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Abstract. Recent studies show that stoichiometric elemental ratios of marine ecosystems are not static at Redfield proportions but vary systematically between biomes. However, the wider Atlantic Ocean is undersampled for particulate organic matter (POM) elemental composition, especially when it comes to phosphorus (i.e., POP). Thus, it is uncertain how environmental variation in this region translates into shifts in the C:N:P ratio. To address this, we analyzed hydrography, genomics, and POM concentrations from 877 stations on the meridional transects AMT28 and C13.5, spanning the Atlantic Ocean. We observed nutrient-replete, high-latitude ecosystem C:N:P to be significantly lower than that in the oligotrophic gyres. Latitudinal and zonal differences in elemental stoichiometry were linked to overall nutrient supply as well as N vs. P stress. C:P and N:P were generally higher in the P-stressed northern region compared to Southern Hemisphere regions. We also detected a zonal difference linked to a westward deepening nutricline and a shift from N to P stress. We also evaluated possible seasonal changes in C:N:P across the basin and predicted these to be limited. Overall, this study confirms latitudinal shifts in surface ocean POM ratios but reveals previously unrecognized hemisphere and zonal gradients. This work demonstrates the importance of understanding how regional shifts in hydrography and type of nutrient stress shape the coupling between Atlantic Ocean nutrient and carbon cycles. 

Abstract Oceanic nutrient cycles are coupled, yet carbon-nitrogen-phosphorus (C:N:P) stoichiometry in marine ecosystems is variable through space and time, with no clear consensus on the controls on variability. Here, we analyze hydrographic, plankton genomic diversity, and particulate organic matter data from 1970 stations sampled during a global ocean observation program (Bio-GO-SHIP) to investigate the biogeography of surface ocean particulate organic matter stoichiometry. We find latitudinal variability in C:N:P stoichiometry, with surface temperature and macronutrient availability as strong predictors of stoichiometry at high latitudes. Genomic observations indicated community nutrient stress and suggested that nutrient supply rate and nitrogen-versus-phosphorus stress are predictive of hemispheric and regional variations in stoichiometry. Our data-derived statistical model suggests that C:P and N:P ratios will increase at high latitudes in the future, however, changes at low latitudes are uncertain. Our findings suggest systematic regulation of elemental stoichiometry among ocean ecosystems, but that future changes remain highly uncertain. 

Abstract Concentrations and elemental stoichiometry of suspended particulate organic carbon, nitrogen, phosphorus, and oxygen demand for respiration (C:N:P:−O 2 ) 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:-O 2 into models will help improve our estimates of key ocean biogeochemical fluxes such as carbon export, nitrogen fixation, and organic matter remineralization. 

Abstract Ocean ecosystem models predict that warming and increased surface ocean stratification will trigger a series of ecosystem events, reducing the biological export of particulate carbon to the ocean interior. We present a nearly three-decade time series from the open ocean that documents a biological response to ocean warming and nutrient reductions wherein particulate carbon export is maintained, counter to expectations. Carbon export is maintained through a combination of phytoplankton community change to favor cyanobacteria with high cellular carbon-to-phosphorus ratios and enhanced shallow phosphorus recycling leading to increased nutrient use efficiency. These results suggest that surface ocean ecosystems may be more responsive and adapt more rapidly to changes in the hydrographic system than is currently envisioned in earth ecosystem models, with positive consequences for ocean carbon uptake. 

Abstract. We describe the third version of Minnesota Earth System Model for Oceanbiogeochemistry (MESMO 3), an Earth system model of intermediate complexity,with a dynamical ocean, dynamic–thermodynamic sea ice, and an energy–moisture-balanced atmosphere. A major feature of version 3 is the flexibleC:N:P ratio for the three phytoplankton functional types represented in themodel. The flexible stoichiometry is based on the power law formulation withenvironmental dependence on phosphate, nitrate, temperature, and light.Other new features include nitrogen fixation, water column denitrification,oxygen and temperature-dependent organic matter remineralization, andCaCO3 production based on the concept of the residual nitrate potentialgrowth. In addition, we describe the semi-labile and refractory dissolved organicpools of C, N, P, and Fe that can be enabled in MESMO 3 as an optionalfeature. The refractory dissolved organic matter can be degraded byphotodegradation at the surface and hydrothermal vent degradation at thebottom. These improvements provide a basis for using MESMO 3 in furtherinvestigations of the global marine carbon cycle to changes in theenvironmental conditions of the past, present, and future. 

