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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Mar. Sci.</journal-id>
<journal-title>Frontiers in Marine Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Mar. Sci.</abbrev-journal-title>
<issn pub-type="epub">2296-7745</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmars.2024.1473208</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Marine Science</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Representing ocean biology-induced heating effects in ROMS-based simulations for the Indo-Pacific Ocean</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Zhang</surname>
<given-names>Wenzhe</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2804610"/>
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</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Gao</surname>
<given-names>Chuan</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Tian</surname>
<given-names>Feng</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
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<contrib contrib-type="author">
<name>
<surname>Yu</surname>
<given-names>Yang</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2026748"/>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Wang</surname>
<given-names>Hongna</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
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</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Zhang</surname>
<given-names>Rong-Hua</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
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<aff id="aff1">
<sup>1</sup>
<institution>Key Laboratory of Ocean Observation and Forecasting and Key Laboratory of Ocean Circulation and Waves, Institute of Oceanology, Chinese Academy of Sciences</institution>, <addr-line>Qingdao</addr-line>, <country>China</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>University of Chinese Academy of Sciences</institution>, <addr-line>Beijing</addr-line>, <country>China</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Laoshan Laboratory</institution>, <addr-line>Qingdao</addr-line>, <country>China</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>School of Marine Sciences, Nanjing University of Information Science and Technology</institution>, <addr-line>Nanjing</addr-line>, <country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Kyung-Ae Park, Seoul National University, Republic of Korea</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Hailun He, Ministry of Natural Resources, China</p>
<p>Tien Anh Tran, Seoul National University, Republic of Korea</p>
<p>Xianqing Lv, Ocean University of China, China</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Chuan Gao, <email xlink:href="mailto:gaochuan@qdio.ac.cn">gaochuan@qdio.ac.cn</email>; Rong-Hua Zhang, <email xlink:href="mailto:rzhang@nuist.edu.cn">rzhang@nuist.edu.cn</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>07</day>
<month>11</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>11</volume>
<elocation-id>1473208</elocation-id>
<history>
<date date-type="received">
<day>30</day>
<month>07</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>14</day>
<month>10</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2024 Zhang, Gao, Tian, Yu, Wang and Zhang</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Zhang, Gao, Tian, Yu, Wang and Zhang</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>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) and the copyright owner(s) 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.</p>
</license>
</permissions>
<abstract>
<p>Incident shortwave radiation can penetrate and heat the upper ocean water column, acting to modulate the stratification, vertical mixing and sea surface temperature. As a light-absorbing constituent, ocean chlorophyll (CHL) plays an important role in regulating these processes; however, its heating effect on the ocean state remains controversial and exhibits strong model dependence on ways the solar radiation transmission and the related CHL-induced heating are represented. In this study, we implement a chlorophyll-based two-way coupling between physical and ecological processes within the Regional Ocean Modeling System (ROMS). The bio-physics coupled model performs well in simulating the structure and variability of oceanic physical and ecological fields in the tropical Indo-Pacific region. Three CHL-related heating terms are analyzed based on the model output to diagnose the ocean biology-induced heating effects, namely the shortwave radiation part penetrating out of the base of the mixed layer (ML; <italic>Q<sub>pen</sub>
</italic>), the portion absorbed within the ML (<italic>Q<sub>abs</sub>
</italic>), and the rate of temperature change of the ML resulting from the <italic>Q<sub>abs</sub>
</italic> effects (<italic>R<sub>sr</sub>
</italic>). Results show that the spatio-temporal distributions of the three heating terms are mainly determined by the ML depth (MLD). However, <italic>Q<sub>pen</sub>
</italic> can also be regulated by the euphotic depth (ED), especially in the western-central equatorial Pacific. This moderating effect is particularly evident during El Ni&#xf1;o when the ED tends to be greater than the MLD; positive ED anomalies act to enhance the positive <italic>Q<sub>pen</sub>
</italic> anomalies caused by negative MLD anomalies. For the first time, the bio-heating effects are quantified within the ROMS-based two-way coupling context between the physical submodel and ecological submodel over the tropical Indo-Pacific Ocean, providing a basis for further understanding of the bio-effects and mechanisms. It is expected that the methodology and understanding developed in this study can help explore the chlorophyll-related processes in the ocean and the interactions with the atmosphere.</p>
