alternative phototrophic organelle for light-driven production of cellular energy. Unlike photosynthetic C fixation, which decreases substantially at colder temperatures, the proton transport activity of PPR was unaffected by decreasing temperature. Cellular PPR levels in cultured SO diatoms increased with decreasing iron concentrations and energy production from PPR photochemistry could substantially augment that of PS, especially under high light intensities, where PS is often photoinhibited. PPR gene expression and high retinal concentrations in phytoplankton in SO waters support its widespread use in polar environments. PPRs are an important adaptation of SO phytoplankton to growth and survival in their cold, iron-limited, and variable light environment. 

Abstract Synechococcus are the most abundant cyanobacteria in high latitude regions and are responsible for an estimated 17% of annual marine net primary productivity. Despite their biogeochemical importance, Synechococcus populations have been unevenly sampled across the ocean, with most studies focused on low-latitude strains. In particular, the near absence of Synechococcus genomes from high-latitude, High Nutrient Low Chlorophyll (HNLC) regions leaves a gap in our knowledge of picocyanobacterial adaptations to iron limitation and their influence on carbon, nitrogen, and iron cycles. We examined Synechococcus populations from the subarctic North Pacific, a well-characterized HNLC region, with quantitative metagenomics. Assembly with short and long reads produced two near complete Synechococcus metagenome-assembled genomes (MAGs). Quantitative metagenome-derived abundances of these populations matched well with flow cytometry counts, and the Synechococcus MAGs were estimated to comprise >99% of the Synechococcus at Station P. Whereas the Station P Synechococcus MAGs contained multiple genes for adaptation to iron limitation, both genomes lacked genes for uptake and assimilation of nitrate and nitrite, suggesting a dependence on ammonium, urea, and other forms of recycled nitrogen leading to reduced iron requirements. A global analysis of Synechococcus nitrate reductase abundance in the TARA Oceans dataset found nitrate assimilation genes are also lower in other HNLC regions. We propose that nitrate and nitrite assimilation gene loss in Synechococcus may represent an adaptation to severe iron limitation in high-latitude regions where ammonium availability is higher. Our findings have implications for models that quantify the contribution of cyanobacteria to primary production and subsequent carbon export. 

Abstract Southern Ocean (SO) diatoms play an important role in global carbon flux, and their influence on carbon export is directly linked to interactions with epiphytic bacteria. Bacterial symbionts that increase diatom growth promote atmospheric carbon uptake, while bacterial degraders divert diatom biomass into the microbial loop where it can then be released as carbon dioxide through respiration. To further explore SO diatom-bacterial associations, a natural model system is needed that is representative of these diverse and important interactions. Here, we use concurrent cultivation to isolate a species of the ecologically-important SO diatom, Pseudo-nitzschia subcurvata, and its co-occurring bacteria. Although vitamin-depleted, axenic Pseudo-nitzschia grew poorly in culture, addition of a co-isolated Roseobacter promoted diatom growth, while addition of a co-isolated Flavobacterium negatively impacted diatom growth. Microscopy revealed both bacterial isolates are physically associated with diatom cells and genome sequencing identified important predicted functions including vitamin synthesis, motility, cell attachment mechanisms, and diverse antimicrobial weapons that could be used for interbacterial competition. These findings revealed the natural coexistence of competing symbiotic strategies of diatom-associated bacteria in the SO, and the utility of this tripartite system, composed of a diatom and two bacterial strains, as a co-culture model to probe ecological-relevant interactions between diatoms and the bacteria that compete for access to the phycosphere.