<?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">NPG</journal-id>
<journal-title-group>
<journal-title>Nonlinear Processes in Geophysics</journal-title>
<abbrev-journal-title abbrev-type="publisher">NPG</abbrev-journal-title>
<abbrev-journal-title abbrev-type="nlm-ta">Nonlin. Processes Geophys.</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1607-7946</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/npg-18-29-2011</article-id>
<title-group>
<article-title>Interactions between marine biota and ENSO: a conceptual model analysis</article-title>
</title-group>
<contrib-group><contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Heinemann</surname>
<given-names>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>Timmermann</surname>
<given-names>A.</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Feudel</surname>
<given-names>U.</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
</contrib-group><aff id="aff1">
<label>1</label>
<addr-line>Max Planck Institute for Meteorology, Bundesstrasse 53, 20146 Hamburg, Germany</addr-line>
</aff>
<aff id="aff2">
<label>2</label>
<addr-line>IPRC and Dept. of Oceanography, SOEST, University of Hawaii, 1680 East-West Road, Honolulu, Hawaii 96822, USA</addr-line>
</aff>
<aff id="aff3">
<label>3</label>
<addr-line>ICBM, C. v. O. University of Oldenburg, Carl von Ossietzky Strasse 9–11, 26111 Oldenburg, Germany</addr-line>
</aff>
<pub-date pub-type="epub">
<day>21</day>
<month>01</month>
<year>2011</year>
</pub-date>
<volume>18</volume>
<issue>1</issue>
<fpage>29</fpage>
<lpage>40</lpage>
<permissions>
<copyright-statement>Copyright: &#x000a9; 2011 M. Heinemann et al.</copyright-statement>
<copyright-year>2011</copyright-year>
<license license-type="open-access">
<license-p>This work is licensed under the Creative Commons Attribution 3.0 Unported License. To view a copy of this licence, visit <ext-link ext-link-type="uri"  xlink:href="https://creativecommons.org/licenses/by/3.0/">https://creativecommons.org/licenses/by/3.0/</ext-link></license-p>
</license>
</permissions>
<self-uri xlink:href="https://npg.copernicus.org/articles/18/29/2011/npg-18-29-2011.html">This article is available from https://npg.copernicus.org/articles/18/29/2011/npg-18-29-2011.html</self-uri>
<self-uri xlink:href="https://npg.copernicus.org/articles/18/29/2011/npg-18-29-2011.pdf">The full text article is available as a PDF file from https://npg.copernicus.org/articles/18/29/2011/npg-18-29-2011.pdf</self-uri>
<abstract>
<p>We develop a conceptual coupled atmosphere-ocean-ecosystem model for the
tropical Pacific to investigate the interaction between marine biota and the
El Niño-Southern Oscillation (ENSO). Ocean and atmosphere are represented
by a two-box model for the equatorial Pacific cold tongue and the warm pool,
including a simplified mixed layer scheme. Marine biota are represented by a
three-component (nutrient, phytoplankton, and zooplankton) ecosystem model.
&lt;br&gt;&lt;br&gt;
The atmosphere-ocean model exhibits an oscillatory state which qualitatively
captures the main physics of ENSO. During an ENSO cycle, the variation of
nutrient upwelling, and, to a small extent, the variation of
photosynthetically available radiation force an ecosystem oscillation. The
simplified ecosystem in turn, due to the effect of phytoplankton on the
absorption of shortwave radiation in the water column, leads to (1) a warming
of the tropical Pacific, (2) a reduction of the ENSO amplitude, and
(3) a prolongation of the ENSO period. We qualitatively investigate these
bio-physical coupling mechanisms using continuation methods. It is
demonstrated that bio-physical coupling may play a considerable role in
modulating ENSO variability.</p>
</abstract>
