<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing DTD v3.0 20080202//EN" "https://jats.nlm.nih.gov/nlm-dtd/publishing/3.0/journalpublishing3.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" article-type="research-article" dtd-version="3.0" xml:lang="en">
<front>
<journal-meta>
<journal-id journal-id-type="publisher">BG</journal-id>
<journal-title-group>
<journal-title>Biogeosciences</journal-title>
<abbrev-journal-title abbrev-type="publisher">BG</abbrev-journal-title>
<abbrev-journal-title abbrev-type="nlm-ta">Biogeosciences</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1726-4189</issn>
<publisher><publisher-name>Copernicus Publications</publisher-name>
<publisher-loc>Göttingen, Germany</publisher-loc>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.5194/bg-4-353-2007</article-id>
<title-group>
<article-title>Biotic stoichiometric controls on the deep ocean N:P ratio</article-title>
</title-group>
<contrib-group><contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Lenton</surname>
<given-names>T. M.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Klausmeier</surname>
<given-names>C. A.</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
</contrib>
</contrib-group><aff id="aff1">
<label>1</label>
<addr-line>School of Environmental Sciences, University of East Anglia, Norwich NR4 7TJ, UK</addr-line>
</aff>
<aff id="aff2">
<label>2</label>
<addr-line>W. K. Kellogg Biological Station, Michigan State University, Hickory Corners, MI 49060, USA</addr-line>
</aff>
<pub-date pub-type="epub">
<day>20</day>
<month>06</month>
<year>2007</year>
</pub-date>
<volume>4</volume>
<issue>3</issue>
<fpage>353</fpage>
<lpage>367</lpage>
<permissions>
<copyright-statement>Copyright: &#x000a9; 2007 T. M. Lenton</copyright-statement>
<copyright-year>2007</copyright-year>
<license license-type="open-access">
<license-p>This work is licensed under the Creative Commons Attribution-NonCommercial-ShareAlike 2.5 Generic License. To view a copy of this licence, visit <ext-link ext-link-type="uri"  xlink:href="https://creativecommons.org/licenses/by-nc-sa/2.5/">https://creativecommons.org/licenses/by-nc-sa/2.5/</ext-link></license-p>
</license>
</permissions>
<self-uri xlink:href="https://bg.copernicus.org/articles/4/353/2007/bg-4-353-2007.html">This article is available from https://bg.copernicus.org/articles/4/353/2007/bg-4-353-2007.html</self-uri>
<self-uri xlink:href="https://bg.copernicus.org/articles/4/353/2007/bg-4-353-2007.pdf">The full text article is available as a PDF file from https://bg.copernicus.org/articles/4/353/2007/bg-4-353-2007.pdf</self-uri>
<abstract>
<p>We re-examine what controls the deep ocean N:P ratio in the light of
recent findings that the C:N:P stoichiometry of phytoplankton varies
with growth rate, nutrient and light limitation, species and phylum,
and that N&lt;sub&gt;2&lt;/sub&gt;-fixation may be limited by Fe, temperature and/or
light in large parts of the world ocean. In particular, we assess
whether a systematic change in phytoplankton stoichiometry can alter
the deep ocean N:P ratio. To do this we adapt recent models to
include non-Redfieldian stoichiometry of phytoplankton and
restriction of N&lt;sub&gt;2&lt;/sub&gt;-fixers to a fraction of the surface ocean. We
show that a systematic change in phytoplankton C:N:P can alter the
concentrations of NO&lt;sub&gt;3&lt;/sub&gt; and PO&lt;sub&gt;4&lt;/sub&gt; in the deep ocean but cannot
greatly alter their ratio, unless it also alters the N:P threshold
for N&lt;sub&gt;2&lt;/sub&gt;-fixation. This occurs if competitive dynamics set the
N:P threshold for N&lt;sub&gt;2&lt;/sub&gt;-fixation, in which case it remains close
to the N:P requirement of non-fixers (rather than that of
N&lt;sub&gt;2&lt;/sub&gt;-fixers) and consequently so does the deep ocean N:P ratio.