Abstract Using a global ocean biogeochemistry model, we examined three drivers of global ocean production C:N:P ratio: flexible phytoplankton stoichiometry, phytoplankton community composition, and regional production shifts. For a middle-of-the-road warming scenario (SSP2), the model predicts a substantial increase in the global export C:P ratio from 113:1 to 119:1 by the year 2100. The most important physiological driver of this stoichiometric change is the effect of the worldwide warming on cyanobacteria, followed by the effect of phosphate depletion on eukaryotes in the Southern Ocean. Also, there is a modest global shift in the phytoplankton community in favor of cyanobacteria at the expense of eukaryotes with a minimal effect on the global production stoichiometry. We find that shifts in the regional production, even in the absence of any change in phytoplankton stoichiometry or taxonomy, can change the global production C:N:P ratio. For example, enhancing the production in the polar waters, which typically have low C:N:P ratios, will have the effect of lowering the global ratio. In our model, the retreat of Antarctic sea ice has this very effect but is offset by production changes downstream and elsewhere. This study thus provides an understanding of how regional production changes can affect the global production C:N:P ratio. However, the current literature indicates substantial uncertainty in the future projections of regional production changes, so it is unclear at this time what their net effect is on the global production C:N:P ratio. Finally, our model predicts that the overall increase in the carbon content of organic matter due to flexible C:N:P ratio helps to stabilize carbon export in the face of reduced nutrient export (i.e. the decrease in C export is ~30% smaller than expected from the decrease in P export by 2100) but has a minimal effect on atmospheric CO₂uptake (~1%). 

Abstract. The elemental stoichiometry of marine phytoplankton plays a critical role in global biogeochemical cycles through its impact on nutrient cycling, secondary production, and carbon export. Although extensive laboratory experiments have been carried out over the years to assess the influence of different environmental drivers on the elemental composition of phytoplankton, a comprehensive quantitative assessment of the processes is still lacking. Here, we synthesized the responses of P:C and N:C ratios of marine phytoplankton to five major drivers (inorganic phosphorus, inorganic nitrogen, inorganic iron, irradiance, and temperature) by a meta-analysis of experimental data across 366 experiments from 104 journal articles. Our results show that the response of these ratios to changes in macronutrients is consistent across all the studies, where the increase in nutrient availability is positively related to changes in P:C and N:C ratios. We found that eukaryotic phytoplankton are more sensitive to the changes in macronutrients compared to prokaryotes, possibly due to their larger cell size and their abilities to regulate their gene expression patterns quickly. The effect of irradiance was significant and constant across all studies, where an increase in irradiance decreased both P:C and N:C. The P:C ratio decreased significantly with warming, but the response to temperature changes was mixed depending on the culture growth mode and the growth phase at the time of harvest. Along with other oceanographic conditions of the subtropical gyres (e.g., low macronutrient availability), the elevated temperature may explain why P:C is consistently low in subtropical oceans. Iron addition did not systematically change either P:C or N:C. Overall, our findings highlight the high stoichiometric plasticity of eukaryotes and the importance of macronutrients in determining P:C and N:C ratios, which both provide us insights on how to understand and model plankton diversity and productivity. 

Abstract The strength of the biological soft tissue pump in the ocean critically depends on how much organic carbon is produced via photosynthesis and how efficiently the carbon is transferred to the ocean interior. For a given amount of limiting nutrient, phosphate, soft tissue pump would be strengthened if the carbon (C) to phosphorus (P) ratio of sinking organic matter increases as the remineralization length scale of C increases. Here, we present a new data compilation of particle flux stoichiometry and show that C:P of sinking particulate organic matter (POM) in the ocean twilight zone on average is likely to be higher than the C:P ratio of surface suspended POM. We further demonstrate using a physics‐biology coupled global ocean model combined with a theory from first principles that an increase in C:P export flux ratio in the ocean's twilight zone can lead to a considerable drawdown of atmosphericpCO₂.