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  • 1
    Publication Date: 2022-05-26
    Description: © The Author(s), 2020. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Wilson, S. T., Al-Haj, A. N., Bourbonnais, A., Frey, C., Fulweiler, R. W., Kessler, J. D., Marchant, H. K., Milucka, J., Ray, N. E., Suntharalingam, P., Thornton, B. F., Upstill-Goddard, R. C., Weber, T. S., Arevalo-Martinez, D. L., Bange, H. W., Benway, H. M., Bianchi, D., Borges, A., V., Chang, B. X., Crill, P. M., del Valle, D. A., Farias, L., Joye, S. B., Kock, A., Labidi, J., Manning, C. C., Pohlman, J. W., Rehder, G., Sparrow, K. J., Tortell, P. D., Treude, T., Valentine, D. L., Ward, B. B., Yang, S., & Yurganov, L. N. Ideas and perspectives: a strategic assessment of methane and nitrous oxide measurements in the marine environment. Biogeosciences, 17(22), (2020): 5809-5828, https://doi.org/10.5194/bg-17-5809-2020.
    Description: In the current era of rapid climate change, accurate characterization of climate-relevant gas dynamics – namely production, consumption, and net emissions – is required for all biomes, especially those ecosystems most susceptible to the impact of change. Marine environments include regions that act as net sources or sinks for numerous climate-active trace gases including methane (CH4) and nitrous oxide (N2O). The temporal and spatial distributions of CH4 and N2O are controlled by the interaction of complex biogeochemical and physical processes. To evaluate and quantify how these mechanisms affect marine CH4 and N2O cycling requires a combination of traditional scientific disciplines including oceanography, microbiology, and numerical modeling. Fundamental to these efforts is ensuring that the datasets produced by independent scientists are comparable and interoperable. Equally critical is transparent communication within the research community about the technical improvements required to increase our collective understanding of marine CH4 and N2O. A workshop sponsored by Ocean Carbon and Biogeochemistry (OCB) was organized to enhance dialogue and collaborations pertaining to marine CH4 and N2O. Here, we summarize the outcomes from the workshop to describe the challenges and opportunities for near-future CH4 and N2O research in the marine environment.
    Description: This article was an outcome of a workshop organized by the Ocean Carbon and Biogeochemistry (OCB) project office, which is supported by the US National Science Foundation (grant no. 1558412) and the National Aeronautics and Space Administration (grant no. NNX17AB17G). The workshop received additional funding from the Scientific Committee on Ocean Research (SCOR) which receives funding from the US National Science Foundation (grant no. 1840868) and contributions by additional national SCOR committees. The Chilean COPAS N2O time-series measurements were supported by Agencia Nacional de Investigación y Desarrollo (grant no. 1200861).
    Repository Name: Woods Hole Open Access Server
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  • 2
    Publication Date: 2022-05-27
    Description: © The Author(s), 2021. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Diaz, M. A., Gardner, C. B., Welch, S. A., Jackson, W. A., Adams, B. J., Wall, D. H., Hogg, I. D., Fierer, N., & Lyons, W. B. Geochemical zones and environmental gradients for soils from the central Transantarctic Mountains, Antarctica. Biogeosciences, 18(5), (2021): 1629-1644. https://doi.org/10.5194/bg-18-1629-2021.
    Description: Previous studies have established links between biodiversity and soil geochemistry in the McMurdo Dry Valleys, Antarctica, where environmental gradients are important determinants of soil biodiversity. However, these gradients are not well established in the central Transantarctic Mountains, which are thought to represent some of the least hospitable Antarctic soils. We analyzed 220 samples from 11 ice-free areas along the Shackleton Glacier (∼ 85∘ S), a major outlet glacier of the East Antarctic Ice Sheet. We established three zones of distinct geochemical gradients near the head of the glacier (upper), its central part (middle), and at the mouth (lower). The upper zone had the highest water-soluble salt concentrations with total salt concentrations exceeding 80 000 µg g−1, while the lower zone had the lowest water-soluble N:P ratios, suggesting that, in addition to other parameters (such as proximity to water and/or ice), the lower zone likely represents the most favorable ecological habitats. Given the strong dependence of geochemistry on geographic parameters, we developed multiple linear regression and random forest models to predict soil geochemical trends given latitude, longitude, elevation, distance from the coast, distance from the glacier, and soil moisture (variables which can be inferred from remote measurements). Confidence in our random forest model predictions was moderately high with R2 values for total water-soluble salts, water-soluble N:P, ClO−4, and ClO−3 of 0.81, 0.88, 0.78, and 0.74, respectively. These modeling results can be used to predict geochemical gradients and estimate salt concentrations for other Transantarctic Mountain soils, information that can ultimately be used to better predict distributions of soil biota in this remote region.
