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Abstract. Across the Arctic, vast areas of permafrost are being degraded by climatechange, which has the potential to release substantial quantities ofnutrients, including nitrogen into large Arctic rivers. These rivers heavilyinfluence the biogeochemistry of the Arctic Ocean, so it is important tounderstand the potential changes to rivers from permafrost degradation. Thisstudy utilized dissolved nitrogen species (nitrate and dissolved organicnitrogen (DON)) along with nitrogen isotope values (δ15N-NO3- and δ15N-DON) of samples collectedfrom permafrost sites in the Kolyma River and the six largest Arctic rivers.Large inputs of DON and nitrate with a unique isotopically heavy δ15N signature were documented in the Kolyma, suggesting the occurrenceof denitrification and highly invigorated nitrogen cycling in the Yedomapermafrost thaw zones along the Kolyma. We show evidence for permafrost-derived DON being recycled to nitrate as it passes through the river,transferring the high 15N signature to nitrate. However, the potentialto observe these thaw signals at the mouths of rivers depends on the spatialscale of thaw sites, permafrost degradation, and recycling mechanisms. Incontrast with the Kolyma, with near 100 % continuous permafrost extent,the Ob River, draining large areas of discontinuous and sporadicpermafrost, shows large seasonal changes in both nitrate and DON isotopicsignatures. During winter months, water percolating through peat soilsrecords isotopically heavy denitrification signals in contrast with thelighter summer values when surface flow dominates. This early yeardenitrification signal was present to a degree in the Kolyma, but the abilityto relate seasonal nitrogen signals across Arctic Rivers to permafrostdegradation could not be shown with this study. Other large rivers in theArctic show different seasonal nitrogen trends. Based on nitrogen isotopevalues, the vast majority of nitrogen fluxes in the Arctic rivers is fromfresh DON sourced from surface runoff through organic-rich topsoil and notfrom permafrost degradation. However, with future permafrost thaw, otherArctic rivers may begin to show nitrogen trends similar to the Ob. Ourstudy demonstrates that nitrogen inputs from permafrost thaw can beidentified through nitrogen isotopes, but only on small spatial scales.Overall, nitrogen isotopes show potential for revealing integrated catchmentwide nitrogen cycling processes. 

Abstract. Trichodesmium is a globally important marine microbe that provides fixednitrogen (N) to otherwise N-limited ecosystems. In nature, nitrogen fixationis likely regulated by iron or phosphate availability, but the extent andinteraction of these controls are unclear. From metaproteomics analysesusing established protein biomarkers for nutrient stress, we foundthat iron–phosphate co-stress is the norm rather than the exception for Trichodesmium colonies in theNorth Atlantic Ocean. Counterintuitively, the nitrogenase enzyme was moreabundant under co-stress as opposed to single nutrient stress. This isconsistent with the idea that Trichodesmium has a specific physiological state duringnutrient co-stress. Organic nitrogen uptake was observed and occurredsimultaneously with nitrogen fixation. The quantification of the phosphate ABCtransporter PstA combined with a cellular model of nutrient uptake suggestedthat Trichodesmium is generally confronted by the biophysical limits of membrane spaceand diffusion rates for iron and phosphate acquisition in the field. Colonyformation may benefit nutrient acquisition from particulate and organicsources, alleviating these pressures. The results highlight that topredict the behavior of Trichodesmium, both Fe and P stress must be evaluatedsimultaneously. 

Abstract Nano‐ and picophytoplankton are a major component of open‐ocean ecosystems and one of the main plankton functional types in biogeochemical models, yet little is known about their trace metal contents. In cultures of the picoeukaryoteOstreococcus lucimarinus, iron limitation reduced iron quotas by 68%, a fraction of the plasticity known in diatoms. In contrast, a commonly co‐occurring cyanobacterium,Prochlorococcus, showed variable iron contents with iron availability in culture. Synchrotron X‐ray fluorescence was used to measure single‐cell metal (Mn, Fe, Co, Ni, Zn) quotas of autotrophic flagellates (1.4–16.8‐μm diameter) collected from four ocean regions. Iron quotas were tightly constrained and showed little response to iron availability, similar to culturedOstreococcus. Zinc quotas also did not vary with zinc availability but appeared to vary with phosphorus availability. These results suggest that macronutrient and metal availability may be equally important for controlling metal contents of small eukaryotic open‐ocean phytoplankton. 

Abstract. Marine diazotrophs convert dinitrogen (N2) gas intobioavailable nitrogen (N), supporting life in the global ocean. In 2012, thefirst version of the global oceanic diazotroph database (version 1) waspublished. Here, we present an updated version of the database (version 2),significantly increasing the number of in situ diazotrophic measurements from13 565 to 55 286. Data points for N2 fixation rates, diazotrophic cellabundance, and nifH gene copy abundance have increased by 184 %, 86 %, and809 %, respectively. Version 2 includes two new data sheets for the nifH genecopy abundance of non-cyanobacterial diazotrophs and cell-specific N2fixation rates. The measurements of N2 fixation rates approximatelyfollow a log-normal distribution in both version 1 and version 2. However,version 2 considerably extends both the left and right tails of thedistribution. Consequently, when estimating global oceanic N2 fixationrates using the geometric means of different ocean basins, version 1 andversion 2 yield similar rates (43–57 versus 45–63 Tg N yr−1; rangesbased on one geometric standard error). In contrast, when using arithmeticmeans, version 2 suggests a significantly higher rate of 223±30 Tg N yr−1 (mean ± standard error; same hereafter) compared to version 1(74±7 Tg N yr−1). Specifically, substantial rate increases areestimated for the South Pacific Ocean (88±23 versus 20±2 Tg N yr−1), primarily driven by measurements in the southwestern subtropics,and for the North Atlantic Ocean (40±9 versus 10±2 Tg N yr−1). Moreover, version 2 estimates the N2 fixation rate in theIndian Ocean to be 35±14 Tg N yr−1, which could not be estimatedusing version 1 due to limited data availability. Furthermore, a comparisonof N2 fixation rates obtained through different measurement methods atthe same months, locations, and depths reveals that the conventional15N2 bubble method yields lower rates in 69 % cases compared tothe new 15N2 dissolution method. This updated version of thedatabase can facilitate future studies in marine ecology andbiogeochemistry. The database is stored at the Figshare repository(https://doi.org/10.6084/m9.figshare.21677687; Shao etal., 2022).