Then, even if N&lt;sub&gt;2&lt;/sub&gt;-fixers are restricted to a fraction of the
surface ocean, they reach higher densities there, minimising
variations in deep ocean N:P. Theoretical limits on the N:P
requirements of phytoplankton suggest that whilst the deep ocean has
been well oxygenated (i.e. since the Neoproterozoic, with the
possible exception of Oceanic Anoxic Events), its N:P ratio is
unlikely to have varied by more than a factor of two in either
direction. Within these bounds, evolutionary changes in
phytoplankton composition, and increased phosphorus weathering due
to the biological colonisation of the land surface, are predicted to
have driven long-term changes in ocean composition.</p>
</abstract>
<counts><page-count count="15"/></counts>
</article-meta>
</front>
<body/>
<back>
<ref-list>
<title>References</title>
<ref id="ref1">
<label>1</label><mixed-citation publication-type="other" xlink:type="simple"> Anbar, A. D. and Knoll, A. H.: Proterozoic Ocean Chemistry and Evolution: A Bioinorganic Bridge?, Science, 297, 1137&amp;ndash;1142, 2002. </mixed-citation>
</ref>
<ref id="ref2">
<label>2</label><mixed-citation publication-type="other" xlink:type="simple"> Anderson, L. A. and Sarmiento, J. L.: Redfield ratios of remineralization determined by nutrient data analysis, Global Biogeochem. Cy., 8, 65&amp;ndash;80, 1994. </mixed-citation>
</ref>
<ref id="ref3">
<label>3</label><mixed-citation publication-type="other" xlink:type="simple"> Arrigo, K. R.: Marine microorganisms and global nutrient cycles, Nature, 437, 349&amp;ndash;355, 2005. </mixed-citation>
</ref>
<ref id="ref4">
<label>4</label><mixed-citation publication-type="other" xlink:type="simple"> Bergman, N. M., Lenton, T. M., and Watson, A. J.: COPSE: a new model of biogeochemical cycling over Phanerozoic time, Am. J. Sci., 304, 397&amp;ndash;437, 2004. </mixed-citation>
</ref>
<ref id="ref5">
<label>5</label><mixed-citation publication-type="other" xlink:type="simple"> Berman-Frank, I., Cullen, J. T., Shaked, Y., Sherrell, R. M., and Falkowski, P. G.: Iron availability, cellular iron quotas, and nitrogen fixation in Trichodesmium, Limnol. Oceanogr., 46, 1249&amp;ndash;1260, 2001. </mixed-citation>
</ref>
<ref id="ref6">
<label>6</label><mixed-citation publication-type="other" xlink:type="simple"> Bjerrum, C. J. and Canfield, D. E.: Ocean productivity before about 1.9 Gyr ago limited by phosphorus adsorption onto iron oxides, Nature, 417, 159&amp;ndash;162, 2002. </mixed-citation>
</ref>
<ref id="ref7">
<label>7</label><mixed-citation publication-type="other" xlink:type="simple"> Breitbarth, E., Oeschlies, A., and LaRoche, J.: Physiological constraints on the global distribution of \it Trichodesmium &amp;ndash; effect of temperature on diazotrophy, Biogeosciences, 4, 53&amp;ndash;61, 2007. </mixed-citation>
</ref>
<ref id="ref8">
<label>8</label><mixed-citation publication-type="other" xlink:type="simple"> Broecker, W. S. and Peng, T.-H.: Tracers in the Sea, Eldigio Press, New York, 1982. </mixed-citation>
</ref>
<ref id="ref9">
<label>9</label><mixed-citation publication-type="other" xlink:type="simple"> Canfield, D. E.: A new model for Proterozoic ocean chemistry, Nature, 396, 450&amp;ndash;453, 1998. </mixed-citation>
</ref>
<ref id="ref10">
<label>10</label><mixed-citation publication-type="other" xlink:type="simple"> Codispoti, L. A.: Is the ocean losing nitrate?, Nature, 376, 724, 1995. </mixed-citation>
</ref>
<ref id="ref11">
<label>11</label><mixed-citation publication-type="other" xlink:type="simple"> Cullen, J. J.: Iron, nitrogen and phosphorus in the ocean, Nature, 402, 372, 1999. </mixed-citation>
</ref>
<ref id="ref12">
<label>12</label><mixed-citation publication-type="other" xlink:type="simple"> Dalsgaard, T., Canfield, D. E., Petersen, J., Thamdrup, B., and Acuna-Gonzalez, J.: N&lt;sub&gt;2&lt;/sub&gt; production by the anammox reaction in the anoxic water of Golfo Dulce, Costa Rica, Nature, 422, 606&amp;ndash;608, 2003. </mixed-citation>
</ref>
<ref id="ref13">
<label>13</label><mixed-citation publication-type="other" xlink:type="simple"> Deutsch, C., Sarmiento, J. L., Sigman, D. M., Gruber, N., and Dunne, J. P.: Spatial coupling of nitrogen inputs and losses in the ocean, Nature, 445, 163&amp;ndash;167, 2007. </mixed-citation>