    Description: This research has been supported by the National Science Foundation (grant nos. OPP 1341631, GRFP 60041697, OPP 1341618, OPP 1341629, and OPP 1341736).
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  • 3
    Publication Date: 2022-10-26
    Description: © The Author(s), 2021. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Gu, S., Liu, Z., Oppo, D. W., Lynch-Stieglitz, J., Jahn, A., Zhang, J., Lindsay, K., & Wu, L. Remineralization dominating the δ13 C decrease in the mid-depth Atlantic during the last deglaciation. Earth and Planetary Science Letters, 571, (2021): 117106, https://doi.org/10.1016/j.epsl.2021.117106.
    Description: δ 13 C records from the mid-depth Atlantic show a pronounced decrease during the Heinrich Stadial 1 (HS1), a deglacial episode of dramatically weakened Atlantic Meridional Ocean Circulation (AMOC). Proposed explanations for this mid-depth decrease include a greater fraction of δ 13 C -depleted southern sourced water (SSW), a δ 13 C decrease in the North Atlantic Deep Water (NADW) end-member, and accumulation of the respired organic carbon. However, the relative importance of these proposed mechanisms cannot be quantitatively constrained from current available observations alone. Here we diagnose the individual contributions to the deglacial Atlantic mid-depth δ 13 C change from these mechanisms using a transient simulation with carbon isotopes and idealized tracers. We find that although the fraction of the low- δ 13 C SSW increases in response to a weaker AMOC during HS1, the water mass mixture change only plays a minor role in the mid-depth Atlantic δ 13 C decrease. Instead, increased remineralization due to the AMOC-induced mid-depth ocean ventilation decrease is the dominant cause. In this study, we differentiate between the deep end-members, which are assigned to deep water regions used in previous paleoceanography studies, and the surface end-members, which are from the near-surface water defined from the physical origin of deep water masses. We find that the deep NADW end-member includes additional remineralized material accumulated when sinking from the surface (surface NADW end-member). Therefore, the surface end-members should be used in diagnosing mechanisms of changes. Furthermore, our results suggest that remineralization in the surface end-member is more critical than the remineralization along the transport pathway from the near-surface formation region to the deep ocean, especially during the early deglaciation.
    Description: This work is supported by US National Science Foundation (NSF) P2C2 projects (1401778, 1401802, and 1566432), and the National Science Foundation of China No. 41630527. S.G. is supported by Shanghai Pujiang program.
    Keywords: δ13 C ; Water mass composition ; Remineralization ; End-member ; HS1
    Repository Name: Woods Hole Open Access Server
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  • 4
    Publication Date: 2022-05-27
    Description: © The Author(s), 2021. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Kim, H. H., Luo, Y.-W., Ducklow, H. W., Schofield, O. M., Steinberg, D. K., & Doney, S. C. WAP-1D-VAR v1.0: development and evaluation of a one-dimensional variational data assimilation model for the marine ecosystem along the West Antarctic Peninsula. Geoscientific Model Development, 14(8), (2021): 4939–4975, https://doi.org/10.5194/gmd-14-4939-2021.
    Description: The West Antarctic Peninsula (WAP) is a rapidly warming region, with substantial ecological and biogeochemical responses to the observed change and variability for the past decades, revealed by multi-decadal observations from the Palmer Antarctica Long-Term Ecological Research (LTER) program. The wealth of these long-term observations provides an important resource for ecosystem modeling, but there has been a lack of focus on the development of numerical models that simulate time-evolving plankton dynamics over the austral growth season along the coastal WAP. Here, we introduce a one-dimensional variational data assimilation planktonic ecosystem model (i.e., the WAP-1D-VAR v1.0 model) equipped with a model parameter optimization scheme. We first demonstrate the modified and newly added model schemes to the pre-existing food web and biogeochemical components of the other ecosystem models that WAP-1D-VAR model was adapted from, including diagnostic sea-ice forcing and trophic interactions specific to the WAP region. We then present the results from model experiments where we assimilate 11 different data types from an example Palmer LTER growth season (October 2002–March 2003) directly related to corresponding model state variables and flows between these variables. The iterative data assimilation procedure reduces the misfits between observations and model results by 58 %, compared to before optimization, via an optimized set of 12 parameters out of a total of 72 free parameters. The optimized model results capture key WAP ecological features, such as blooms during seasonal sea-ice retreat, the lack of macronutrient limitation, and modeled variables and flows comparable to other studies in the WAP region, as well as several important ecosystem metrics. One exception is that the model slightly underestimates particle export flux, for which we discuss potential underlying reasons. The data assimilation scheme of the WAP-1D-VAR model enables the available observational data to constrain previously poorly understood processes, including the partitioning of primary production by different phytoplankton groups, the optimal chlorophyll-to-carbon ratio of the WAP phytoplankton community, and the partitioning of dissolved organic carbon pools with different lability. The WAP-1D-VAR model can be successfully employed to link the snapshots collected by the available data sets together to explain and understand the observed dynamics along the coastal WAP.