<counts><page-count count="12"/></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">Anderson, W., Gnanadesikan, A., Hallberg, R., Dunne, J., and Samuels, B.&amp;nbsp;L.: Impact of ocean color on the maintenance of the Pacific Cold Tongue, Geophys. Res. Lett., 34, L11609, https://doi.org/10.1029/2007GL030100, 2007.</mixed-citation>
</ref>
<ref id="ref2">
<label>2</label><mixed-citation publication-type="other" xlink:type="simple">Anderson, W., Gnanadesikan, A., and Wittenberg, A.: Regional impacts of ocean color on tropical Pacific variability, Ocean Sci., 5, 313–327, https://doi.org/10.5194/os-5-313-2009, 2009.</mixed-citation>
</ref>
<ref id="ref3">
<label>3</label><mixed-citation publication-type="other" xlink:type="simple">Ballabrera-Poy, J., Murtugudde, R.&amp;nbsp;G., Zhang, R.-H., and Busalacchi, A.&amp;nbsp;J.: Coupled ocean-atmosphere response to seasonal modulation of ocean color: Impact on interannual climate simulations in the tropical Pacific, J. Climate, 20, 353–374, https://doi.org/10.1175/JCLI3958.1, 2007.</mixed-citation>
</ref>
<ref id="ref4">
<label>4</label><mixed-citation publication-type="other" xlink:type="simple">Barber, R.&amp;nbsp;T. and Chavez, F.&amp;nbsp;P.: Biological Consequences of El Niño, Science, 222, 1203–1210, https://doi.org/10.1126/science.222.4629.1203, 1983.</mixed-citation>
</ref>
<ref id="ref5">
<label>5</label><mixed-citation publication-type="other" xlink:type="simple">Bjerknes, J.: Atmospheric teleconnections from the equatorial Pacific, Mon. Weather Rev., 97, 163–172, 1969.</mixed-citation>
</ref>
<ref id="ref6">
<label>6</label><mixed-citation publication-type="other" xlink:type="simple">Bromwich, D., Monaghan, A., and Guo, Z.: Modeling the ENSO modulation of Antarctic climate in the late 1990s with Polar MM5, J. Climate, 17, 109–132, 2004.</mixed-citation>
</ref>
<ref id="ref7">
<label>7</label><mixed-citation publication-type="other" xlink:type="simple">Chavez, F.&amp;nbsp;P., Strutton, P.&amp;nbsp;G., Friederich, G.&amp;nbsp;E., Feely, R.&amp;nbsp;A., Feldman, G.&amp;nbsp;C., Foley, D.&amp;nbsp;G., and McPhaden, M.&amp;nbsp;J.: Biological and Chemical Response of the Equatorial Pacific Ocean to the 1997–98 El Niño, Science, 286, 2126–2131, https://doi.org/10.1126/science.286.5447.2126, 1999.</mixed-citation>
</ref>
<ref id="ref8">
<label>8</label><mixed-citation publication-type="other" xlink:type="simple">Cloern, J.&amp;nbsp;E., Grenz, C., and Vidergar-Lucas, L.: An empirical model of the phytoplankton chlorophyll : carbon ratio – the conversion factor between productivity and growth rate, Limnol. Oceanogr., 40, 1313–1321, https://doi.org/10.2307/2838689, 1995.</mixed-citation>
</ref>
<ref id="ref9">
<label>9</label><mixed-citation publication-type="other" xlink:type="simple">Collins, M., An, S.-I., Cai, W., Ganachaud, A., Guilyardi, E., Jin, F.-F., Jochum, M., Lengaigne, M., Power, S., Timmermann, A., Vecchi, G., and Wittenberg, A.: The Impact of Global Warming on the Tropical Pacific and El Niño, Nat. Geosci., 3, 391–397, https://doi.org/10.1038/ngeo868, 2010.</mixed-citation>
</ref>
<ref id="ref10">
<label>10</label><mixed-citation publication-type="other" xlink:type="simple">Edwards, A.&amp;nbsp;M.: A Rational Dynamical-Systems Approach to Plankton Population Modelling, Ph.D.&amp;nbsp;thesis, University of Leeds, 1997.</mixed-citation>
</ref>
<ref id="ref11">
<label>11</label><mixed-citation publication-type="other" xlink:type="simple">Edwards, A.&amp;nbsp;M. and Brindley, J.: Oscillatory behaviour in a three-component plankton population model, Dynam. Stabil. Syst., 11, 347–370, https://doi.org/10.1080/02681119608806231, 1996.</mixed-citation>
</ref>
<ref id="ref12">
<label>12</label><mixed-citation publication-type="other" xlink:type="simple">Edwards, A.&amp;nbsp;M., Wright, D.&amp;nbsp;G., and Platt, T.: Biological heating effect of a band of phytoplankton, J. Marine Syst., 49, 89–103, https://doi.org/10.1016/j.jmarsys.2003.05.011, 2003.</mixed-citation>
