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	<journal>
		<journal_title>Biogeosciences</journal_title>
		<journal_url>www.biogeosciences.net</journal_url>
		<issn>1726-4170</issn>
		<eissn>1726-4189</eissn>
		<volume_number>6</volume_number>
		<issue_number>5</issue_number>
		<publication_year>2009</publication_year>
	</journal>
	<doi>10.5194/bg-6-923-2009</doi>
	<article_url>http://www.biogeosciences.net/6/923/2009/</article_url>
	<abstract_html>http://www.biogeosciences.net/6/923/2009/bg-6-923-2009.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/6/923/2009/bg-6-923-2009.pdf</fulltext_pdf>
	<start_page>923</start_page>
	<end_page>935</end_page>
	<publication_date>2009-05-28</publication_date>
	<article_title content_type="html">Stoichiometries of remineralisation and denitrification in global biogeochemical ocean models</article_title>
	<authors>
		<author numeration="1" affiliations="1,2">
			<name>A. Paulmier</name>
			<email>apaulmier@ifm-geomar.de</email>
		</author>
		<author numeration="2" affiliations="1">
			<name>I. Kriest</name>
		</author>
		<author numeration="3" affiliations="1">
			<name>A. Oschlies</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Leibniz-Institute of Marine Sciences  IFM-GEOMAR, Düsternbrooker Weg 20, 24105 Kiel, Germany</affiliation>
		<affiliation numeration="2" content_type="html">currently at: Max Planck Institute for Marine Microbiology,  Celsiusstr. 1, 28359 Bremen, Germany</affiliation>
	</affiliations>
	<abstract content_type="html">Since the seminal paper of Redfield (1934), constant
stoichiometric elemental ratios linking biotic carbon  and nutrient
fluxes are often assumed in marine biogeochemistry, and especially in
coupled biogeochemical circulation models, to
couple the global oxygen,
carbon and nutrient cycles. However, when looking in more detail, some
deviations from the classical Redfield stoichiometry have been
reported, in particular with respect to remineralization of organic matter
changing with depth or with ambient oxygen levels.
We here compare the assumptions about the stoichiometry of organic
matter and its remineralization
that are used explicitly and implicitly in common biogeochemical
ocean models. We find that the implicit assumptions made about the
hydrogen content of organic matter can lead to
inconsistencies in
the modeled remineralization and denitrification stoichiometries.
It is suggested that
future marine biogeochemical
models explicitly
state the chemical composition
assumed for the organic matter, including its oxygen and hydrogen content.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Altabet, M. A., Francois, R., Murray, D. W., and Prell, W. L.: Climate-related variations in denitrification in the Arabian Sea from sediment $^15$N/$^14$N ratios, Nature, 373, 506–509, 1995. </reference>
		<reference numeration="2" content_type="text"> Anderson, L.: On the hydrogen and oxygen content of marine phytoplankton, Deep-Sea Res. I, 42, 1675–1680, 1995. </reference>
		<reference numeration="3" content_type="text"> Anderson, L. and Sarmiento, J.: Redfield ratios of remineralization determined by nutrient data analysis, Global Biogeochem. Cy., 8, 65–80, 1994. </reference>
		<reference numeration="4" content_type="text"> Aumont, O.: Etude du cycle naturel du carbone dans un modele 3D de l&apos;ocean mondial, Doctoral Thesis, Paris VI, 346~pp., 1998. </reference>
		<reference numeration="5" content_type="text"> Aumont, O. and Bopp, L.: Globalizing results from ocean in situ iron fertilization studies, Global Biogeochem. Cy., 20, GB2017, %30, \blackbox\bf article number or page range?, \doi10.1029/2005GB002591, 2006. </reference>
		<reference numeration="6" content_type="text"> Boulahdid, M. and Minster, J.-F.: Oxygen consumption and nutrient regeneration ratios along isopycnal horizons in the Pacific Ocean, Mar. Chem., 26, 133–153, 1989. </reference>
		<reference numeration="7" content_type="text"> Brewer, P. and Goldman, J.: Alkalinity changes generated by phytoplankton growth, Limnol. Oceanogr., 21, 108–117, 1976. </reference>
		<reference numeration="8" content_type="text"> Canfield, D.: Models of oxic respiration, denitrification and sulfate reduction in zones of coastal upwelling, Geochim. Cosmochim. Ac., 70, 5753–5765, 2006. </reference>
		<reference numeration="9" content_type="text"> Codispoti, L., Brandes, J., Christensen, J., Devol, A., Naqvi, S., Paerl, H., and Yoshinari, T.: The oceanic fixed nitrogen and nitrous oxide budgets: moving targets as we enter the anthropocene?, Sci. Mar., 65, 85–105, 2001. </reference>
