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Abstract Each spring, the North Atlantic experiences one of the largest open‐ocean phytoplankton blooms in the global ocean. Diatoms often dominate the initial phase of the bloom with succession driven by exhaustion of silicic acid. The North Atlantic was sampled over 3.5 weeks in spring 2021 following the demise of the main diatom bloom, allowing mechanisms that sustain continued diatom contributions to be examined. Diatom biomass was initially relatively high with biogenic silica concentrations up to 2.25 μmol Si L⁻¹. A low initial silicic acid concentration of 0.1–0.3 μM imposed severe Si limitation of silica production and likely limited the diatom growth rate. Four storms over the next 3.5 weeks entrained silicic acid into the mixed layer, relieving growth limitation, but uptake limitation persisted. Silica production was modest and dominated by the >5.0 μm size fraction although specific rates were highest in the 0.6–5.0 μm size fraction over most of the cruise. Silica dissolution averaged 68% of silica production. The resupply of silicic acid via storm entrainment and silica dissolution supported a cumulative post‐bloom silica production that was 32% of that estimated during the main bloom event. Diatoms contributed significantly to new and to primary production after the initial bloom, possibly dominating both. Diatom contribution to organic‐carbon export was also significant at 40%–70%. Thus, diatoms can significantly contribute to regional biogeochemistry following initial silicic acid depletion, but that contribution relies on physical processes that resupply the nutrient to surface waters. 

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. 

This dataset includes trace metal (iron, manganese, cobalt, nickel, copper, zinc, cadmium, lead) and macronutrient (nitrate&nitrite, nitrite, phosphate, silicic acid) concentration data from incubation experiments conducted on board the RRS Discovery during the EXPORTS North Atlantic campaign at the Porcupine Abyssal Plain-Sustained Observatory (PAP-SO) site (DY131). In these experiments, additions of macronutrients (N, P, Si) and Fe were used to assess the level of Si, N, and Fe stress being experienced by the phytoplankton and to contextualize taxa-specific metatranscriptome responses for resolving gene expression profiles in the in-situ communities. This research project focuses on the vertical export of the carbon associated with a major group of phytoplankton, the diatoms in the North Atlantic near the Porcupine Abyssal Plain. The major objective is to understand how diatom community composition and the prevailing nutrient conditions create taxonomic differences in metabolic state that combine to direct diatom taxa to different carbon export pathways. The focus is on diatoms, given their large contribution to global marine primary productivity and carbon export which translates into a significant contribution to the biogeochemical cycling of carbon (C), nitrogen (N), phosphorus (P), iron (Fe) and silicon (Si). It is hypothesized that the type and degree of diatom physiological stress are vital aspects of ecosystem state that drive export. To test this hypothesis, combined investigator expertise in phytoplankton physiology, genomics, and trace element chemistry is used to assess the rates of nutrient use and the genetic composition and response of diatom communities, with measurements of silicon and iron stress to evaluate stress as a predictor of the path of diatom carbon export. The EXPORTS field campaign in the North Atlantic sampled a retentive eddy over nearly a month in May 2021, which coincided with the decline of the North Atlantic Spring Bloom. 

This dataset includes concentrations of dissolved (<0.4 micrometers (µm)) and labile particulate (0.4-5 µm and >5 µm) phosphorus (P), vanadium (V), manganese (Mn), iron (Fe), cobalt (Co), nickel (Ni), copper (Cu), zinc (Zn), cadmium (Cd), and lead (Pb) in shipboard incubation samples collected during the EXports Processes in the Oceans from RemoTe Sensing (EXPORTS) North Pacific (NP) cruise RR1813 on the R/V Roger Revelle near Ocean Station PAPA (Station P). 

This dataset includes concentrations of dissolved (<0.2 micrometers (µm)) manganese (Mn), iron (Fe), cobalt (Co), nickel (Ni), copper (Cu), zinc (Zn), cadmium (Cd), and lead (Pb) in depth profile samples collected on 8 September 2018 during the EXports Processes in the Oceans from RemoTe Sensing (EXPORTS) cruise aboard R/V Roger Revelle cruise RR1813 at Ocean Station PAPA (Station P). 