    Description: Hyewon Heather Kim and Scott C. Doney were supported by the National Aeronautics and Space Administration Ocean Biology and Biogeochemistry Program (grant no. NNX14AL86G) and the US National Science Foundation Office of Polar Programs (grant no. PLR-1440435 to Hugh W. Ducklow at Columbia University; Palmer LTER). Hyewon Heather Kim was additionally supported by the Investment in Science Fund and the Reuben F. and Elizabeth B. Richards Endowed Fund from Woods Hole Oceanographic Institution. Oscar M. Schofield and Deborah K. Steinberg were supported by US NSF grant no. PLR-1440435. Ya-Wei Luo was supported by National Natural Science Foundation of China project no. 41890802.
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  • 5
    Publication Date: 2022-05-27
    Description: © The Author(s), 2021. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Almayrac, M. G., Broadley, M. W., Bekaert, D. V., Hofmann, A., & Marty, B. Possible discontinuous evolution of atmospheric xenon suggested by Archean barites. Chemical Geology, 581, (2021): 120405, https://doi.org/10.1016/j.chemgeo.2021.120405.
    Description: The Earth's atmosphere has continually evolved since its formation through interactions with the mantle as well as through loss of volatile species to space. Atmospheric xenon isotopes show a unique and progressive evolution during the Archean that stopped around the Archean-Proterozoic transition. The Xe isotope composition of the early atmosphere has been previously documented through the analysis of fluid inclusions trapped within quartz and barite. Whether this evolution was continuous or not is unclear, requiring additional analyses of ancient samples, which may potentially retain remnants of the ancient atmosphere. Here we present new argon, krypton and xenon isotopic data from a suite of Archean and Proterozoic barites ranging in age from 3.5 to 1.8 Ga, with the goal of providing further insights in to the evolution of atmospheric Xe, whilst also outlining the potential complications that can arise when using barites as a record of past atmospheres. Xenon released by low temperature pyrolysis and crushing of two samples which presumably formed around 2.8 and 2.6 Ga show Xe isotope mass dependent fractionation (MDF) of 11‰.u−1 and 3.4‰.u−1, respectively, relative to modern atmosphere. If trapped Xe is contemporaneous with the respective formation age, the significant difference in the degree of fractionation between the two samples provides supporting evidence for a plateau in the MDF-Xe evolution between 3.3 Ga and 2.8 Ga, followed by a rapid evolution at 2.8–2.6 Ga. This sharp decrease in MDF-Xe degree suggests the potential for a discontinuous temporal evolution of atmospheric Xe isotopes, which could have far reaching implications regarding current physical models of the early evolution of the Earth's atmosphere.
    Description: This work was funded by the ERC grant No. 695618 to B.M. We thank the S.A.R.M for providing elemental bulk analyses of the barites. We thank Laurent Zimmerman for technical mentorship and assistance.
    Keywords: Archean barite ; Noble gases ; Xenon anomalies ; Archean atmosphere
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  • 6
    Publication Date: 2022-05-27
    Description: © The Author(s), 2021. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Wehmiller, J. F., Brothers, L. L., Ramsey, K. W., Foster, D. S., Mattheus, C. R., Hein, C. J., & Shawler, J. L. Molluscan aminostratigraphy of the US Mid-Atlantic Quaternary coastal system: implications for onshore-offshore correlation, paleochannel and barrier island evolution, and local late Quaternary sea-level history. Quaternary Geochronology, 66, (2021): 101177, https://doi.org/10.1016/j.quageo.2021.101177.