		<reference numeration="10" content_type="text"> Copin-Montegut, C. and Copin-Montegut, G.: Stoichiometry of carbon, nitrogen, and phosphorous in marine particulate matter, Deep-Sea Res. I, 30, 31–46, 1983. </reference>
		<reference numeration="11" content_type="text"> Ganeshram, R. S., Pedersen, T. F., Calvert, S. E., and Fran\c cois, R.: Reduced nitrogen fixation in the glacial ocean inferred from changes in marine nitrogen and phosphorus inventories, Nature, 415, 156–159, 2002. </reference>
		<reference numeration="12" content_type="text"> Devol, A., Uhlenhopp, A., Naqvi, S., Brandes, J., Jayakumar, A., Naik, H., Gaurin, S., Codispoti, L., and Yoshinari, T.: Denitrification rates and excess nitrogen gas concentrations in the Arabian Sea oxygen deficient zone, Deep-Sea Res. I, 53, 1533–1547, 2006. </reference>
		<reference numeration="13" content_type="text"> Fraga, F., Rios, A., Perez, F., and Figueras, F.: Theoretical limits of oxygen:carbon and oxygen:nitrogen ratios during photosynthesis and mineralisation of organic matter in the sea, Mar. Chem., 62, 161–168, 1998. </reference>
		<reference numeration="14" content_type="text"> Fraga, F. and Alvarez-Salgado, X.: On the variation of alkalinity during phytoplankton photosynthesis, Cienc. Mar., 31, 627–639, 2005. </reference>
		<reference numeration="15" content_type="text"> Goldman, J. and Brewer, P.: Effect of nitrogen source and growth rate on phytoplankton-mediated changes in alkalinity, Limnol. Oceanogr., 25, 352–357, 1980. </reference>
		<reference numeration="16" content_type="text"> Hedges, J. I., Baldock, J. A., Gélinas, Y., Peterson, M. L., and Wakeham, S. G.: The biochemical and elemental compositions of marine plankton/a NMR perspective, Mar. Chem., 47, 47–63, 2002. </reference>
		<reference numeration="17" content_type="text"> Koeve, W.: Spring bloom carbon to nitrogen ratio of net community production in the temperate N. Atlantic, Deep-Sea Res., 51, 1579–1600, 2004. </reference>
		<reference numeration="18" content_type="text"> Lam, P., Lavik, G., Jensen, M. M., van de Vossenber, J., Schmidt, M., Woebken, D., Gutierrez, D., Amann, R., Jetten, M. S. M, and Kuypers, M. M.: Revising the nitrogen cycle in the Peruvian oxygen minimum zone, P. Natl. Acad. Sci., 106, 4752–4757, 2009. </reference>
		<reference numeration="19" content_type="text"> Laws, E. A.: Photosynthetic quotients, new production and net community production in the open ocean, Deep-Sea Res., 38, 143–167, 1991. </reference>
		<reference numeration="20" content_type="text"> Maier-Reimer, E.: Geochemical cycles in an ocean general circulation model – Preindustrial tracer distributions, Global Biogeochem. Cy., 7, 645–677, 1993. </reference>
		<reference numeration="21" content_type="text"> Maier-Reimer, E., Kriest, I., Segschneider, J., and Wetzel, P.: The HAMburg Ocean Carbon Cycle Model HAMOCC 5.1 – Technical Description Release 1.1, Reports on Earth System Science~14, Max-Planck-Institute for Meteorology, Hamburg, online available at prefixhttp://www.mpimet.mpg.de/fileadmin/publikationen/erdsystem_14.pdf, 2005. </reference>
		<reference numeration="22" content_type="text"> Meissner, K. J., Galbraith, E. D., and Völker, C.: Denitrification under glacial and interglacial conditions: A physical approach, Paleoceanography, 20, PA3001, doi:10.1029/2004PA001083, 2005. </reference>
		<reference numeration="23" content_type="text"> Moore, J., Doney, S., Kleypas, J., Glover, D., and Fung, I.: An intermediate complexity marine ecosystem model for the global domain, Deep-Sea Res. II, 49, 403–462, 2002. </reference>
		<reference numeration="24" content_type="text"> Moore, J., Doney, S., and Lindsay, K.: Upper ocean ecosystem dynamics and iron cycling in a global three-dimensional model, Global Biogeochem. Cy., 18, %CORRECTED: %BG4028, GB4028, \doi10.1029/2004GB002220, 2004. %\blackbox\bf article number or page range? </reference>
		<reference numeration="25" content_type="text"> Moore, J. and Doney, S.: Iron availability limits the ocean nitrogen inventory stabilizing feedbacks between marine denitrification and nitrogen fixationg, Global Biogeochem. Cy., 21, %CORRECTED: the article number is BG2001, and there is no page range %BG2001, GB2001, \doi10.1029/2006GB002762, 2007. %\blackbox\bf article number or page range? </reference>
		<reference numeration="26" content_type="text"> Oschlies, A., Schulz, K. G., Riebesell, U., and Schmittner, A.: Simulated 21st century&apos;s increase in oceanic suboxia by CO&lt;sub&gt;2&lt;/sub&gt;-enhanced biological carbon export, Global Biogeochem. Cy., 22, GB4008, \doi10.1029/2007GB003147, 2008. %in press, 2009.\blackbox\bf update? </reference>