</ref>
<ref id="ref14">
<label>14</label><mixed-citation publication-type="other" xlink:type="simple"> Falkowski, P. G.: Evolution of the nitrogen cycle and its influence on the biological sequestration of CO2 in the ocean, Nature, 387, 272&amp;ndash;275, 1997. </mixed-citation>
</ref>
<ref id="ref15">
<label>15</label><mixed-citation publication-type="other" xlink:type="simple"> Falkowski, P. G. and Davis, C. S.: Natural proportions, Nature, 431, 131, 2004. </mixed-citation>
</ref>
<ref id="ref16">
<label>16</label><mixed-citation publication-type="other" xlink:type="simple"> Fennel, K., Follows, M., and Falkowski, P. G.: The co-evolution of the nitrogen, carbon and oxygen cycles in the Proterozoic ocean, Am. J. Sci., 305, 526&amp;ndash;545, 2005. </mixed-citation>
</ref>
<ref id="ref17">
<label>17</label><mixed-citation publication-type="other" xlink:type="simple"> Geider, R. J. and La Roche, J.: Redfield revisited: variability of C:N:P in marine microalgae and its biochemical basis, Eur. J. Phycol., 37, 1&amp;ndash;17, 2002. </mixed-citation>
</ref>
<ref id="ref18">
<label>18</label><mixed-citation publication-type="other" xlink:type="simple"> Goldman, J. C.: On phytoplankton growth rates and particulate C:N:P ratios at low light, Limnol. Oceanogr., 31, 1358&amp;ndash;1361, 1986. </mixed-citation>
</ref>
<ref id="ref19">
<label>19</label><mixed-citation publication-type="other" xlink:type="simple"> Goldman, J. C., McCarthy, J. J., and Peavey, D. G.: Growth rate influence on the chemical composition of phytoplankton in oceanic waters, Nature, 279, 210&amp;ndash;215, 1979. </mixed-citation>
</ref>
<ref id="ref20">
<label>20</label><mixed-citation publication-type="other" xlink:type="simple"> Handoh, I. C. and Lenton, T. M.: Periodic mid-Cretaceous Oceanic Anoxic Events linked by oscillations of the phosphorus and oxygen biogeochemical cycles, Global Biogeochem. Cy., 17, 1092, https://doi.org/10.1029/2003GB002039, 2003. </mixed-citation>
</ref>
<ref id="ref21">
<label>21</label><mixed-citation publication-type="other" xlink:type="simple"> Hood, R. R., Coles, V. J., and Capone, D. G.: Modeling the distribution of Trichodesmium and nitrogen fixation in the Atlantic Ocean, J. Geophys. Res., 109, 1&amp;ndash;25, 2004. </mixed-citation>
</ref>
<ref id="ref22">
<label>22</label><mixed-citation publication-type="other" xlink:type="simple"> Karl, D. M., Letelier, R., Hebel, D. V., Bird, D. F., and Winn, C. D.: Trichodesmium blooms and new nitrogen in the North Pacific Gyre, in: Marine Pelagic Cyanobacteria: Trichodesmium and Other Diazotrophs, edited by: Carpenter, E. J., Kluwer Academic Publishers, pp 219&amp;ndash;237, 1992. </mixed-citation>
</ref>
<ref id="ref23">
<label>23</label><mixed-citation publication-type="other" xlink:type="simple"> Klausmeier, C. A., Litchman, E., Daufresne, T., and Levin, S. A.: Optimal nitrogen-to-phosphorus stoichiometry of phytoplankton, Nature, 429, 171&amp;ndash;174, 2004. </mixed-citation>
</ref>
<ref id="ref24">
<label>24</label><mixed-citation publication-type="other" xlink:type="simple"> Kuypers, M. M. M., Sliekers, A. O., Lavik, G., Schmid, M., Jorgensen, B. B., Kuenen, J. G., Sinninghe Damste, J. S., Strous, M., and Jetten, M. S. M.: Anaerobic ammonium oxidation by anammox bacteria in the Black Sea, Nature, 422, 608&amp;ndash;611, 2003. </mixed-citation>
</ref>
<ref id="ref25">
<label>25</label><mixed-citation publication-type="other" xlink:type="simple"> Kuypers, M. M. M., van Breugel, Y., Schouten, S., Erba, E., and Sinninghe Damste, J. S.: N&lt;sub&gt;2&lt;/sub&gt;-fixing cyanobacteria supplied nutrient N for Cretaceous oceanic anoxic events, Geology, 32, 853&amp;ndash;856, 2004. </mixed-citation>
</ref>
<ref id="ref26">
<label>26</label><mixed-citation publication-type="other" xlink:type="simple"> Lenton, T. M.: The role of land plants, phosphorus weathering and fire in the rise and regulation of atmospheric oxygen, Glob. Change Biol., 7, 613&amp;ndash;629, 2001. </mixed-citation>
</ref>
<ref id="ref27">