</abstract>
<kwd-group>
<kwd>ocean physical-ecological model</kwd>
<kwd>shortwave radiation penetration</kwd>
<kwd>ocean biology-induced heating</kwd>
<kwd>two-way coupling</kwd>
<kwd>Regional Ocean Modeling System (ROMS)</kwd>
</kwd-group>    <contract-num rid="cn001">NSFC; Grant Nos. 42176032, 42030410&#x56fd;&#x5bb6;&#x81ea;&#x7136;&#x79d1;&#x5b66;&#x57fa;&#x91d1;;&#x7b2c;42176032&#x53f7;&#x6388;&#x6743;&#x4e66;&#xff0c;42030410</contract-num>    <contract-sponsor id="cn001">National Natural Science Foundation of China<named-content content-type="fundref-id">10.13039/501100001809</named-content>
</contract-sponsor>
<counts>
<fig-count count="10"/>
<table-count count="2"/>
<equation-count count="7"/>
<ref-count count="78"/>
<page-count count="17"/>
<word-count count="7791"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Physical Oceanography</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Solar radiation is the primary energy source for physical, biological and chemical processes in the Earth&#x2019;s surface system (<xref ref-type="bibr" rid="B1">Allen, 1997</xref>). Solar radiation can penetrate through the sea surface and heat a few meters of the upper ocean water column, which thus affects the stratification and mixing in the upper ocean. In addition, solar radiation provides energy for photosynthesis and helps fix carbon dioxide in the presence of chlorophyll in phytoplankton. The ecological processes in the ocean are more complex than those on land, because they are affected by the geofluid dynamic processes in the marine physical environment. For example, the ocean stratification causes the retention of phytoplankton in the upper layer, wherein light is more readily available with inorganic nutrients being limited (<xref ref-type="bibr" rid="B37">Mann and Lazier, 2006</xref>). In addition, chlorophyll-a is the main light-absorbing constituent in the ocean (<xref ref-type="bibr" rid="B41">Mobley, 2001</xref>), acting to trap more solar radiation in the surface layer, which in turn alters the vertical distribution of light and heat in the ocean (<xref ref-type="bibr" rid="B37">Mann and Lazier, 2006</xref>).</p>
<p>In the tropical upper ocean, phytoplankton is usually not light-limited (<xref ref-type="bibr" rid="B54">Shell et&#xa0;al., 2003</xref>; <xref ref-type="bibr" rid="B37">Mann and Lazier, 2006</xref>; <xref ref-type="bibr" rid="B23">Jochum et&#xa0;al., 2010</xref>). In particular, in the eastern equatorial Pacific, as the strong equatorial upwelling brings abundant nutrients, there is a triangular region with higher phytoplankton biomass expressed as the chlorophyll concentration (<xref ref-type="bibr" rid="B29">Lewis et&#xa0;al., 1990</xref>; <xref ref-type="bibr" rid="B32">Lin et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B25">Khanna et&#xa0;al., 2009</xref>). In the equatorial Pacific, the impact of chlorophyll on SST can be further amplified through the ocean-atmosphere interaction and then can influence SST interannual variations in association with the El Ni&#xf1;o-Southern Oscillation (ENSO) events (<xref ref-type="bibr" rid="B48">Philander, 1983</xref>; <xref ref-type="bibr" rid="B72">Zhang and Gao, 2016</xref>; <xref ref-type="bibr" rid="B14">Gao and Zhang, 2023</xref>; <xref ref-type="bibr" rid="B13">Gao et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B78">Zhang et&#xa0;al., 2020</xref>, <xref ref-type="bibr" rid="B73">2022</xref>, <xref ref-type="bibr" rid="B75">2024</xref>). In addition to interactions among the hydrosphere, biosphere and atmosphere, there are also inter-basin linkages between the tropical Pacific and Indian Ocean, which may cause inter-basin effects through the Walker circulation in the atmosphere (<xref ref-type="bibr" rid="B27">Latif and Barnett, 1995</xref>) and the Indonesian Throughflow in the ocean (<xref ref-type="bibr" rid="B58">Sprintall et&#xa0;al., 2014</xref>). Thus, it is necessary to take into account chlorophyll variability and its effect in the Indo-Pacific region (<xref ref-type="bibr" rid="B42">Murtugudde et&#xa0;al., 2002</xref>; <xref ref-type="bibr" rid="B39">Marzeion et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B65">Wetzel et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B4">Ballabrera-Poy et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B28">Lengaigne et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B44">Park et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B24">Kang et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B77">Zhang et&#xa0;al., 2019b</xref>; <xref ref-type="bibr" rid="B61">Tian et&#xa0;al., 2021</xref>). At present, it is difficult to quantify the effects of ocean biology-induced heating (OBH) on the climate through <italic>in situ</italic> observations. Fortunately, since the launch of the Coastal Zone Color Scanner in 1979, the Sea-viewing Wide Field-of-view Sensor (SeaWiFS), the Moderate-resolution Imaging Spectroradiometer and the Medium Resolution Imaging Spectrometer have been launched successively (<xref ref-type="bibr" rid="B18">Gordon and Morel, 1983</xref>; <xref ref-type="bibr" rid="B37">Mann and Lazier, 2006</xref>; <xref ref-type="bibr" rid="B63">Valiela, 2015</xref>). The development of satellite remote sensing makes it possible to obtain satellite color images of the ocean surface and to retrieve chlorophyll by inverse modeling techniques (<xref ref-type="bibr" rid="B50">Rast et&#xa0;al., 1999</xref>). With the mapping of chlorophyll concentrations over large areas, the influence of chlorophyll on shortwave radiation penetration in the vertical direction can be considered in large-scale ocean models.</p>
