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Concentrations and elemental stoichiometry of suspended particulate organic carbon, nitrogen, phosphorus, and oxygen demand for respiration (C:N:P:-O2) play a vital role in characterizing and quantifying marine elemental cycles. Here, we present Version 2 of the Global Ocean Particulate Organic Phosphorus, Carbon, Oxygen for Respiration, and Nitrogen (GO-POPCORN) dataset. Version 1 is a previously published dataset of particulate organic matter from 70 different studies between 1971 and 2010, while Version 2 is comprised of data collected from recent cruises between 2011 and 2020. The combined GO-POPCORN dataset contains 2673 paired surface POC/N/P measurements from 70°S to 73°N across all major ocean basins at high spatial resolution. Version 2 also includes 965 measurements of oxygen demand for organic carbon respiration. This new dataset can help validate and calibrate the next generation of global ocean biogeochemical models with flexible elemental stoichiometry. We expect that incorporating variable C:N:P:-O2 into models will help improve our estimates of key ocean biogeochemical fluxes such as carbon export, nitrogen fixation, and organic matter remineralization.

Linking 'omics measurements with biogeochemical cycles is a widespread challenge in microbial community ecology. Here, we propose applying genomic adaptation as 'biosensors' for microbial investments to overcome nutrient stress. We then integrate this genomic information with a trait-based model to predict regional shifts in the elemental composition of marine plankton communities. We evaluated this approach using metagenomic and particulate organic matter samples from the Atlantic, Indian and Pacific Oceans. We find that our genome-based trait model significantly improves our prediction of particulate C : P (carbon : phosphorus) across ocean regions. Furthermore, we detect previously unrecognized ocean areas of iron, nitrogen and phosphorus stress. In many ecosystems, it can be very challenging to quantify microbial stress. Thus, a carefully calibrated genomic approach could become a widespread tool for understanding microbial responses to environmental changes and the biogeochemical outcomes. This article is part of the theme issue 'Conceptual challenges in microbial community ecology'.

Surface ocean phosphate is commonly below the standard analytical detection limits, leading to an incomplete picture of the global variation and biogeochemical role of phosphate. A global compilation of phosphate measured using high-sensitivity methods revealed several previously unrecognized low-phosphate areas and clear regional differences. Both observational climatologies and Earth system models (ESMs) systematically overestimated surface phosphate. Furthermore, ESMs misrepresented the relationships between phosphate, phytoplankton biomass, and primary productivity. Atmospheric iron input and nitrogen fixation are known important controls on surface phosphate, but model simulations showed that differences in the iron-to-macronutrient ratio in the vertical nutrient supply and surface lateral transport are additional drivers of phosphate concentrations. Our study demonstrates the importance of accurately quantifying nutrients for understanding the regulation of ocean ecosystems and biogeochemistry now and under future climate conditions.

The Indian Ocean accounts for about one fifth of global ocean net primary production but remains undersampled relative to other major ocean basins. The eastern tropical Indian Ocean is characterized by extremely low concentrations of both macronutrients and the micronutrient iron. We measured concentrations of dissolved and particulate trace metals (Fe, Mn, Zn, Pb) in the upper ocean along the GO-SHIP IO9N transect (28˚S to 17˚N, mostly along the 95˚E meridian) during a cruise in April 2016. Cellular quotas (metal/C) of Fe, Mn, Co, Ni, Cu, and Zn were measured in small eukaryotic flagellates (2–7 μm). Deckboard bottle incubation experiments were conducted at one station in each of three putative biogeochemical regions: southern Indian Ocean gyre (SIO, 28-10˚S); equatorial Indian Ocean (EqIO, 10˚S - 5˚N); Bay of Bengal (BoB, 5-17˚N). Nitrate and phosphate were below detection limits in surface waters across the transect. Dissolved and particulate Fe were

The globally abundant marine Cyanobacteria Prochlorococcus and Synechococcus share many physiological traits but presumably have different evolutionary histories and associated phylogeography. In Prochlorococcus, there is a clear phylogenetic hierarchy of ecotypes, whereas multiple Synechococcus clades have overlapping physiologies and environmental distributions. However, microbial traits are associated with different phylogenetic depths. Using this principle, we reclassified diversity at different phylogenetic levels and compared the phylogeography. We sequenced the genetic diversity of Prochlorococcus and Synechococcus from 339 samples across the tropical Pacific Ocean and North Atlantic Ocean using a highly variable phylogenetic marker gene (rpoC1). We observed clear parallel niche distributions of ecotypes leading to high Pianka's Index values driven by distinct shifts at two transition points. The first transition point at 6°N distinguished ecotypes adapted to warm waters but separated by macronutrient content. At 39°N, ecotypes adapted to warm, low macronutrient vs. colder, high macronutrient waters shifted. Finally, we detected parallel vertical and regional single-nucleotide polymorphism microdiversity within clades from both Prochlorococcus and Synechococcus, suggesting uniquely adapted populations at very specific depths, as well as between the Atlantic and Pacific Oceans. Overall, this study demonstrates that Prochlorococcus and Synechococcus have shared phylogenetic organization of traits and associated phylogeography.