</ref>
<ref id="ref13">
<label>13</label><mixed-citation publication-type="other" xlink:type="simple">Gildor, H., Sobel, A.&amp;nbsp;H., Cane, M.&amp;nbsp;A., and Sambrutto, R.&amp;nbsp;N.: A role of ocean biota in tropical intraseasonal atmospheric variability, Geophys. Res. Lett., 30(9), 1460, https://doi.org/10.1029/2002GL016759, 2003.</mixed-citation>
</ref>
<ref id="ref14">
<label>14</label><mixed-citation publication-type="other" xlink:type="simple">Gill, A.&amp;nbsp;E.: Atmosphere-Ocean Dynamics, Academic Press Inc., London, UK, 1982.</mixed-citation>
</ref>
<ref id="ref15">
<label>15</label><mixed-citation publication-type="other" xlink:type="simple">Gnanadesikan, A. and Anderson, W.: Ocean water clarity and the ocean general circulation in a coupled climate model, J. Phys. Oceanogr., 39, 314–332, https://doi.org/10.1175/2008JPO3935.1, 2009.</mixed-citation>
</ref>
<ref id="ref16">
<label>16</label><mixed-citation publication-type="other" xlink:type="simple">Guilyardi, E., Wittenberg, A., Fedorov, A., Collins, M., Wang, C., Capotondi, A., van Oldenborgh, G., and Stockdale, T.: Understanding El Niño in Ocean-Atmosphere General Circulation Models : progress and challenges, B. Am. Meteorol. Soc., 90, 325–340, https://doi.org/10.1175/2008BAMS2387.1, 2009.</mixed-citation>
</ref>
<ref id="ref17">
<label>17</label><mixed-citation publication-type="other" xlink:type="simple">Jin, F.&amp;nbsp;F.: Tropical ocean-atmosphere interaction, the Pacific cold tongue and the El Niño-Southern Oscillation, Science, 274, 76–78, https://doi.org/10.1126/science.274.5284.76, 1996.</mixed-citation>
</ref>
<ref id="ref18">
<label>18</label><mixed-citation publication-type="other" xlink:type="simple">Jin, F.&amp;nbsp;F.: A Simple Model for the Pacific Cold Tongue and ENSO, J. Atmos. Sci., 2458–2469, 1998.</mixed-citation>
</ref>
<ref id="ref19">
<label>19</label><mixed-citation publication-type="other" xlink:type="simple">Jin, F.&amp;nbsp;F. and Neelin, J.&amp;nbsp;D.: Modes of interannual tropical ocean-atmosphere interaction – A unified view. Part&amp;nbsp;I: Numerical results, J. Atmos. Sci., 50, 3477–3502, 1993.</mixed-citation>
</ref>
<ref id="ref20">
<label>20</label><mixed-citation publication-type="other" xlink:type="simple">Kraus, E.&amp;nbsp;B. and Turner, J.&amp;nbsp;S.: A one-dimensional model of the seasonal thermocline. II: The general theory and its consequences, Tellus, 19, 98–109, https://doi.org/10.1111/j.2153-3490.1967.tb01462.x, 1967.</mixed-citation>
</ref>
<ref id="ref21">
<label>21</label><mixed-citation publication-type="other" xlink:type="simple">Kuznetsov, Y.&amp;nbsp;A. and Levitin, V.&amp;nbsp;V.: CONTENT: A multiplatform environment for continuation and bifurcation analysis of dynamical systems, Tech. rep., Centrum voor Wiskunde en Informatica, Amsterdam (Netherlands), 1997.</mixed-citation>
</ref>
<ref id="ref22">
<label>22</label><mixed-citation publication-type="other" xlink:type="simple">Latif, M., Anderson, D., Barnett, T., Cane, M., Kleeman, R., Leetmaa, A., O&apos;Brien, J., Rosati, A., and Schneider, E.: A review of predictability and prediction of ENSO, J. Geophys. Res., 103, 14375–14393, 1998.</mixed-citation>
</ref>
<ref id="ref23">
<label>23</label><mixed-citation publication-type="other" xlink:type="simple">Lengaigne, M., Menkes, C., Aumont, O., Gorgues, T., Bopp, L., André, J.-M., and Madec, G.: Influence of the oceanic biology on the tropical Pacific climate in a Coupled General Circulation Model, Clim. Dynam., 28, 503–516, https://doi.org/10.1007/s00382-006-0200-2, 2007.</mixed-citation>
</ref>
<ref id="ref24">