<p>Numerical models have been developed to study the bio-physical feedback induced by chlorophyll (CHL) in the tropical Pacific. Several types of ocean general circulation models (OGCMs) based on different vertical coordinate systems have been employed to study the ocean biology-induced heating effects, including the most popularly used level OGCMs developed by GFDL/NOAA (<xref ref-type="bibr" rid="B52">Sathyendranath et&#xa0;al., 1991</xref>; <xref ref-type="bibr" rid="B60">Sweeney et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B44">Park et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B24">Kang et&#xa0;al., 2017</xref>), and layer ocean model (e.g., <xref ref-type="bibr" rid="B67">Zhang, 2015a</xref>; <xref ref-type="bibr" rid="B15">Gao et&#xa0;al., 2020</xref>). Historically, CHL-induced heating effects in oceanic models have been differently represented, which evolves with different complexities, including horizontally uniform penetration depth of solar radiation (<xref ref-type="bibr" rid="B65">Wetzel et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B4">Ballabrera-Poy et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B17">Gnanadesikan and Anderson, 2009</xref>), the spatially-varying sea surface CHL from satellite observations (<xref ref-type="bibr" rid="B2">Anderson et&#xa0;al., 2007</xref>, <xref ref-type="bibr" rid="B3">2009</xref>; <xref ref-type="bibr" rid="B44">Park et&#xa0;al., 2014</xref>), the empirical statistical model for CHL interannual anomalies (<xref ref-type="bibr" rid="B69">Zhang et&#xa0;al., 2009</xref>, <xref ref-type="bibr" rid="B70">2011</xref>; <xref ref-type="bibr" rid="B68">Zhang, 2015b</xref>), and the interactive oceanic ecological models for horizontally and vertically varying chlorophyll depictions (<xref ref-type="bibr" rid="B65">Wetzel et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B28">Lengaigne et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B45">Patara et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B44">Park et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B30">Lim et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B76">Zhang et&#xa0;al., 2018</xref>, <xref ref-type="bibr" rid="B74">2019a</xref>; <xref ref-type="bibr" rid="B61">Tian et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B55">Shi et&#xa0;al., 2023</xref>). For example, <xref ref-type="bibr" rid="B69">Zhang et&#xa0;al. (2009)</xref> coupled the Gent-Cane ocean general circulation model (OGCM) with an empirical statistical model for the penetration depth (<inline-formula>
<mml:math display="inline" id="im1">
<mml:mrow>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mi>p</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>, defined as the inverse of the attenuation coefficient for the penetrative solar radiation), and diagnosed the impact of interannual <inline-formula>
<mml:math display="inline" id="im2">
<mml:mrow>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mi>p</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> variability on bio-heating. Previous studies indicated that chlorophyll can cause contrasting effects on sea surface temperature (SST) and its interannual variability through direct thermodynamic heating (<xref ref-type="bibr" rid="B60">Sweeney et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B35">L&#xf6;ptien et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B44">Park et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B30">Lim et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B16">Gera et&#xa0;al., 2020</xref>) or indirect dynamic cooling (<xref ref-type="bibr" rid="B42">Murtugudde et&#xa0;al., 2002</xref>; <xref ref-type="bibr" rid="B39">Marzeion et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B69">Zhang et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B16">Gera et&#xa0;al., 2020</xref>), respectively. Currently, the nature of the bio-effects in oceanic and coupled ocean-atmosphere models remains elusive and the underlying processes are controversial. Evidently, the results from previous modeling studies are quite inconsistent and strongly model-dependent in terms of the ocean biology-induced heating effects on mean SST and its variability in the eastern equatorial Pacific (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>), underscoring the need for alternative modeling approaches.</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Representative studies investigating the influence of chlorophyll on the eastern equatorial Pacific mean sea surface temperature (SST) using models with different vertical coordinate systems.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">vertical coordinate</th>
<th valign="middle" align="left">Warm</th>
<th valign="middle" align="left">Cool</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">z-coordinate</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B65">Wetzel et&#xa0;al. (2006)</xref> <break/>
<xref ref-type="bibr" rid="B45">Patara et&#xa0;al. (2012)</xref>
</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B36">Manizza (2005)</xref> <break/>
<xref ref-type="bibr" rid="B35">L&#xf6;ptien et&#xa0;al. (2009)</xref> <break/>
<xref ref-type="bibr" rid="B23">Jochum et&#xa0;al. (2010)</xref> <break/>
<xref ref-type="bibr" rid="B44">Park et&#xa0;al.  (2014)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">&#x3c1;-coordinate</td>
<td valign="top" align="left"/>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B43">Nakamoto et&#xa0;al. (2001)</xref> <break/>