    Description: The Quaternary record of the US Mid-Atlantic coastal system includes onshore emergent late Pleistocene shoreline deposits, offshore inner shelf and barrier island units, and paleovalleys formed during multiple glacial stage sea-level lowstands. The geochronology of this coastal system is based on uranium series, radiocarbon, amino acid racemization (AAR), and optically stimulated luminescence (OSL) methods. We report over 600 mollusk AAR results from 93 sites between northeastern North Carolina and the central New Jersey shelf, representing samples from both onshore cores or outcrops, sub-barrier and offshore cores, and transported shells from barrier island beaches. AAR age estimates are constrained by paired 14C analyses on specific shells and associated U-series coral ages from onshore sites. AAR data from offshore cores are interpreted in the context of detailed seismic stratigraphy. The distribution of Pleistocene-age shells on the island beaches is linked to the distribution of inner shelf or sub-barrier source units. Age mixing over a range of time-scales (~1 ka to ~100 ka) is identified by AAR results from onshore, beach, and shelf collections, often contributing insights into the processes forming individual barrier islands. The regional aminostratigraphic framework identifies a widespread late Pleistocene (Marine Isotope Stage 5) aminozone, with isolated records of middle and early Pleistocene deposition. AAR results provide age estimates for the timing of formation of the three major paleochannels that underlie the Delmarva Peninsula: Persimmon Point paleochannel ≥800 ka; Exmore paleochannel ~400–500 ka (MIS 12); and Eastville paleochannel 〉 125 ka (MIS 6). The results demonstrate the value of synthesizing abundant AAR chronologic data across various coastal environments, integrating multiple distinct geologic studies. The ages and elevations of the Quaternary units are important for current hypotheses about relative sea-level history and crustal dynamics in the region, which was likely influenced by the Laurentide ice sheet, the margin just ~400 km to the north.
    Description: This project was funded through a cooperative agreement with the Bureau of Ocean Energy Management of the U.S. Department of the Interior, Offshore Sand Resources for Coastal Resilience and Restoration Planning: M14AC00003 and M16AC00001. We thank J. Waldner (BOEM) for support and encouragement during this project. We also thank S. Howard and K. Luciano, South Carolina Geological Survey, and numerous colleagues in both the Mid-Atlantic and Southeast Atlantic BOEM ASAP projects, active from 2015 through 2019. This paper is contribution #3999 of the Virginia Institute of Marine Science, William & Mary. Partial support was also provided to Hein by the Mid-Atlantic Sea Grant program (NOAA) award numbers R/71856G and R/71856H and a Virginia Sea Grant (NOAA) Fellowship award NA18OAR4170083 supported Shawler. JFW acknowledges support from the University of Delaware Retired Faculty Research Program.
    Keywords: Quaternary sea-level ; Delmarva peninsula ; US Mid-Atlantic shelf ; Paleovalley ; Amino acid racemization ; Geochronology ; Age-mixing ; Seismic stratigraphy ; Mollusks
    Repository Name: Woods Hole Open Access Server
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  • 7
    Publication Date: 2022-05-27
    Description: © The Author(s), 2022. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Leech, V., Hazel, J. W., Gatlin, J. C., Lindsay, A. E., & Manhart, A. Mathematical modeling accurately predicts the dynamics and scaling of nuclear growth in discrete cytoplasmic volumes. Journal of Theoretical Biology, 533, (2022): 110936, https://doi.org/10.1016/j.jtbi.2021.110936.
    Description: Scaling of nuclear size with cell size has been observed in many species and cell types. In this work we formulate a modeling framework based on the limiting component hypothesis. We derive a family of spatio-temporal mathematical models for nuclear size determination based on different transport and growth mechanisms. We analyse model properties and use in vitro experimental data to identify the most probable mechanism. This suggests that nuclear volume scales with cell volume and that a nucleus controls its import rate as it grows. We further test the model by comparing to data of early frog development, where rapid cell divisions set the relevant time scales.