Variation in ocean C:N:P of particulate organic matter (POM) has led to competing hypotheses for the underlying drivers. Each hypothesis predicts C:N:P equally well due to regional co-variance in environmental conditions and biodiversity. The Indian Ocean offers a unique positive temperature and nutrient supply relationship to test these hypotheses. Here we show how elemental concentrations and ratios vary over daily and regional scales. POM concentrations were lowest in the southern gyre, elevated across the equator, and peaked in the Bay of Bengal. Elemental ratios were highest in the gyre, but approached Redfield proportions northwards. As Prochlorococcus dominated the phytoplankton community, biodiversity changes could not explain the elemental variation. Instead, our data supports the nutrient supply hypothesis. Finally, gyre dissolved iron concentrations suggest extensive iron stress, leading to depressed ratios compared to other gyres. We propose a model whereby differences in iron supply and N2-fixation influence C:N:P levels across ocean gyres.

Current hypotheses suggest that cellular elemental stoichiometry of marine eukaryotic phytoplankton such as the ratios of cellular carbon:nitrogen:phosphorus (C:N:P) vary between phylogenetic groups. To investigate how phylogenetic structure, cell volume, growth rate, and temperature interact to affect the cellular elemental stoichiometry of marine eukaryotic phytoplankton, we examined the C:N:P composition in 30 isolates across 7 classes of marine phytoplankton that were grown with a sufficient supply of nutrients and nitrate as the nitrogen source. The isolates covered a wide range in cell volume (5 orders of magnitude), growth rate (

The U.S. Culture Collection Network held a meeting to share information about how culture collections are responding to the requirements of the recently enacted Nagoya Protocol on Access to Genetic Resources and the Fair and Equitable Sharing of Benefits Arising from their Utilization to the Convention on Biological Diversity (CBD). The meeting included representatives of many culture collections and other biological collections, the U.S. Department of State, U.S. Department of Agriculture, Secretariat of the CBD, interested scientific societies, and collection groups, including Scientific Collections International and the Global Genome Biodiversity Network. The participants learned about the policies of the United States and other countries regarding access to genetic resources, the definition of genetic resources, and the status of historical materials and genetic sequence information. Key topics included what constitutes access and how the CBD Access and Benefit-Sharing Clearing-House can help guide researchers through the process of obtaining Prior Informed Consent on Mutually Agreed Terms. U.S. scientists and their international collaborators are required to follow the regulations of other countries when working with microbes originally isolated outside the United States, and the local regulations required by the Nagoya Protocol vary by the country of origin of the genetic resource. Managers of diverse living collections in the United States described their holdings and their efforts to provide access to genetic resources. This meeting laid the foundation for cooperation in establishing a set of standard operating procedures for U.S. and international culture collections in response to the Nagoya Protocol.

Phosphate (P) is an important nutrient potentially limiting for primary productivity, yet, we currently know little about the relationship between growth rate and physiological response to P limitation in abundant marine Cyanobacteria. Thus, the aim of this research was to determine how variation in growth rate affected the physiology of marine Synechococcus WH8102 and CC9311 when growing under high N:P conditions. Experiments were carried out in chemostats with a media input N:P of 441 and we estimated the half saturation concentration for growth under P limiting conditions (K s,p ) and cellular C:N:P ratios. The K s,p values were the lowest measured for any phytoplankton and on par with ambient P concentrations in oligotrophic regions. We also observed that both strains were able draw down P below 3 nM. Both K s,p and drawdown concentration were lower for the open ocean vs. coastal Synechococcus strain, which may be linked to differences in P acquisition genes in these strains. Cellular C:P and N:P ratios were significantly higher in relation to the Redfield ratio for both Synechococcus strains but we saw no difference in these ratios among growth rates or strains. These results demonstrate that Synechococcus can proliferate under very low P conditions and also that genetically different strains have unique physiological responses to P limitation.