<xref ref-type="bibr" rid="B2">Anderson et&#xa0;al. (2007</xref>, <xref ref-type="bibr" rid="B3">2009</xref>) <break/>
<xref ref-type="bibr" rid="B17">Gnanadesikan and Anderson (2009)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">&#x3b7;-coordinate</td>
<td valign="top" align="left"/>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B32">Lin et&#xa0;al. (2007)</xref>, <xref ref-type="bibr" rid="B33">Lin et&#xa0;al. (2008)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">&#x3c3;-coordinate</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B42">Murtugudde et&#xa0;al. (2002)</xref> <break/>
<xref ref-type="bibr" rid="B39">Marzeion et&#xa0;al. (2005)</xref> <break/>
<xref ref-type="bibr" rid="B61">Tian et&#xa0;al. (2021)</xref>
</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B61">Tian et&#xa0;al. (2021)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>The Regional Ocean Modeling System (ROMS) is an ocean model which is based on free-surface, terrain-following and primitive-equations, which is widely utilized among oceanographers (<xref ref-type="bibr" rid="B53">Shchepetkin and McWilliams, 2005</xref>; <xref ref-type="bibr" rid="B20">Hedstr&#xf6;m, 2018</xref>). The ROMS offers flexible vertical coordinate systems that incorporate two kinds of vertical transformation equations and a variety of vertical stretching functions. The stretched coordinates allow higher resolution in the areas of interest, such as the upper mixed layer (<xref ref-type="bibr" rid="B56">Song and Haidvogel, 1994</xref>; <xref ref-type="bibr" rid="B53">Shchepetkin and McWilliams, 2005</xref>). The ROMS includes several vertical mixing parameterization schemes for turbulence closure and modules for ecological, sediment and sea ice applications. Although the ROMS offers various ecosystem sub-models with varying levels of complexity (<xref ref-type="bibr" rid="B19">Haidvogel et&#xa0;al., 2008</xref>), the coupling between its physical and ecological sub-models is in one way from the physics to ecological state, in which the scheme of <xref ref-type="bibr" rid="B46">Paulson and Simpson (1977)</xref> is adopted for solar radiation penetration parameterization. For example, in current ROMS setting, a simplified empirical formula is adopted to represent this intricate radiation transfer process. The bimodal-exponential function of <xref ref-type="bibr" rid="B46">Paulson and Simpson (1977)</xref> is a traditional parameterization scheme to represent the attenuation of shortwave radiation with depth (z, which is defined to be negative below the ocean surface). The formula is expressed as follows:</p>
<disp-formula id="eq1">
<label>(1)</label>
<mml:math display="block" id="M1">
<mml:mrow>
<mml:mi>I</mml:mi>
<mml:mo stretchy="false">/</mml:mo>
<mml:msub>
<mml:mi>I</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:msub>
<mml:mi>A</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
<mml:mtext>exp</mml:mtext>
<mml:mrow>
<mml:mo>(</mml:mo>
<mml:mrow>
<mml:mfrac>
<mml:mi>z</mml:mi>
<mml:mrow>
<mml:msub>
<mml:mi>B</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
<mml:mo>)</mml:mo>
</mml:mrow>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>A</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:mtext>exp</mml:mtext>
<mml:mrow>
<mml:mo>(</mml:mo>
<mml:mrow>
<mml:mfrac>
<mml:mi>z</mml:mi>
<mml:mrow>
<mml:msub>
<mml:mi>B</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
<mml:mo>)</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <italic>I</italic> and <inline-formula>
<mml:math display="inline" id="im3">
<mml:mrow>
<mml:msub>
<mml:mi>I</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> denote the shortwave radiation at depth <italic>z</italic> and the sea surface, respectively. The proportion of <italic>I</italic> to <inline-formula>
<mml:math display="inline" id="im4">
<mml:mrow>
<mml:msub>
<mml:mi>I</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> represents the decay rate of shortwave radiation. <inline-formula>
<mml:math display="inline" id="im5">
<mml:mrow>
<mml:msub>
<mml:mi>A</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>, <inline-formula>
<mml:math display="inline" id="im6">
<mml:mrow>
<mml:msub>
<mml:mi>A</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>, <inline-formula>
<mml:math display="inline" id="im7">
<mml:mrow>
<mml:msub>
<mml:mi>B</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> and <inline-formula>
<mml:math display="inline" id="im8">
<mml:mrow>
<mml:mo>&#xa0;</mml:mo>
<mml:msub>
<mml:mi>B</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> are tunable parameters determined empirically by fitting the function to downward irradiance observations according to the water types of <xref ref-type="bibr" rid="B22">Jerlov (1968)</xref>. The descriptive classification scheme defines five types of water based on the ocean transparency that can be considered as a proxy for chlorophyll concentration. For example, a clear open ocean is classified as Type I (<xref ref-type="bibr" rid="B22">Jerlov, 1968</xref>). In such a way, when the water type is given, the decay rate at a particular depth is identical everywhere regardless of the distribution of chlorophyll. Clearly, this parameterization scheme for the penetration of solar radiation in the upper ocean cannot represent the effects of temporal and spatial variations in chlorophyll concentrations; only the impacts of ocean dynamics on ecological processes are taken into account, while the impacts of chlorophyll on the physical fields are not considered. So, there is no feedback from ocean ecological processes to physical processes. Note that <xref ref-type="bibr" rid="B34">Liu et&#xa0;al. (2020)</xref> previously tested four shortwave radiation transmission schemes using an idealized one-dimensional ROMS model, and they found that the simulation results with chlorophyll-based schemes exhibit obvious differences in the vertical temperature and upper mixing when compared with the default water-type scheme, with SST differences reaching 1.5&#x2013;2.0&#xb0;C. These results clearly indicate the importance of incorporating chlorophyll-based shortwave radiation parameterization schemes into the ROMS. Despite its capabilities and wide applications to regional oceanic problems, the ROMS has not yet been extensively utilized in simulating and understanding large-scale ocean circulations. Previous studies using ROMS-based ecological models have been limited by their shortwave radiation scheme. The ocean biology-induced heating effects have not been adequately explored.</p>