This dataset includes depth profiles in the euphotic zone of nutrient (nitrate, silicate, phosphate) concentrations and profiles of silicic acid uptake rates from seawater samples collected on EXPORTS cruise DY131 during May 2021. This research focuses on the vertical export of the carbon associated with a major group of phytoplankton, the diatoms in the North Atlantic near the Porcupine Abyssal Plain. The major objective is to understand how diatom community composition and the prevailing nutrient conditions create taxonomic differences in metabolic state that combine to direct diatom taxa to different carbon export pathways. The focus is on diatoms, given their large contribution to global marine primary productivity and carbon export which translates into a significant contribution to the biogeochemical cycling of carbon (C), nitrogen (N), phosphorus (P), iron (Fe) and silicon (Si). It is hypothesized that the type and degree of diatom physiological stress are vital aspects of ecosystem state that drive export. To test this hypothesis, combined investigator expertise in phytoplankton physiology, genomics, and trace element chemistry is used to assess the rates of nutrient use and the genetic composition and response of diatom communities, with measurements of silicon and iron stress to evaluate stress as a predictor of the path of diatom carbon export. The EXPORTS field campaign in the North Atlantic sampled a retentive eddy over nearly a month. At the beginning of the cruise, nitrate was abundant while silicic acid was nearly undetectable. Such low dissolved Si concentrations significantly limit diatom silicification resulting in diatoms with reduced mineral ballast and low Si:C and Si:N ratios that would reduce sinking rates and competition for Si can alter diatom taxonomic composition. Both factors can the path cells follow through the food web ultimately altering diatom carbon export. Within each ecosystem state examined in the EXPORTS program, nutrient biogeochemistry, diatom and phytoplankton community structure, and global diatom gene expression patterns (metatranscriptomics) are characterized in the ocean. Nutrient amendment experiments with tracer addition (14C, 32Si) are used to quantify the level of Si, N, and Fe stress being experienced by the phytoplankton and to contextualize taxa-specific metatranscriptome responses for resolving gene expression profiles in the in situ communities. 

This dataset includes concentrations of dissolved (<0.2 micrometers (µm)) manganese (Mn), iron (Fe), cobalt (Co), nickel (Ni), copper (Cu), zinc (Zn), cadmium (Cd), and lead (Pb) in surface (~2 meters) samples collected between 15 August 2018 and 6 September 2018 during the EXports Processes in the Oceans from RemoTe Sensing (EXPORTS) cruise aboard R/V Roger Revelle cruise RR1813 at Ocean Station PAPA (Station P). 

This dataset includes data from the nutrient amendment experiments. In these experiments, tracer additions (14C, 32Si) were used to quantify the level of Si, N, and Fe stress being experienced by the phytoplankton and to contextualize taxa-specific metatranscriptome responses for resolving gene expression profiles in the in situ communities. Seawater samples were collected on EXPORTS cruise DY131 during May 2021. This research focuses on the vertical export of the carbon associated with a major group of phytoplankton, the diatoms in the North Atlantic near the Porcupine Abyssal Plain. The major objective is to understand how diatom community composition and the prevailing nutrient conditions create taxonomic differences in metabolic state that combine to direct diatom taxa to different carbon export pathways. The focus is on diatoms, given their large contribution to global marine primary productivity and carbon export which translates into a significant contribution to the biogeochemical cycling of carbon (C), nitrogen (N), phosphorus (P), iron (Fe) and silicon (Si). It is hypothesized that the type and degree of diatom physiological stress are vital aspects of ecosystem state that drive export. To test this hypothesis, combined investigator expertise in phytoplankton physiology, genomics, and trace element chemistry is used to assess the rates of nutrient use and the genetic composition and response of diatom communities, with measurements of silicon and iron stress to evaluate stress as a predictor of the path of diatom carbon export. The EXPORTS field campaign in the North Atlantic sampled a retentive eddy over nearly a month. At the beginning of the cruise, nitrate was abundant while silicic acid was nearly undetectable. Such low dissolved Si concentrations significantly limit diatom silicification resulting in diatoms with reduced mineral ballast and low Si:C and Si:N ratios that would reduce sinking rates and competition for Si can alter diatom taxonomic composition. Both factors can the path cells follow through the food web ultimately altering diatom carbon export. Within each ecosystem state examined in the EXPORTS program, nutrient biogeochemistry, diatom and phytoplankton community structure, and global diatom gene expression patterns (metatranscriptomics) are characterized in the ocean. Nutrient amendment experiments with tracer addition (14C, 32Si) are used to quantify the level of Si, N, and Fe stress being experienced by the phytoplankton and to contextualize taxa-specific metatranscriptome responses for resolving gene expression profiles in the in situ communities. 