<label>27</label><mixed-citation publication-type="other" xlink:type="simple"> Lenton, T. M. and Watson, A. J.: Redfield revisited: 1. Regulation of nitrate, phosphate and oxygen in the ocean, Global Biogeochem. Cy., 14, 225&amp;ndash;248, 2000. </mixed-citation>
</ref>
<ref id="ref28">
<label>28</label><mixed-citation publication-type="other" xlink:type="simple"> Lenton, T. M. and Watson, A. J.: Biotic enhancement of weathering, atmospheric oxygen and carbon dioxide in the Neoproterozoic, Geophys. Res. Lett., 31, L05202, https://doi.org/10.1029/2003GL018802, 2004. </mixed-citation>
</ref>
<ref id="ref29">
<label>29</label><mixed-citation publication-type="other" xlink:type="simple"> Mills, M. M., Ridame, C., Davey, M., La Roche, J., and Geider, R. J.: Iron and phosphorus co-limit nitrogen fixation in the eastern tropical North Atlantic, Nature, 429, 292&amp;ndash;294, 2004. </mixed-citation>
</ref>
<ref id="ref30">
<label>30</label><mixed-citation publication-type="other" xlink:type="simple"> Piper, D. Z. and Codispoti, L. A.: Marine Phosphorite Deposits and the Nitrogen Cycle, Science, 188, 15&amp;ndash;18, 1975. </mixed-citation>
</ref>
<ref id="ref31">
<label>31</label><mixed-citation publication-type="other" xlink:type="simple"> Quigg, A., Finkel, Z. V., Irwin, A. J., Rosenthal, Y., Ho, T.-Y., Reinfelder, J. R., Schofield, O., Morel, F. M. M., and Falkowski, P. G.: The evolutionary inheritance of elemental stoichiometry in marine phytoplankton, Nature, 425, 291&amp;ndash;294, 2003. </mixed-citation>
</ref>
<ref id="ref32">
<label>32</label><mixed-citation publication-type="other" xlink:type="simple"> Redfield, A. C.: On the proportions of organic derivatives in sea water and their relation to the composition of plankton, James Johnstone Memorial Volume, University of Liverpool, Liverpool, pp 176&amp;ndash;192, 1934. </mixed-citation>
</ref>
<ref id="ref33">
<label>33</label><mixed-citation publication-type="other" xlink:type="simple"> Redfield, A. C.: The biological control of chemical factors in the environment, Am. Sci., 46, 205&amp;ndash;221, 1958. </mixed-citation>
</ref>
<ref id="ref34">
<label>34</label><mixed-citation publication-type="other" xlink:type="simple"> Ruttenberg, K. C.: Reassessment of the oceanic residence time of phosphorus, Chem. Geol., 107, 405&amp;ndash;409, 1993. </mixed-citation>
</ref>
<ref id="ref35">
<label>35</label><mixed-citation publication-type="other" xlink:type="simple"> Sa&amp;ntilde;udo-Wilhelmy, S. A., Kustka, A. B., Gobler, C. J., Hutchins, D. A., Yang, M., Lwiza, K., Burns, J., Capone, D. G., Raven, J. A., and Carpenter, E. J.: Phosphorus limitation of nitrogen fixation by \textitTrichodesmium in the central Atlantic Ocean, Nature, 411, 66&amp;ndash;69, 2001. </mixed-citation>
</ref>
<ref id="ref36">
<label>36</label><mixed-citation publication-type="other" xlink:type="simple"> Schade, J. D., Espelata, J. F., Klausmeier, C. A., McGroddy, M. E., Thomas, S. A., and Zhang, L.: A conceptual framework for ecosystem stoichiometry: balancing resource supply and demand, Oikos, 109, 40&amp;ndash;51, 2005. </mixed-citation>
</ref>
<ref id="ref37">
<label>37</label><mixed-citation publication-type="other" xlink:type="simple"> Schulz, H. N. and Schulz, H. D.: Large Sulfur Bacteria and the Formation of Phosphorite, Science, 307, 416&amp;ndash;418, 2005. </mixed-citation>
</ref>
<ref id="ref38">
<label>38</label><mixed-citation publication-type="other" xlink:type="simple"> Tyrrell, T.: The relative influences of nitrogen and phosphorus on oceanic primary production, Nature, 400, 525&amp;ndash;531, 1999. </mixed-citation>
</ref>
<ref id="ref39">
<label>39</label><mixed-citation publication-type="other" xlink:type="simple"> Van Cappellen, P. and Ingall, E. D.: Redox stabilisation of the atmosphere and oceans by phosphorus-limited marine productivity, Science, 271, 493&amp;ndash;496, 1996. </mixed-citation>
</ref>
<ref id="ref40">
<label>40</label><mixed-citation publication-type="other" xlink:type="simple"> Wallmann, K.: Feedbacks between oceanic redox states and marine productivity: A model perspective focused on benthic phosphorus cycling, Global Biogeochem. Cy., 17, 1084, https://doi.org/10.1029/2002GB001968, 2003. </mixed-citation>
</ref>
</ref-list>
</back>
</article>