<p>This study addresses this gap by implementing a two-way coupling between the physical and ecological processes within the ROMS, in which the default shortwave penetration parameterization scheme within the ROMS is substituted by a chlorophyll-based scheme. This modeling application to a basinwide Indo-Pacific region allows to represent the CHL-induced heating feedback onto physics, including SST, a field that is important to the climate system. Furthermore, processes underlying the bio-effects are analyzed based on the ROMS simulations for the Indo-Pacific region; the corresponding bio-heating effects are quantified by adopting a diagnostic scheme and separating the shortwave radiation part penetrating out of the base of the mixed layer (ML; <italic>Q<sub>pen</sub>
</italic>), the portion absorbed within the ML (<italic>Q<sub>abs</sub>
</italic>), and the rate of temperature change of the ML (<italic>R<sub>sr</sub>
</italic>), respectively. These dedicated efforts provide a basis for further understanding of the thermodynamic and dynamic effects associated with ocean biology-induced heating within the ROMS. It is expected that the methodology and understanding developed in this study can help explore the CHL-related processes in the ocean and further the interactions with the atmosphere.</p>
<p>A detailed description of the model, experiments and data sources is provided in Section 2. The model results are described in Section 3, in which following the method of <xref ref-type="bibr" rid="B70">Zhang et&#xa0;al. (2011</xref>; <xref ref-type="bibr" rid="B67">Zhang, 2015a</xref>), three bio-heating terms are analyzed to isolate and quantify the direct OBH effects in association with the mixed layer depth (MLD) and chlorophyll. A brief discussion and conclusion are given in Sections 4 and 5. <xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref> presents the nomenclature and abbreviation adopted in this paper.</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>The nomenclature and abbreviation adopted in this paper.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">Symbol</th>
<th valign="top" align="left">Meaning</th>
<th valign="top" align="left">Value and/or unit</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">A<sub>1</sub>, A<sub>2</sub>, B<sub>1</sub>, B<sub>2</sub>
</td>
<td valign="top" align="left">parameters determined empirically by fitting the function to observed downward irradiance according to the water types</td>
<td valign="top" align="left">for Type I water:<break/>A<sub>1</sub> = 0.58, B<sub>1</sub> = 0.35 m<break/>A<sub>2</sub> = 0.42, B<sub>2</sub> = 23.0 m</td>
</tr>
<tr>
<td valign="top" align="left">CHL</td>
<td valign="top" align="left">chlorophyll</td>
<td valign="top" align="left">mg Chl m<sup>-3</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">Chl(&#x3b6;)</td>
<td valign="top" align="left">the chlorophyll concentration at a certain point &#x3b6;&#x2208;[0,<italic>z</italic>]</td>
<td valign="top" align="left">mg Chl m<sup>-3</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">Chl: N</td>
<td valign="top" align="left">the ratio of chlorophyll to nitrogen</td>
<td valign="top" align="left">0.53 gChl molN<sup>&#x2212;1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">C: N</td>
<td valign="top" align="left">the Redfield ratio of carbon to nitrogen</td>
<td valign="top" align="left">106:16 molC molN<sup>&#x2212;1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">C: Chl</td>
<td valign="top" align="left">the ratio of carbon to chlorophyll</td>
<td valign="top" align="left">150:1 gC gChl<sup>&#x2212;1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">C<sub>p</sub>
</td>
<td valign="top" align="left">specific heat capacity of seawater</td>
<td valign="top" align="left">4000 J K<sup>-1</sup> kg<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">&#x3b3;</td>
<td valign="top" align="left">photosynthetic available radiation</td>
<td valign="top" align="left">0.43</td>
</tr>
<tr>
<td valign="top" align="left">&#x3c1;<sub>0</sub>
</td>
<td valign="top" align="left">the mean density of seawater</td>
<td valign="top" align="left">1025 kg m<sup>-3</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">&#x3c1;</td>
<td valign="top" align="left">the density of seawater</td>
<td valign="top" align="left">kg m<sup>-3</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">&#x3c1;<sub>MLD&#x2003;</sub>
</td>
<td valign="top" align="left">the density of seawater at the MLD</td>
<td valign="top" align="left">kgm<sup>-3</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">ECMWF</td>
<td valign="top" align="left">the European Centre for Medium-range Weather Forecasts</td>
<td valign="top" align="left">
<bold>/</bold>
</td>
</tr>
<tr>
<td valign="top" align="left">ED</td>
<td valign="top" align="left">the euphotic depth</td>