We have a limited understanding of the consequences of variations in microbial biodiversity on ocean ecosystem functioning and global biogeochemical cycles. A core process is macronutrient uptake by microorganisms, as the uptake of nutrients controls ocean CO2 fixation rates in many regions. Here, we ask whether variations in ocean phytoplankton biodiversity lead to novel functional relationships between environmental variability and phosphate (Pi) uptake. We analyzed Pi uptake capabilities and cellular allocations among phytoplankton groups and the whole community throughout the extremely Pi-depleted western North Atlantic Ocean. Pi uptake capabilities of individual populations were well described by a classic uptake function but displayed adaptive differences in uptake capabilities that depend on cell size and nutrient availability. Using an eco-evolutionary model as well as observations of in situ uptake across the region, we confirmed that differences among populations lead to previously uncharacterized relationships between ambient Pi concentrations and uptake. Supported by novel theory, this work provides a robust empirical basis for describing and understanding assimilation of limiting nutrients in the oceans. Thus, it demonstrates that microbial biodiversity, beyond cell size, is important for understanding the global cycling of nutrients.

The ratio of carbon (C) to phosphorus (P) in marine phytoplankton is thought to be constant throughout the worlds'oceans. Known as the Redfield ratio, this relationship describes the links between carbon and phosphorus cycling and marine ecosystems. However, variations in the stoichiometry of phytoplankton have recently been identified, in particular strong latitudinal variability. Here we assess the impact of this variability in the C:P ratio of biomass on the C:P ratio of organic matter that is exported to the deep ocean using a biogeochemical inverse-model based on a data-constrained ocean circulation model and a global database of dissolved inorganic carbon and phosphate measurements. We identify global patterns of variability in the C:P ratios of exported organic matter, with higher values in the nutrient-depleted subtropical gyres, where organic matter export is relatively low, and lower ratios in nutrient-rich upwelling zones and high-latitude regions, where organic matter export is high. This suggests that total carbon export is relatively constant throughout the oceans, in agreement with recent estimates of carbon fluxes. We conclude that the latitudinal patterns of C:P in exported organic matter are consistent with the large-scale stoichiometric variations in phytoplankton C:P. We suggest that a future expansion of nutrient-depleted waters could result in a shift to more efficient C export that compensates for the expected decline in productivity.

Knowledge of concentrations and elemental ratios of suspended particles are important for understanding many biogeochemical processes in the ocean. These include patterns of phytoplankton nutrient limitation as well as linkages between the cycles of carbon and nitrogen or phosphorus. To further enable studies of ocean biogeochemistry, we here present a global dataset consisting of 100,605 total measurements of particulate organic carbon, nitrogen, or phosphorus analyzed as part of 70 cruises or time-series. The data are globally distributed and represent all major ocean regions as well as different depths in the water column. The global median C:P, N:P, and C:N ratios are 163, 22, and 6.6, respectively, but the data also includes extensive variation between samples from different regions. Thus, this compilation will hopefully assist in a wide range of future studies of ocean elemental ratios.

The concept of constant elemental ratios in plankton communities—the Redfield ratio—is of central importance to ocean biogeochemistry. Recently, several studies have demonstrated regional differences in the plankton C:P and N:P ratio. However, less is known about potential systematic variations in the C:N ratio. Here we present an analysis of the particulate organic carbon to nitrogen ratio of 40,482 globally distributed samples from the upper 200 m of the ocean water column. Particulate organic carbon and nitrogen concentrations are highly correlated (R 2 = 0.86) with a median value of 6.5. Using an artificial neural network analysis, we find regional variations in the C:N ratio linked to differences in environmental conditions. The ratio is lower in upper latitude cold water as well as upwelling regions in comparison to the warm oligotrophic gyres. We find substantial differences between ocean gyres that might be associated with differences in the nutrient supply ratio. Using cell sorting, we also quantified the C:N ratio of Prochlorococcus,Synechococcus, and picoeukaryotic field populations. The analysis demonstrates that picophytoplankton lineages exhibit a significantly higher ratio than the bulk particulate material but are only marginally significantly different from each other. Thus, the dominance of picophytoplankton in ocean gyres may contribute to the elevated ratios observed in these regions. Overall, the median C:N ratio derived from 40,482 samples is close to the canonical Redfield ratio, but significant regional deviations from this value are observed. These differences could be important for marine biogeochemistry and the regional coupling between the ocean's carbon and nitrogen cycles.