    Keywords: Nuclear Growth ; Partial differential equations ; Free boundary problems
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  • 8
    Publication Date: 2022-05-27
    Description: © The Author(s), 2021. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Chu, H., Luo, X., Ouyang, Z., Chan, W. S., Dengel, S., Biraud, S. C., Torn, M. S., Metzger, S., Kumar, J., Arain, M. A., Arkebauer, T. J., Baldocchi, D., Bernacchi, C., Billesbach, D., Black, T. A., Blanken, P. D., Bohrer, G., Bracho, R., Brown, S., Brunsell, N. A., Chen, J., Chen, X., Clark, K., Desai, A. R., Duman, T., Durden, D., Fares, S., Forbrich, I., Gamon, J. A., Gough, C. M., Griffis, T., Helbig, M., Hollinger, D., Humphreys, E., Ikawa, H., Iwata, H., Ju, Y., Knowles, J. F., Knox, S. H., Kobayashi, H., Kolb, T., Law, B., Lee, X., Litvak, M., Liu, H., Munger, J. W., Noormets, A., Novick, K., Oberbauer, S. F., Oechel, W., Oikawa, P., Papuga, S. A., Pendall, E., Prajapati, P., Prueger, J., Quinton, W. L., Richardson, A. D., Russell, E. S., Scott, R. L., Starr, G., Staebler, R., Stoy, P. C., Stuart-Haentjens, E., Sonnentag, O., Sullivan, R. C., Suyker, A., Ueyama, M., Vargas, R., Wood, J. D., & Zona, D. Representativeness of eddy-covariance flux footprints for areas surrounding AmeriFlux sites. Agricultural and Forest Meteorology, 301, (2021): 108350, https://doi.org/10.1016/j.agrformet.2021.108350.
    Description: Large datasets of greenhouse gas and energy surface-atmosphere fluxes measured with the eddy-covariance technique (e.g., FLUXNET2015, AmeriFlux BASE) are widely used to benchmark models and remote-sensing products. This study addresses one of the major challenges facing model-data integration: To what spatial extent do flux measurements taken at individual eddy-covariance sites reflect model- or satellite-based grid cells? We evaluate flux footprints—the temporally dynamic source areas that contribute to measured fluxes—and the representativeness of these footprints for target areas (e.g., within 250–3000 m radii around flux towers) that are often used in flux-data synthesis and modeling studies. We examine the land-cover composition and vegetation characteristics, represented here by the Enhanced Vegetation Index (EVI), in the flux footprints and target areas across 214 AmeriFlux sites, and evaluate potential biases as a consequence of the footprint-to-target-area mismatch. Monthly 80% footprint climatologies vary across sites and through time ranging four orders of magnitude from 103 to 107 m2 due to the measurement heights, underlying vegetation- and ground-surface characteristics, wind directions, and turbulent state of the atmosphere. Few eddy-covariance sites are located in a truly homogeneous landscape. Thus, the common model-data integration approaches that use a fixed-extent target area across sites introduce biases on the order of 4%–20% for EVI and 6%–20% for the dominant land cover percentage. These biases are site-specific functions of measurement heights, target area extents, and land-surface characteristics. We advocate that flux datasets need to be used with footprint awareness, especially in research and applications that benchmark against models and data products with explicit spatial information. We propose a simple representativeness index based on our evaluations that can be used as a guide to identify site-periods suitable for specific applications and to provide general guidance for data use.
    Description: We thank the AmeriFlux site teams for sharing their data and metadata with the network. Funding for these flux sites is acknowledged in the site data DOI, shown in Table S1. This analysis was supported in part by funding provided to the AmeriFlux Management Project by the U.S. Department of Energy's Office of Science under Contract No. DE-AC02-05CH11231. All footprint climatologies, site-level representativeness indices, and monthly EVI and sensor location biases can be accessed via the Zenodo Data Repository (Datasets S1–S6, http://doi.org/10.5281/zenodo.4015350).
    Keywords: Flux footprint ; Spatial representativeness ; Landsat EVI ; Land cover ; Sensor location bias ; Model-data benchmarking
    Repository Name: Woods Hole Open Access Server
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  • 9
    Publication Date: 2022-05-26
    Description: © The Author(s), 2021. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Celander, M. C., Goldstone, J. V., Brun, N. R., Clark, B., Jayaraman, S., Nacci, D., & Stegeman, J. J. Resistance to Cyp3a induction by polychlorinated biphenyls, including non-dioxin-like PCB153, in gills of killifish (Fundulus heteroclitus) from New Bedford Harbor. Environmental Toxicology and Pharmacology, 83, (2021): 103580. doi: 10.1016/j.etap.2020.103580.