<td valign="top" align="left">m</td>
</tr>
<tr>
<td valign="top" align="left">ENSO</td>
<td valign="top" align="left">the El Ni&#xf1;o/Southern Oscillation</td>
<td valign="top" align="left">
<bold>/</bold>
</td>
</tr>
<tr>
<td valign="top" align="left">EP El Ni&#xf1;o</td>
<td valign="top" align="left">the eastern Pacific El Ni&#xf1;o</td>
<td valign="top" align="left">
<bold>/</bold>
</td>
</tr>
<tr>
<td valign="top" align="left">CP El Ni&#xf1;o</td>
<td valign="top" align="left">the central Pacific El Ni&#xf1;o</td>
<td valign="top" align="left">
<bold>/</bold>
</td>
</tr>
<tr>
<td valign="top" align="left">F<sub>p</sub>
</td>
<td valign="top" align="left">bioavailable iron incorporated within phytoplankton cells</td>
<td valign="top" align="left">mmolFe m<sup>-3</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">F<sub>d</sub>
</td>
<td valign="top" align="left">dissolved iron concentration</td>
<td valign="top" align="left">mmolFe m<sup>-3</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">H<sub>p</sub>
</td>
<td valign="top" align="left">the penetration depth</td>
<td valign="top" align="left">m</td>
</tr>
<tr>
<td valign="top" align="left">H<sub>m</sub>
</td>
<td valign="top" align="left">the mixed layer depth&#x2003;</td>
<td valign="top" align="left">m</td>
</tr>
<tr>
<td valign="top" align="left">I, I<sub>0</sub>
</td>
<td valign="top" align="left">the shortwave radiation at depth <italic>z</italic> and the sea surface</td>
<td valign="top" align="left">W m<sup>-2</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">I<sub>ED</sub>
</td>
<td valign="top" align="left">the shortwave radiation at the euphotic depth</td>
<td valign="top" align="left">
<bold>/</bold>
</td>
</tr>
<tr>
<td valign="top" align="left">K<sub>w</sub>
</td>
<td valign="top" align="left">the vertical attenuation coefficients of light for water</td>
<td valign="top" align="left">0.028 m<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">K<sub>chl</sub>
</td>
<td valign="top" align="left">the vertical attenuation coefficients of light for chlorophyll</td>
<td valign="top" align="left">0.058 m mg Chl<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">ML</td>
<td valign="top" align="left">the mixed layer</td>
<td valign="top" align="left">/</td>
</tr>
<tr>
<td valign="top" align="left">MLD</td>
<td valign="top" align="left">the mixed layer depth</td>
<td valign="top" align="left">m</td>
</tr>
<tr>
<td valign="top" align="left">NPZD</td>
<td valign="top" align="left">nutrient-phytoplankton-zooplankton-detritus</td>
<td valign="top" align="left">
<bold>/</bold>
</td>
</tr>
<tr>
<td valign="top" align="left">N</td>
<td valign="top" align="left">nitrate concentration</td>
<td valign="top" align="left">mmolN m<sup>-3</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">P</td>
<td valign="top" align="left">phytoplankton biomass</td>
<td valign="top" align="left">mmolN m<sup>-3</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">Z</td>
<td valign="top" align="left">zooplankton biomass</td>
<td valign="top" align="left">mmolN m<sup>-3</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">D</td>
<td valign="top" align="left">detritus concentration</td>
<td valign="top" align="left">mmolN m<sup>-3</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">OBH</td>
<td valign="top" align="left">ocean biology-induced heating</td>
<td valign="top" align="left">
<bold>/</bold>
</td>
</tr>
<tr>
<td valign="top" align="left">OGCM</td>
<td valign="top" align="left">ocean general circulation model</td>
<td valign="top" align="left">
<bold>/</bold>
</td>
</tr>
<tr>
<td valign="top" align="left">OISST</td>
<td valign="top" align="left">the Optimum Interpolation Sea Surface Temperature</td>
<td valign="top" align="left">
<bold>/</bold>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Q<sub>abs</sub>
</italic>
</td>
<td valign="top" align="left">absorbed shortwave radiation within the ML</td>
<td valign="top" align="left">W m<sup>-2</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Q<sub>pen</sub>
</italic>
</td>
<td valign="top" align="left">the penetrative shortwave radiation out of the base of ML</td>
<td valign="top" align="left">W m<sup>-2</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">Q<sub>sr</sub>
</td>
<td valign="top" align="left">the incident solar radiation flux at the sea surface</td>
<td valign="top" align="left">W m<sup>-2</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>R<sub>sr</sub>
</italic>
</td>
<td valign="top" align="left">the rate of temperature change resulting from the <italic>Q<sub>abs</sub>
</italic> effect in the ML</td>
<td valign="top" align="left">&#xb0;C month<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">ROMS</td>
<td valign="top" align="left">the Regional Ocean Modeling System</td>
<td valign="top" align="left">
<bold>/</bold>
</td>
</tr>
<tr>
<td valign="top" align="left">SeaWiFS</td>
<td valign="top" align="left">the Sea-viewing Wide Field-of-view Sensor</td>
<td valign="top" align="left">/</td>
</tr>
<tr>
<td valign="top" align="left">SST</td>
<td valign="top" align="left">sea surface temperature</td>
<td valign="top" align="left">&#xb0;C</td>
</tr>
<tr>
<td valign="top" align="left">WOA</td>
<td valign="top" align="left">the World Ocean Atlas</td>
<td valign="top" align="left">
<bold>/</bold>
</td>
</tr>
<tr>
<td valign="top" align="left">T<sub>0</sub>, S<sub>0</sub>, p<sub>0</sub>
</td>
<td valign="top" align="left">the temperature, salinity and pressure values at the sea surface in determining the MLD</td>
<td valign="top" align="left"/>
</tr>
<tr>
<td valign="top" align="left">&#x394;T</td>
<td valign="top" align="left">the temperature threshold</td>