Diatoms are major contributors to marine primary productivity and carbon export due to their rapid growth in high-nutrient environments and their heavy silica ballast. Their contributions are highly modified in high-nutrient low-chlorophyll regions due to the decoupling of upper-ocean silicon and carbon cycling caused by low iron (Fe). The Si cycle and the role of diatoms in the biological carbon pump was examined at Ocean Station Papa (OSP) in the HNLC region of the northeastern subarctic Pacific during the NASA EXport Processes in the Ocean from RemoTe Sensing (EXPORTS) field study. Sampling occurred during the annual minimum in surface silicic acid (Si(OH)4) concentration. Biogenic silica (bSi) concentrations were low, being in the tens of nanomolar range, despite high Si(OH)4 concentrations of about 15 μM. On average, the >5.0-µm particle size fraction dominated Si dynamics, accounting for 65% of bSi stocks and 81% of Si uptake compared to the small fraction (0.6–5.0 μm). Limitation of Si uptake was detected in the small, but not the large, size fraction. Growth rate in small diatoms was limited by Fe, while their Si uptake was restricted by Si(OH)4 concentration, whereas larger diatoms were only growth-limited by Fe. About a third of bSi production was exported out of the upper 100 m. The contribution of diatoms to carbon export (9–13%) was about twice their contribution to primary productivity (3–7%). The combination of low bSi production, low diatom primary productivity and high bSi export efficiency at OSP was more similar to the dynamics in the subtropical gyres than to other high-nutrient low-chlorophyll regions. 

The goal of the EXport Processes in the Ocean from RemoTe Sensing (EXPORTS) field campaign is to develop a predictive understanding of the export, fate, and carbon cycle impacts of global ocean net primary production. To accomplish this goal, observations of export flux pathways, plankton community composition, food web processes, and optical, physical, and biogeochemical (BGC) properties are needed over a range of ecosystem states. Here we introduce the first EXPORTS field deployment to Ocean Station Papa in the Northeast Pacific Ocean during summer of 2018, providing context for other papers in this special collection. The experiment was conducted with two ships: a Process Ship, focused on ecological rates, BGC fluxes, temporal changes in food web, and BGC and optical properties, that followed an instrumented Lagrangian float; and a Survey Ship that sampled BGC and optical properties in spatial patterns around the Process Ship. An array of autonomous underwater assets provided measurements over a range of spatial and temporal scales, and partnering programs and remote sensing observations provided additional observational context. The oceanographic setting was typical of late-summer conditions at Ocean Station Papa: a shallow mixed layer, strong vertical and weak horizontal gradients in hydrographic properties, sluggish sub-inertial currents, elevated macronutrient concentrations and low phytoplankton abundances. Although nutrient concentrations were consistent with previous observations, mixed layer chlorophyll was lower than typically observed, resulting in a deeper euphotic zone. Analyses of surface layer temperature and salinity found three distinct surface water types, allowing for diagnosis of whether observed changes were spatial or temporal. The 2018 EXPORTS field deployment is among the most comprehensive biological pump studies ever conducted. A second deployment to the North Atlantic Ocean occurred in spring 2021, which will be followed by focused work on data synthesis and modeling using the entire EXPORTS data set.