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      Bridging Food Webs, Ecosystem Metabolism, and Biogeochemistry Using Ecological Stoichiometry Theory

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          Abstract

          Although aquatic ecologists and biogeochemists are well aware of the crucial importance of ecosystem functions, i.e., how biota drive biogeochemical processes and vice-versa, linking these fields in conceptual models is still uncommon. Attempts to explain the variability in elemental cycling consequently miss an important biological component and thereby impede a comprehensive understanding of the underlying processes governing energy and matter flow and transformation. The fate of multiple chemical elements in ecosystems is strongly linked by biotic demand and uptake; thus, considering elemental stoichiometry is important for both biogeochemical and ecological research. Nonetheless, assessments of ecological stoichiometry (ES) often focus on the elemental content of biota rather than taking a more holistic view by examining both elemental pools and fluxes (e.g., organismal stoichiometry and ecosystem process rates). ES theory holds the promise to be a unifying concept to link across hierarchical scales of patterns and processes in ecology, but this has not been fully achieved. Therefore, we propose connecting the expertise of aquatic ecologists and biogeochemists with ES theory as a common currency to connect food webs, ecosystem metabolism, and biogeochemistry, as they are inherently concatenated by the transfer of carbon, nitrogen, and phosphorous through biotic and abiotic nutrient transformation and fluxes. Several new studies exist that demonstrate the connections between food web ecology, biogeochemistry, and ecosystem metabolism. In addition to a general introduction into the topic, this paper presents examples of how these fields can be combined with a focus on ES. In this review, a series of concepts have guided the discussion: (1) changing biogeochemistry affects trophic interactions and ecosystem processes by altering the elemental ratios of key species and assemblages; (2) changing trophic dynamics influences the transformation and fluxes of matter across environmental boundaries; (3) changing ecosystem metabolism will alter the chemical diversity of the non-living environment. Finally, we propose that using ES to link nutrient cycling, trophic dynamics, and ecosystem metabolism would allow for a more holistic understanding of ecosystem functions in a changing environment.

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          Most cited references 128

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          Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater, marine and terrestrial ecosystems.

          The cycles of the key nutrient elements nitrogen (N) and phosphorus (P) have been massively altered by anthropogenic activities. Thus, it is essential to understand how photosynthetic production across diverse ecosystems is, or is not, limited by N and P. Via a large-scale meta-analysis of experimental enrichments, we show that P limitation is equally strong across these major habitats and that N and P limitation are equivalent within both terrestrial and freshwater systems. Furthermore, simultaneous N and P enrichment produces strongly positive synergistic responses in all three environments. Thus, contrary to some prevailing paradigms, freshwater, marine and terrestrial ecosystems are surprisingly similar in terms of N and P limitation.
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            Body size and metabolism

             Max Kleiber (1932)
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              Control of nitrogen export from watersheds by headwater streams.

              A comparative (15)N-tracer study of nitrogen dynamics in headwater streams from biomes throughout North America demonstrates that streams exert control over nutrient exports to rivers, lakes, and estuaries. The most rapid uptake and transformation of inorganic nitrogen occurred in the smallest streams. Ammonium entering these streams was removed from the water within a few tens to hundreds of meters. Nitrate was also removed from stream water but traveled a distance 5 to 10 times as long, on average, as ammonium. Despite low ammonium concentration in stream water, nitrification rates were high, indicating that small streams are potentially important sources of atmospheric nitrous oxide. During seasons of high biological activity, the reaches of headwater streams typically export downstream less than half of the input of dissolved inorganic nitrogen from their watersheds.
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                Author and article information

                Contributors
                Journal
                Front Microbiol
                Front Microbiol
                Front. Microbiol.
                Frontiers in Microbiology
                Frontiers Media S.A.
                1664-302X
                12 July 2017
                2017
                : 8
                Affiliations
                1Department of Environmental and Biological Sciences, University of Eastern Finland Kuopio, Finland
                2Agriculture and Food, Commonwealth Scientific and Industrial Research Organisation, Adelaide SA, Australia
                3Institute for Chemistry and Biology of the Marine Environment, University of Oldenburg Oldenburg, Germany
                4Stream Biofilm and Ecosystem Research, Ecole Polytechnique Fédérale de Lausanne Lausanne, Switzerland
                5Department of Ecology, Montana State University, Bozeman MT, United States
                6School of Freshwater Sciences, University of Wisconsin-Milwaukee, Milwaukee WI, United States
                7University of Maryland Center for Environmental Science, Cambridge MD, United States
                8The Hirst Lab, Organismal Biology, School of Biological and Chemical Sciences, Queen Mary University of London London, United Kingdom
                9Centre for Ocean Life, National Institute for Aquatic Resources, Technical University of Denmark Copenhagen, Denmark
                10Department of Evolution, Ecology, and Organismal Biology, Aquatic Ecology Laboratory, The Ohio State University, Columbus OH, United States
                11The Kominoski Lab, Department of Biological Sciences, Florida International University, Miami FL, United States
                12Department of Ecology and Evolutionary Biology, Cornell University, Ithaca NY, United States
                13Scripps Institution of Oceanography, University of California, San Diego, La Jolla CA, United States
                14Department of Biology, St. Catherine University, Minneapolis MN, United States
                15Helmholtz-Institute for Functional Marine Biodiversity Oldenburg, Germany
                Author notes

                Edited by: Robert Warner Sterner, University of Minnesota Duluth, United States

                Reviewed by: André Megali Amado, Federal University of Rio Grande do Norte, Brazil; Ian Salter, Alfred-Wegener-Institut für Polar- und Meeresforschung, Germany

                *Correspondence: Maren Striebel, maren.striebel@ 123456uni-oldenburg.de

                These authors have contributed equally to this work.

                This article was submitted to Aquatic Microbiology, a section of the journal Frontiers in Microbiology

                Article
                10.3389/fmicb.2017.01298
                5507128
                Copyright © 2017 Welti, Striebel, Ulseth, Cross, DeVilbiss, Glibert, Guo, Hirst, Hood, Kominoski, MacNeill, Mehring, Welter and Hillebrand.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                Page count
                Figures: 3, Tables: 1, Equations: 0, References: 139, Pages: 14, Words: 0
                Funding
                Funded by: Suomen Akatemia 10.13039/501100002341
                Award ID: 258875
                Funded by: Deutsche Forschungsgemeinschaft 10.13039/501100001659
                Award ID: STR 1383/1-1
                Funded by: Deutsche Forschungsgemeinschaft 10.13039/501100001659
                Award ID: DFG HI 848/11-2
                Categories
                Microbiology
                Review

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