<td valign="top" align="left">0.5 &#xb0;C</td>
</tr>
<tr>
<td valign="top" align="left">z</td>
<td valign="top" align="left">the geographic depth (negative below the ocean surface)</td>
<td valign="top" align="left">m</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s2">
<label>2</label>
<title>The model, experimental design and data</title>
<p>In this study, ocean-only experiments are performed using a basin-scale three-dimensional ocean circulation model and a simple ecological model. The former is the ROMS provided by the ROMS/Terrain-following Ocean Modeling System Group (<ext-link ext-link-type="uri" xlink:href="https://www.myroms.org/">https://www.myroms.org/</ext-link>). The latter is the Nutrient-Phytoplankton-Zooplankton-Detritus Model with Iron Limitation on Phytoplankton Growth (hereinafter referred to as the NPZD-IRON) developed by <xref ref-type="bibr" rid="B11">Fiechter et&#xa0;al. (2009)</xref>. Some details are presented in this section.</p>
<sec id="s2_1">
<label>2.1</label>
<title>Model configuration</title>
<sec id="s2_1_1">
<label>2.1.1</label>
<title>Physical model</title>
<p>The ocean model used is the Regional Ocean Modeling System (ROMS), which is based on free-surface, terrain-following and primitive-equations, and it is widely utilized among oceanographers. The stretched coordinates allow higher resolution in the areas of interest, such as the upper mixed layer (<xref ref-type="bibr" rid="B56">Song and Haidvogel, 1994</xref>; <xref ref-type="bibr" rid="B53">Shchepetkin and McWilliams, 2005</xref>). The ROMS includes several vertical mixing parameterization schemes for turbulence closure and modules for ecological, sediment and sea ice applications. The Mellor-Yamada level-2.5 closure scheme (<xref ref-type="bibr" rid="B40">Mellor and Yamada, 1982</xref>) is chosen for the surface ocean mixing parameterization, and the bathymetric dataset Earth Topography 2 is used to represent the ocean bottom topography. The model domain covers the entire tropical Pacific and Indian Ocean (20&#xb0;E&#x2013;70&#xb0;W, 35&#xb0;S&#x2013;35&#xb0;N), with a zonal resolution of 0.5&#xb0; and meridionally varying resolution of 0.4&#xb0;-0.6&#xb0; and with 50 vertical &#x3c3;-levels. The monthly climatology of temperature and salinity from the World Ocean Atlas (WOA) 2009 is used to provide initial and lateral boundary conditions for the model (<xref ref-type="bibr" rid="B47">Penven et&#xa0;al., 2008</xref>). There are three open boundaries at the northern, southern and western edges. The free surface and barotropic horizontal velocities are respectively subject to the boundary conditions as designed in <xref ref-type="bibr" rid="B7">Chapman (1985)</xref> and <xref ref-type="bibr" rid="B12">Flather (1976)</xref>, while the baroclinic horizontal velocities and tracers use radiation boundary conditions with a nudging to observations (<xref ref-type="bibr" rid="B38">Marchesiello et&#xa0;al., 2001</xref>).</p>
</sec>
<sec id="s2_1_2">
<label>2.1.2</label>
<title>Ecological model</title>
<p>The NPZD-IRON includes four standard state variables (represented in terms of nitrogen concentration): nitrate concentration (N), phytoplankton biomass (P), zooplankton biomass (Z) and detritus concentration (D). Considering the iron limitation on phytoplankton growth, there are also two nutrient components: bioavailable iron incorporated within phytoplankton cells (<inline-formula>
<mml:math display="inline" id="im9">
<mml:mrow>
<mml:msub>
<mml:mi>F</mml:mi>
<mml:mi>p</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>) and dissolved iron concentration (<inline-formula>
<mml:math display="inline" id="im10">
<mml:mrow>
<mml:msub>
<mml:mi>F</mml:mi>
<mml:mi>d</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>).</p>
<p>Initial and boundary conditions for nitrate are obtained from the WOA. As for the phytoplankton, phytoplankton-associated iron, zooplankton and detritus concentration, their values are estimated from the seasonal surface chlorophyll concentrations derived from SeaWiFS under the assumption of constant ratios of 0.4/1 for phytoplankton/chlorophyll, 0.01/1 for phytoplankton-associated iron/chlorophyll, 0.1/1 for zooplankton/chlorophyll, and 0.15/1 for detritus/chlorophyll (<xref ref-type="bibr" rid="B47">Penven et&#xa0;al., 2008</xref>). The available dissolved iron concentration is set to 0.02 mmolFe m<sup>&#x2212;3</sup>. The boundary conditions for all these biogeochemical tracers are the same as temperature and salinity.</p>
</sec>
<sec id="s2_1_3">
<label>2.1.3</label>
<title>Coupling between ocean physics and ecology</title>
<p>In the present study, the default shortwave transmission scheme is replaced by a CHL-dependent scheme, allowing to represent bio-feedback. Specifically, an optics scheme that parameterizes the effect of chlorophyll distribution on solar radiation penetration is employed as a single-exponential function (<xref ref-type="bibr" rid="B10">Fennel et&#xa0;al., 2006</xref>):</p>
<disp-formula id="eq2">
<label>(2)</label>
<mml:math display="block" id="M2">
<mml:mrow>
<mml:mi>I</mml:mi>
<mml:mo stretchy="false">/</mml:mo>
<mml:msub>
<mml:mi>I</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:mi>&#x3b3;</mml:mi>
<mml:mo>&#xb7;</mml:mo>
<mml:mi>exp</mml:mi>
<mml:mrow>
<mml:mo>{</mml:mo>
<mml:mrow>
<mml:mi>z</mml:mi>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mi>K</mml:mi>
<mml:mi>w</mml:mi>
</mml:msub>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>K</mml:mi>
<mml:mrow>
<mml:mi>c</mml:mi>
<mml:mi>h</mml:mi>
<mml:mi>l</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mstyle displaystyle="true">
<mml:mrow>
<mml:msubsup>
<mml:mo>&#x222b;</mml:mo>
<mml:mi>z</mml:mi>