		<reference numeration="27" content_type="text"> Paulmier, A. and Ruiz-Pino, D.: Oxygen minimum zones (OMZs) in the modern ocean, Prog. Oceanogr., 80(3-4), 113-128, \doi10.1029/j.pocean.2008.08.001, 2008 %doi:10,1016/j.pocean.2008,05,01, 2008. % \blackbox\bf volume and article number? </reference>
		<reference numeration="28" content_type="text"> Redfield, A.: On the proportions of organic derivatives in sea water and their relation to the composition of plamkton, in: James Johnstone Memorial Volume, Liverpool, 176–192, 1934. </reference>
		<reference numeration="29" content_type="text"> Redfield, A. C, Ketchum, B. H., and Richards, F. A.: The influence of organisms on the composition of sea-water, in: The Sea, edited by: Hill, M., vol 2, chap 2, Interscience, New York, USA, 26–77, 1963. </reference>
		<reference numeration="30" content_type="text"> Richards, F.: Anoxic basins and fjords, in: Chemical Oceanography, edited by: Riley, J. and Skirrow, G., vol 1, chap 13, Academic Press, New York, USA, 611–645, 1965. </reference>
		<reference numeration="31" content_type="text"> Sambrotto, R. N., Savidge, G., Robinson, C., Boyd, P., Takahashi, T., Karl, D. M., Langdon, C., Chipman, D., Marra, J., and Codispoti, L.: Elevated consumption of carbon relative to nitrogen in the surface ocean, Nature, 363, 248–250, 1993. </reference>
		<reference numeration="32" content_type="text"> Schartau, M. and Oschlies, A.: Simulataneous data-based optimization of a 1D-ecosystem model at three locations in the North Atlantic: Part I – Method and parameter estimates, J. Mar. Res., 61, 765–793, 2003. </reference>
		<reference numeration="33" content_type="text"> Schmittner, A., Oschlies, A., Matthews, H., and Galbraith, E.: Future changes in climate, ocean circulation, ecosystems, and biogeochemical cycling simulated for a business-as-usual CO&lt;sub&gt;2&lt;/sub&gt; emission scenario until year 4000 AD, Global Biogeochem. Cy., 22, GB1013, \doi10.1029/2007GB002953, 2008. %\blackbox\bf article number or page range? </reference>
		<reference numeration="34" content_type="text"> Schneider, B., Schlitzer, R., Fischer, G., and Nötig, E.-M.: Depth-dependent elemental composition of particulate organic matter (POM) in the ocean, Global Biogeochem. Cy., 17(2), 1032, doi:10.1029/2002GB001871, 2003. %\blackbox\bf article number or page range? </reference>
		<reference numeration="35" content_type="text"> Six, K D. and Maier-Reimer, E.: Effects of plankton dynamics on seasonal carbon fluxes in an ocean general circulation model, Global Biogeochem. Cy., 10, 559–583, 1996. </reference>
		<reference numeration="36" content_type="text"> Stramma, L., G. C. Johnson, J. Sprintall, and V. Mohrholz: Expanding oxygen-minimum zones in the tropical oceans, Science, 320, 655–658, 2008. </reference>
		<reference numeration="37" content_type="text"> Suntharalingam, P., Sarmiento, J., and Toggweiler, J.: Global significance of nitrous-oxide production and transport from oceanic low-oxygen zones – A modeling study, Global Biogeochem. Cy., 14, 1353–1370, 2000. </reference>
		<reference numeration="38" content_type="text"> Takahashi, T., Broecker, W., and Langer, S.: Redfield ratio based on chemical data from isopycnal surfaces, J. Geophys. Res., 90, 6907–6924, 1985. </reference>
		<reference numeration="39" content_type="text"> Van Mooy, B., Keil, R., and Devol, A.: Impact of suboxia on sinking particulate organic carbon: Enhanced carbon flux and preferential degradation of amino acids via denitrificiation, Geochim. Cosmochim. Ac., 66, 457–465, 2002. </reference>
		<reference numeration="40" content_type="text"> Weaver, A. J., Eby, M., Wiebe, E. C., Bitz, C. M., Duffy, P. B., Ewen, T. L., Fanning, A. F., Holland, M. M., MacFadyen, A., Matthews, H. D., Meissner, K. J., Saenko, O., Schmittner, A., Wang, H., and Yoshimori, M.: The UVic earth system climate model: Model description, climatology, and applications to past, present and future climates, Atmos.-Ocean, 39, 361–428, 2001. </reference>
		<reference numeration="41" content_type="text"> Wetzel, P.: Interannual and Decadal Variability in the Air-Sea Exchange of CO&lt;sub&gt;2&lt;/sub&gt; – a Model Study, Reports on Earth System Science~7, Max-Planck-Institute for Meteorology, Hamburg, online availabel at: prefixhttp://www.mpimet.mpg.de/fileadmin/publikationen/erdsystem_7.pdf, 2004. </reference>
		<reference numeration="42" content_type="text"> Wolf-Gladrow, D., Zeebe, R., Klaas, C., Körtzinger, A., and Dickson, A.: Total alkalinity: The explicit conservative expression and its application to biogeochemical processes, Mar. Chem., 106, 287–300, 2007. </reference>
	</references>
</article>

