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Creators/Authors contains: "Mulholland, Margaret R."

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  1. Abstract

    Viruses play an important role in the ecology and biogeochemistry of marine ecosystems. Beyond mortality and gene transfer, viruses can reprogram microbial metabolism during infection by expressing auxiliary metabolic genes (AMGs) involved in photosynthesis, central carbon metabolism, and nutrient cycling. While previous studies have focused on AMG diversity in the sunlit and dark ocean, less is known about the role of viruses in shaping metabolic networks along redox gradients associated with marine oxygen minimum zones (OMZs). Here, we analyzed relatively quantitative viral metagenomic datasets that profiled the oxygen gradient across Eastern Tropical South Pacific (ETSP) OMZ waters, assessing whether OMZ viruses might impact nitrogen (N) cycling via AMGs. Identified viral genomes encoded six N-cycle AMGs associated with denitrification, nitrification, assimilatory nitrate reduction, and nitrite transport. The majority of these AMGs (80%) were identified in T4-like Myoviridae phages, predicted to infect Cyanobacteria and Proteobacteria, or in unclassified archaeal viruses predicted to infect Thaumarchaeota. Four AMGs were exclusive to anoxic waters and had distributions that paralleled homologous microbial genes. Together, these findings suggest viruses modulate N-cycling processes within the ETSP OMZ and may contribute to nitrogen loss throughout the global oceans thus providing a baseline for their inclusion in the ecosystem and geochemical models.

     
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  2. Abstract

    Examination of dinitrogen (N2) fixation in the Eastern Tropical South Pacific oxygen deficient zone has raised questions about the range of diazotrophs in the deep sea and their quantitative importance as a source of new nitrogen globally. However, technical considerations in the deployment of stable isotopes in quantifying N2fixation rates have complicated interpretation of this research. Here, we report the findings of a comprehensive survey of N2fixation within, above and below the Eastern Tropical South Pacific oxygen deficient zone. N2fixation rates were measured using a robust15N tracer method (bubble removal) that accounts for the slow dissolution of N2gas and calculated using a conservative approach. N2fixation was only detected in a subset of samples (8 of 125 replicated measurements) collected within suboxic waters (< 20 μmol O2kg−1) or at the oxycline. Most of these detectable rates were measured at nearshore stations, or where surface productivity was high. These findings support the hypothesis that low oxygen/high organic carbon conditions favor non‐cyanobacterial diazotrophs. Nevertheless, this study indicates that N2fixation is neither widespread nor quantitatively important throughout this region.

     
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  3. Abstract

    In the North Atlantic Ocean, dinitrogen (N2) fixation on the western continental shelf represents a significant fraction of basin‐wide nitrogen (N) inputs. However, the factors regulating coastal N2fixation remain poorly understood, in part due to sharp physico‐chemical gradients and dynamic water mass interactions that are difficult to constrain via traditional oceanographic approaches. This study sought to characterize the spatial heterogeneity of N2fixation on the western North Atlantic shelf, at the confluence of Mid‐ and South Atlantic Bight shelf waters and the Gulf Stream, in August 2016. Rates were quantified using the15N2bubble release method and used to build empirical models of regional N2fixation via a random forest machine learning approach. N2fixation rates were then predicted from high‐resolution CTD and satellite data to infer the variability of its depth and surface distributions, respectively. Our findings suggest that the frontal mixing zone created conditions conducive to exceptionally high N2fixation rates (> 100 nmol N L−1d−1), which were likely driven by the haptophyte‐symbiont UCYN‐A. Above and below this hotspot, N2fixation rates were highest on the shelf due to the high particulate N concentrations there. Conversely, specific N2uptake rates, a biomass‐independent metric for diazotroph activity, were enhanced in the oligotrophic slope waters. Broadly, these observations suggest that N2fixation is favored offshore but occurs continuously across the shelf. Nevertheless, our model results indicate that there is a niche for diazotrophs along the coastline as phytoplankton populations begin to decline, likely due to exhaustion of coastal nutrients.

     
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  4. Summary

    Oxygen minimum zones (OMZs) are critical to marine nitrogen cycling and global climate change. While OMZ microbial communities are relatively well‐studied, little is known about their viruses. Here, we assess the viral community ecology of 22 deeply sequenced viral metagenomes along a gradient of oxygenated to anoxic waters (<0.02 μmol/l O2) in the Eastern Tropical South Pacific (ETSP) OMZ. We identified 46 127 viral populations (≥5 kb), which augments the known viruses from ETSP by 10‐fold. Viral communities clustered into six groups that correspond to oceanographic features. Oxygen concentration was the predominant environmental feature driving viral community structure. Alpha and beta diversity of viral communities in the anoxic zone were lower than in surface waters, which parallels the low microbial diversity seen in other studies. ETSP viruses were largely endemic, with the majority of shared viruses (87%) also present in other OMZ samples. We detected 543 putative viral‐encoded auxiliary metabolic genes (AMGs), of which some have a distribution that reflects physico‐chemical characteristics across depth. Together these findings provide an ecological baseline for viral community structure, drivers and population variability in OMZs that will help future studies assess the role of viruses in these climate‐critical environments.

     
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  5. Abstract

    Recent work has suggested that the oxygen deficient zone (ODZ) and overlying surface waters of the eastern tropical South Pacific (ETSP) is a potential niche for dinitrogen (N2) fixation. Rates of dinitrogen fixation were measured in the ETSP above and within the ODZ in July 2013 using a modified15N2bubble addition method, wherein a bubble was added, mixed, and then removed, and the isotopic enrichment of the dissolved N2was measured directly for each incubation. N2fixation rates in the euphotic zone ranged from below detection to 3.9 nmol L−1d−1and were below detection at all depths surveyed within the ODZ. Depth‐integrated rates ranged from below detection to 289.7μmol m−2d−1. DNA and RNA of diversenifHgenes were detected at both surface waters and in the ODZ. However, the results of this study suggest that N2fixation rates were low and contribute little to N cycling in the ETSP.

     
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