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      Regulation of denitrification at the cellular level: a clue to the understanding of N 2O emissions from soils

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          Abstract

          Denitrifying prokaryotes use NO x as terminal electron acceptors in response to oxygen depletion. The process emits a mixture of NO, N 2O and N 2, depending on the relative activity of the enzymes catalysing the stepwise reduction of NO 3 to N 2O and finally to N 2. Cultured denitrifying prokaryotes show characteristic transient accumulation of NO 2 , NO and N 2O during transition from oxic to anoxic respiration, when tested under standardized conditions, but this character appears unrelated to phylogeny. Thus, although the denitrifying community of soils may differ in their propensity to emit N 2O, it may be difficult to predict such characteristics by analysis of the community composition. A common feature of strains tested in our laboratory is that the relative amounts of N 2O produced (N 2O/(N 2+N 2O) product ratio) is correlated with acidity, apparently owing to interference with the assembly of the enzyme N 2O reductase. The same phenomenon was demonstrated for soils and microbial communities extracted from soils. Liming could be a way to reduce N 2O emissions, but needs verification by field experiments. More sophisticated ways to reduce emissions may emerge in the future as we learn more about the regulation of denitrification at the cellular level.

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          Cell biology and molecular basis of denitrification.

          W Zumft (1997)
          Denitrification is a distinct means of energy conservation, making use of N oxides as terminal electron acceptors for cellular bioenergetics under anaerobic, microaerophilic, and occasionally aerobic conditions. The process is an essential branch of the global N cycle, reversing dinitrogen fixation, and is associated with chemolithotrophic, phototrophic, diazotrophic, or organotrophic metabolism but generally not with obligately anaerobic life. Discovered more than a century ago and believed to be exclusively a bacterial trait, denitrification has now been found in halophilic and hyperthermophilic archaea and in the mitochondria of fungi, raising evolutionarily intriguing vistas. Important advances in the biochemical characterization of denitrification and the underlying genetics have been achieved with Pseudomonas stutzeri, Pseudomonas aeruginosa, Paracoccus denitrificans, Ralstonia eutropha, and Rhodobacter sphaeroides. Pseudomonads represent one of the largest assemblies of the denitrifying bacteria within a single genus, favoring their use as model organisms. Around 50 genes are required within a single bacterium to encode the core structures of the denitrification apparatus. Much of the denitrification process of gram-negative bacteria has been found confined to the periplasm, whereas the topology and enzymology of the gram-positive bacteria are less well established. The activation and enzymatic transformation of N oxides is based on the redox chemistry of Fe, Cu, and Mo. Biochemical breakthroughs have included the X-ray structures of the two types of respiratory nitrite reductases and the isolation of the novel enzymes nitric oxide reductase and nitrous oxide reductase, as well as their structural characterization by indirect spectroscopic means. This revealed unexpected relationships among denitrification enzymes and respiratory oxygen reductases. Denitrification is intimately related to fundamental cellular processes that include primary and secondary transport, protein translocation, cytochrome c biogenesis, anaerobic gene regulation, metalloprotein assembly, and the biosynthesis of the cofactors molybdopterin and heme D1. An important class of regulators for the anaerobic expression of the denitrification apparatus are transcription factors of the greater FNR family. Nitrate and nitric oxide, in addition to being respiratory substrates, have been identified as signaling molecules for the induction of distinct N oxide-metabolizing enzymes.
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            Shifts in lake N:P stoichiometry and nutrient limitation driven by atmospheric nitrogen deposition.

            Human activities have more than doubled the amount of nitrogen (N) circulating in the biosphere. One major pathway of this anthropogenic N input into ecosystems has been increased regional deposition from the atmosphere. Here we show that atmospheric N deposition increased the stoichiometric ratio of N and phosphorus (P) in lakes in Norway, Sweden, and Colorado, United States, and, as a result, patterns of ecological nutrient limitation were shifted. Under low N deposition, phytoplankton growth is generally N-limited; however, in high-N deposition lakes, phytoplankton growth is consistently P-limited. Continued anthropogenic amplification of the global N cycle will further alter ecological processes, such as biogeochemical cycling, trophic dynamics, and biological diversity, in the world's lakes, even in lakes far from direct human disturbance.
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              On the fate of anthropogenic nitrogen.

              This article provides a synthesis of literature values to trace the fate of 150 Tg/yr anthropogenic nitrogen applied by humans to the Earth's land surface. Approximately 9 TgN/yr may be accumulating in the terrestrial biosphere in pools with residence times of ten to several hundred years. Enhanced fluvial transport of nitrogen in rivers and percolation to groundwater accounts for approximately 35 and 15 TgN/yr, respectively. Greater denitrification in terrestrial soils and wetlands may account for the loss of approximately 17 TgN/yr from the land surface, calculated by a compilation of data on the fraction of N(2)O emitted to the atmosphere and the current global rise of this gas in the atmosphere. A recent estimate of atmospheric transport of reactive nitrogen from land to sea (NO(x) and NH(x)) accounts for 48 TgN/yr. The total of these enhanced sinks, 124 TgN/yr, is less than the human-enhanced inputs to the land surface, indicating areas of needed additional attention to global nitrogen biogeochemistry. Policy makers should focus on increasing nitrogen-use efficiency in fertilization, reducing transport of reactive N to rivers and groundwater, and maximizing denitrification to its N(2) endproduct.
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                Author and article information

                Journal
                Philos Trans R Soc Lond B Biol Sci
                Philos. Trans. R. Soc. Lond., B, Biol. Sci
                RSTB
                royptb
                Philosophical Transactions of the Royal Society B: Biological Sciences
                The Royal Society
                0962-8436
                1471-2970
                5 May 2012
                5 May 2012
                : 367
                : 1593 , Theo Murphy Meeting Issue 'Nitrous oxide: the forgotten greenhouse gas' organized and edited by A. J. Thomson, E. Baggs and D. J. Richardson
                : 1226-1234
                Affiliations
                [1 ]Department of Plant and Environmental Sciences, Norwegian University of Life Sciences , PO Box 5003, 1432 Aas, Norway
                [2 ]Department of Chemistry, Biotechnology and Food Sciences, Norwegian University of Life Sciences , PO Box 5003, 1432 Aas, Norway
                Author notes
                [* ]Author for correspondence ( lars.bakken@ 123456umb.no ).

                One contribution of 12 to a Theo Murphy Meeting Issue ‘ Nitrous oxide: the forgotten greenhouse gas’.

                Article
                rstb20110321
                10.1098/rstb.2011.0321
                3306626
                22451108
                9c6537cc-775e-4b28-84d4-95bff578b8e8
                This journal is © 2012 The Royal Society

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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                15
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                69
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                Research Article

                Philosophy of science
                physiology,nitrous oxide,mitigation,soil,regulation
                Philosophy of science
                physiology, nitrous oxide, mitigation, soil, regulation

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