<label>24</label><mixed-citation publication-type="other" xlink:type="simple">Löptien, U., Eden, C., Timmermann, A., and Dietze, H.: Effects of biologically induced differential heating in an eddy-permitting coupled ocean-ecosystem model, J. Geophys. Res., 114, C06011, https://doi.org/10.1029/2008JC004936, 2009.</mixed-citation>
</ref>
<ref id="ref25">
<label>25</label><mixed-citation publication-type="other" xlink:type="simple">Manizza, M., Quéré, C.&amp;nbsp;L., Watson, A., and Buitenhuis, E.: Bio-optical feedbacks among phytoplankton upper ocean physics and sea-ice in a global model, Geophys. Res. Lett., 32, L05603, https://doi.org/10.1029/2004GL020 778, 2005.</mixed-citation>
</ref>
<ref id="ref26">
<label>26</label><mixed-citation publication-type="other" xlink:type="simple">Marzeion, B., Timmermann, A., Murtugudde, R., and Jin, F.-F.: Biophysical Feedbacks in the Tropical Pacific, J. Climate, 18, 58–70, 2005.</mixed-citation>
</ref>
<ref id="ref27">
<label>27</label><mixed-citation publication-type="other" xlink:type="simple">Murtugudde, R., Beauchamp, J., and Busalacchi, A.: Effects of Penetrative Radiation on the Upper Tropical Ocean Circulation, J. Climate, 15, 470–486, 2002.</mixed-citation>
</ref>
<ref id="ref28">
<label>28</label><mixed-citation publication-type="other" xlink:type="simple">Nakamoto, S., Kumar, S., Oberhuber, J., Ishizaka, J., Muneyama, K., and Frouin, R.: Response of the equatorial Pacific to chlorophyll pigment in a mixed layer isopycnal ocean general circulation model, Geophys. Res. Lett., 28(10), 2021–2024, https://doi.org/10.1029/2000GL012494, 2001.</mixed-citation>
</ref>
<ref id="ref29">
<label>29</label><mixed-citation publication-type="other" xlink:type="simple">Neelin, J.&amp;nbsp;D.: The slow sea surface temperature mode and the fast-wave limit: Analytic theory for tropical interannual oscillations and experiments in a hybrid coupled model, J. Atmos. Sci., 48, 584–606, 1991.</mixed-citation>
</ref>
<ref id="ref30">
<label>30</label><mixed-citation publication-type="other" xlink:type="simple">Philip, S. and van&amp;nbsp;Oldenborgh, G.: Shifts in ENSO coupling processes under global warming, Geophys. Res. Lett., 33, L11704, https://doi.org/10.1029/2006GL026196, 2006.</mixed-citation>
</ref>
<ref id="ref31">
<label>31</label><mixed-citation publication-type="other" xlink:type="simple">Platt, T., Sathyendranath, S., Edwards, A.&amp;nbsp;M., Broomhead, D.&amp;nbsp;S., and Ulloa, O.: Nitrate supply and demand in the mixed layer of the ocean, Mar. Ecol.-Prog. Ser., 254, 3–9, 2003.</mixed-citation>
</ref>
<ref id="ref32">
<label>32</label><mixed-citation publication-type="other" xlink:type="simple">Schopf, P.&amp;nbsp;S. and Cane, M.&amp;nbsp;A.: On equatorial dynamics, mixed layer physics and sea surface temperature, J. Phys. Oceanogr., 13, 917–935, 1983.</mixed-citation>
</ref>
<ref id="ref33">
<label>33</label><mixed-citation publication-type="other" xlink:type="simple">Seager, R., Zebiak, S.&amp;nbsp;E., and Cane, M.&amp;nbsp;A.: A Model for the Tropical Pacific Sea Surface Temperature Climatology, J. Geophys. Res., 93, 1265–1280, 1988.</mixed-citation>
</ref>
<ref id="ref34">
<label>34</label><mixed-citation publication-type="other" xlink:type="simple">Seo, H., Jochum, M., Murtugudde, R., and Miller, A.: Effect of ocean mesoscale variability on the mean state of tropical Atlantic climate, Geophys. Res. Lett., 33, L09606, https://doi.org/10.1029/2005GL025651, 2006.</mixed-citation>
</ref>
<ref id="ref35">
<label>35</label><mixed-citation publication-type="other" xlink:type="simple">Steele, J.&amp;nbsp;H. and Henderson, E.&amp;nbsp;W.: A simple plankton model, Am. Nat., 117, 676–691, 1981.</mixed-citation>
</ref>
<ref id="ref36">
<label>36</label><mixed-citation publication-type="other" xlink:type="simple">Strutton, P. and Chavez, F.: Biological heating in the equatorial Pacific: Observed variability and potential for real-time calculation, J. Climate, 17, 1097–1109, 2004.</mixed-citation>