Nearly 75 years ago, Alfred C. Redfield observed a similarity between the elemental composition of marine plankton in the surface ocean and dissolved nutrients in the ocean interior. This stoichiometry, referred to as the Redfield ratio, continues to be a central tenet in ocean biogeochemistry, and is used to infer a variety of ecosystem processes, such as phytoplankton productivity and rates of nitrogen fixation and loss2-4. Model, field and laboratory studies have shown that different mechanisms can explain both constant and variable ratios of carbon to nitrogen and phosphorus among ocean plankton communities. The range of C/N/P ratios in the ocean, and their predictability, are the subject of much active research. Here we assess global patterns in the elemental composition of phytoplankton and particulate organic matter in the upper ocean, using published and unpublished observations of particulate phosphorus, nitrogen and carbon from a broad latitudinal range, supplemented with elemental data for surface plankton populations. We show that the elemental ratios of marine organic matter exhibit large spatial variations, with a global average that differs substantially from the canonical Redfield ratio. However, elemental ratios exhibit a clear latitudinal trend. Specifically, we observed a ratio of 195:28:1 in the warm nutrient-depleted low-latitude gyres, 137:18:1 in warm, nutrient-rich upwelling zones, and 78:13:1 in cold, nutrient-rich high-latitude regions. We suggest that the coupling between oceanic carbon, nitrogen and phosphorus cycles may vary systematically by ecosystem. © 2013 Macmillan Publishers Limited. All rights reserved.

This study investigated the impact of atmospheric metal deposition on natural phytoplankton communities at open-ocean and coastal sites in the Sargasso Sea during the spring bloom. Locally collected aerosols with different metal contents were added to natural phytoplankton assemblages from each site, and changes in nitrate, dissolved metal concentration, and phytoplankton abundance and carbon content were monitored. Addition of aerosol doubled the concentrations of cadmium (Cd), cobalt (Co), copper (Cu), iron (Fe), manganese (Mn), and nickel (Ni) in the incubation water. Over the 3-day experiments, greater drawdown of dissolved metals occurred in the open ocean water, whereas little metal drawdown occurred in the coastal water. Two populations of picoeukaryotic algae and Synechococcus grew in response to aerosol additions in both experiments. Particulate organic carbon increased and was most sensitive to changes in picoeukaryote abundance. Phytoplankton community composition differed depending on the chemistry of the aerosol added. Enrichment with aerosol that had higher metal content led to a 10-fold increase in Synechococcus abundance in the oceanic experiment but not in the coastal experiment. Enrichment of aerosol-derived Co, Mn, and Ni were particularly enhanced in the oceanic experiment, suggesting the Synechococcus population may have been fertilized by these aerosol metals. Cu-binding ligand concentrations were in excess of dissolved Cu in both experiments, and increased with aerosol additions. Bioavailable free hydrated Cu2+ concentrations were below toxicity thresholds throughout both experiments. These experiments show (1) atmospheric deposition contributes biologically important metals to seawater, (2) these metals are consumed over time scales commensurate with cell growth, and (3) growth responses can differ between distinct Synechococcus or eukaryotic algal populations despite their relatively close geographic proximity and taxonomic similarity.

The performance of 36 models (22 ocean color models and 14 biogeochemical ocean circulation models (BOGCMs)) that estimate depth-integrated marine net primary productivity (NPP) was assessed by comparing their output to in situ 14C data at the Bermuda Atlantic Time series Study (BATS) and the Hawaii Ocean Time series (HOT) over nearly two decades. Specifically, skill was assessed based on the models' ability to estimate the observed mean, variability, and trends of NPP. At both sites, more than 90% of the models underestimated mean NPP, with the average bias of the BOGCMs being nearly twice that of the ocean color models. However, the difference in overall skill between the best BOGCM and the best ocean color model at each site was not significant. Between 1989 and 2007, in situ NPP at BATS and HOT increased by an average of nearly 2% per year and was positively correlated to the North Pacific Gyre Oscillation index. The majority of ocean color models produced in situ NPP trends that were closer to the observed trends when chlorophyll-a was derived from high-performance liquid chromatography (HPLC), rather than fluorometric or SeaWiFS data. However, this was a function of time such that average trend magnitude was more accurately estimated over longer time periods. Among BOGCMs, only two individual models successfully produced an increasing NPP trend (one model at each site). We caution against the use of models to assess multiannual changes in NPP over short time periods. Ocean color model estimates of NPP trends could improve if more high quality HPLC chlorophyll-a time series were available.