    Description: Previous reports suggested that non-dioxin-like (NDL) PCB153 effects on cytochrome P450 3A (Cyp3a) expression in Atlantic killifish (Fundulus heteroclitus) gills differed between F0 generation fish from a PCB site (New Bedford Harbor; NBH) and a reference site (Scorton Creek; SC). Here, we examined effects of PCB153, dioxin-like (DL) PCB126, or a mixture of both, on Cyp3a56 mRNA in killifish generations removed from the wild, without environmental PCB exposures. PCB126 effects in liver and gills differed between populations, as expected. Gill Cyp3a56 was not affected by either congener in NBH F2 generation fish, but was induced by PCB153 in SC F1 fish, with females showing a greater response. PCB153 did not affect Cyp3a56 in liver of either population. Results suggest a heritable resistance to NDL-PCBs in killifish from NBH, in addition to that reported for DL PCBs. Induction of Cyp3a56 in gills may be a biomarker of exposure to NDL PCBs in fish populations that are not resistant to PCBs.
    Description: This study was supported by the sabbatical program from the Faculty of Science at the University of Gothenburg (MC), and by the Swiss National Science Foundation P2EZP2-165200 (NRB). The study was supported in part by the Superfund Hazardous Substances Research Program at Boston University NIH P42ES007381 (JVG, JJS). This research was also funded partly by the US Environmental Protection Agency (SJ, DN), including an appointment (BC) with the Postdoctoral Research Program at the US Environmental Protection (US-EPA) Office of Research and Development administered by the Oak Ridge Institute for Science and Education (ORISE), through Interagency Agreement No. DW92429801 between the US Department of Energy and the US-EPA. The contents do not reflect the views of the US-EPA, nor does mention of trade names or commercial products constitute endorsement or recommendation for use by the US-EPA. We thank Rene Francolini at the Woods Hole Oceanographic Institutions for excellent technical assistance and Dr. Sibel Karchner and Dr. Mark Hahn at the Woods Hole Oceanographic Institutions for valuable discussions and comments on the manuscript.
    Keywords: PCB153 ; Fish ; Gills ; Cyp3a ; Resistance
    Repository Name: Woods Hole Open Access Server
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  • 10
    Publication Date: 2022-10-26
    Description: © The Author(s), 2021. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Elling, F. J., Hemingway, J. D., Kharbush, J. J., Becker, K. W., Polik, C. A., & Pearson, A. Linking diatom-diazotroph symbioses to nitrogen cycle perturbations and deep-water anoxia: insights from Mediterranean sapropel events. Earth and Planetary Science Letters, 571, (2021): 117110, https://doi.org/10.1016/j.epsl.2021.117110.
    Description: Elevated organic matter (OM) export flux promotes marine anoxia, thus increasing carbon sequestration efficiency and decreasing atmospheric carbon dioxide levels. However, the mechanisms that trigger and sustain anoxic events—particularly those associated with nutrient-poor, oligotrophic surface waters—remain poorly constrained. Mediterranean Sea sapropels are well-preserved sediments deposited during episodic anoxic events throughout the Plio-Pleistocene; as such, they may provide unique insight into the biogeochemical and ecological drivers of—and responses to—marine anoxia. Using biomarker distributions, we demonstrate that anaerobic ammonium oxidizing (anammox) bacteria and diazotrophic endosymbionts of mat- and/or raft-forming diatoms were both abundant during sapropel events, particularly in the Ionian and Libyan seas. In these sapropels, the carbon isotope compositions of anammox biomarkers directly capture progressive 13C-depletion in deep-water dissolved inorganic carbon, indicating sustained carbon sequestration. To explain these observations, we propose a reinforcing feedback whereby initial nutrient and/or circulation perturbations promote fixed nitrogen loss via intensified anammox and heterotrophic denitrification, which in turn favors proliferation of rapidly sinking diatom-diazotroph symbiotic consortia, increases OM burial flux, and sustains anoxia. This mechanism resolves the long-standing conundrum that small and buoyant diazotrophs are apparently associated with high OM export during periods of marine anoxia and oligotrophy.
    Description: This work was funded through the Gordon and Betty Moore Foundation and US National Science Foundation grants 1843285, 1702262 and 1349126 (to A.P.). Additional financial support was provided by the Postdoctoral Program at the Woods Hole Oceanographic Institution and U.S. Geological Survey (K.W.B.).
    Keywords: Nammox ; Anoxia ; Compound-specific, δ13, C ; Diatom-diazotroph symbioses ; N2-fixation ; Mediterranean sapropels
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