</ref>
<ref id="ref37">
<label>37</label><mixed-citation publication-type="other" xlink:type="simple">Sweeney, C., Gnanadesikan, A., Griffies, S., Harrison, M., Rosati, A., and Samuel, B.: Impacts of shortwave penetration depth on the large-scale circulation and heat transport, J. Phys. Oceanogr., 35, 1103–1119, 2005.</mixed-citation>
</ref>
<ref id="ref38">
<label>38</label><mixed-citation publication-type="other" xlink:type="simple">Taylor, A.&amp;nbsp;H., Geider, R.&amp;nbsp;J., and Gilbert, F.&amp;nbsp;J.&amp;nbsp;H.: Seasonal and latitudinal dependencies of phytoplankton carbon-to-chlorophyll a ratios: results of a modelling study, Mar. Ecol.-Prog. Ser., 152, 51–66, https://doi.org/10.3354/meps152051, 1997.</mixed-citation>
</ref>
<ref id="ref39">
<label>39</label><mixed-citation publication-type="other" xlink:type="simple">Timmermann, A. and Jin, F.-F.: Phytoplankton influences on tropical climate, Geophys. Res. Lett., 29(23), 2104, https://doi.org/10.1029/2002GL015434, 2002.</mixed-citation>
</ref>
<ref id="ref40">
<label>40</label><mixed-citation publication-type="other" xlink:type="simple">Timmermann, A., Latif, M., Bacher, A., Oberhuber, J., and Roeckner, E.: Increased El Niño frequency in a climate model forced by future greenhouse warming, Nature, 398, 694–696, https://doi.org/10.1038/19505, 1999.</mixed-citation>
</ref>
<ref id="ref41">
<label>41</label><mixed-citation publication-type="other" xlink:type="simple">van Oldenborgh, G., Burgers, G., and Tank, A.&amp;nbsp;K.: On the El Niño Teleconnection to spring precipitation in Europe, Int. J. Climatol., 20, 565–574, https://doi.org/10.1002/(SICI)1097-0088(200004), 2000.</mixed-citation>
</ref>
<ref id="ref42">
<label>42</label><mixed-citation publication-type="other" xlink:type="simple">Wang, Y., Xie, S.-P., Xu, H., and Wang, B.: Regional model simulations of marine boundary layer clouds over the Southeast Pacific off South America. Part I: Control experiment, Mon. Weather Rev., 132, 274–296, 2004.</mixed-citation>
</ref>
<ref id="ref43">
<label>43</label><mixed-citation publication-type="other" xlink:type="simple">Wang, J., Cota, G., and Comiso, J.: Phytoplankton in the Beaufort and Chukchi Seas: Distribution, dynamics, and environmental forcing, Deep-Sea Research Pt.&amp;nbsp;II, 52, 3355–3368, https://doi.org/ 10.1016/j.dsr2.2005.10.014, 2005.</mixed-citation>
</ref>
<ref id="ref44">
<label>44</label><mixed-citation publication-type="other" xlink:type="simple">Wetzel, P., Maier-Reimer, E., Botzet, M., Jungclaus, J., Keenlyside, N., and Latif, M.: Effects of ocean biology on the penetrative radiation in a coupled climate model, J. Climate, 19, 3973–3987, 2006.</mixed-citation>
</ref>
<ref id="ref45">
<label>45</label><mixed-citation publication-type="other" xlink:type="simple">Xie, S.-P.: Effects of Seasonal Forcing on Latitudinal Asymmetry of the ITCZ, J. Climate, 9, 2945–2950, 1996.</mixed-citation>
</ref>
<ref id="ref46">
<label>46</label><mixed-citation publication-type="other" xlink:type="simple">Yu, J.-Y. and Mechoso, C.: Links between annual variations of Peruvian stratocumulus clouds and of SST in the eastern equatorial Pacific, J. Climate, 12, 3305–3318, 1999.</mixed-citation>
</ref>
<ref id="ref47">
<label>47</label><mixed-citation publication-type="other" xlink:type="simple">Zhang, R.-H., Busalacchi, A.&amp;nbsp;J., Wang, X., Ballabrera-Poy, J., Murtugudde, R.&amp;nbsp;G., Hackert, E.&amp;nbsp;C., and Chen, D.: Role of ocean biology-induced climate feedback in the modulation of El Niño-Southern Oscillation, Geophys. Res. Lett., 36, 1–6, https://doi.org/10.1029/2008GL036568, 2009.</mixed-citation>
</ref>
</ref-list>
</back>
</article>