<mml:mn>0</mml:mn>
</mml:msubsup>
<mml:mrow>
<mml:mi>c</mml:mi>
<mml:mi>h</mml:mi>
<mml:mi>l</mml:mi>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>&#x3b6;</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mtext>d</mml:mtext>
<mml:mi>&#x3b6;</mml:mi>
</mml:mrow>
</mml:mrow>
</mml:mstyle>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mrow>
<mml:mo>}</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <inline-formula>
<mml:math display="inline" id="im11">
<mml:mi>&#x3b3;</mml:mi>
</mml:math>
</inline-formula> is the fraction of shortwave radiation that is photosynthetically active and equals 0.43; <inline-formula>
<mml:math display="inline" id="im12">
<mml:mrow>
<mml:msub>
<mml:mi>K</mml:mi>
<mml:mi>w</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> and <inline-formula>
<mml:math display="inline" id="im13">
<mml:mrow>
<mml:msub>
<mml:mi>K</mml:mi>
<mml:mrow>
<mml:mi>c</mml:mi>
<mml:mi>h</mml:mi>
<mml:mi>l</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> are the vertical attenuation coefficients of light for water and chlorophyll, which are set to 0.028 m<sup>&#x2212;1</sup> and 0.058 m mgChl<sup>&#x2212;1</sup>, respectively; <inline-formula>
<mml:math display="inline" id="im14">
<mml:mrow>
<mml:mi>c</mml:mi>
<mml:mi>h</mml:mi>
<mml:mi>l</mml:mi>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>&#x3b6;</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:math>
</inline-formula> is the chlorophyll concentration at a certain vertical point <inline-formula>
<mml:math display="inline" id="im15">
<mml:mrow>
<mml:mtext>&#x3b6;</mml:mtext>
<mml:mo>&#x2208;</mml:mo>
<mml:mrow>
<mml:mo stretchy="false">[</mml:mo>
<mml:mrow>
<mml:mn>0</mml:mn>
<mml:mo>,</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>z</mml:mi>
</mml:mrow>
<mml:mo stretchy="false">]</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:math>
</inline-formula>. In this model, the chlorophyll concentrations are estimated by multiplying simulated phytoplankton nitrogen concentrations by a Chl: N ratio (0.53 gChl molN<sup>&#x2212;1</sup>), which is calculated from a C: Chl ratio of 150:1 gC gChl<sup>&#x2212;1</sup> and a C: N Redfield ratio of 106:16 molC molN<sup>&#x2212;1</sup>. Note that in this function, the self-shading effect is considered, which means that light is reduced by the chlorophyll concentration itself and subsequently the phytoplankton growth is restrained.</p>
</sec>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Experimental design</title>
<p>The ocean-only simulation, initialized by using the WOA hydrography with no flow, is forced by monthly mean 10-meter wind speed, solar shortwave radiation flux, downwelling longwave radiation flux, surface air temperature, surface air specific humidity, surface air pressure and precipitation derived from the National Centers for Environmental Prediction (NCEP)/National Center for Atmospheric Research (NCAR) reanalysis product with the bulk flux algorithm (<xref ref-type="bibr" rid="B9">Fairall et&#xa0;al., 2003</xref>).</p>
<p>After the first 80-year spin-up period, the ROMS is coupled to the NPZD-IRON and is integrated from 1979 to 2019 for a real-ocean model setting. Atmospheric forcing fields (the variables are the same as the spin-up run) are extracted from the monthly average output of the European Centre for Medium-range Weather Forecasts (ECMWF) Reanalysis V5 (<xref ref-type="bibr" rid="B21">Hersbach et&#xa0;al., 2020</xref>). The model outputs are extracted from the 40-year simulation in the form of monthly mean values for the years from 1995 to 2018.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Reanalysis and observational data</title>
<p>Atmospheric fields are used to force the ocean model, including climatological monthly-mean fields from the NCEP/NCAR reanalysis product, and interannually varying monthly-mean fields from the ECMWF Reanalysis V5 (<xref ref-type="bibr" rid="B21">Hersbach et&#xa0;al., 2020</xref>). Various observational datasets are used to validate the model performance. SST data are from the Optimum Interpolation Sea Surface Temperature (OISST) (<ext-link ext-link-type="uri" xlink:href="https://psl.noaa.gov/data/gridded/data.noaa.oisst.v2.html">https://psl.noaa.gov/data/gridded/data.noaa.oisst.v2.html</ext-link>) with a spatial resolution of 1&#xb0; and a monthly temporal resolution provided by the National Climate Data Center of the National Ocean and Atmospheric Administration. Monthly climatology of surface chlorophyll concentration is the satellite-retrieved data from the SeaWiFS provided by the Asia-Pacific Data-Research Center of the International Pacific Research Center (<ext-link ext-link-type="uri" xlink:href="http://apdrc.soest.hawaii.edu/las/v6/dataset?catitem=13078">http://apdrc.soest.hawaii.edu/las/v6/dataset?catitem=13078</ext-link>).</p>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Definitions of variables of interest</title>
<sec id="s2_4_1">
<label>2.4.1</label>
<title>The euphotic depth</title>
<p>Ocean phytoplankton needs light to maintain growth and reproduction. Because of the unidirectional nature of light, the solar radiation that reaches the sea surface passes downward in a finite layer, named euphotic zone (<xref ref-type="bibr" rid="B26">Kirk, 1994</xref>), in which there is sufficient light for photosynthesis and phytoplankton growth. Usually, the euphotic depth (ED) is defined as the depth where the solar radiation declines to 1% of its value below the sea surface (<xref ref-type="bibr" rid="B25">Khanna et&#xa0;al., 2009</xref>). At the ED, thus, the decay rate of shortwave radiation equals 0.01, which can be expressed as follows:</p>
<disp-formula id="eq3">
<label>(3)</label>
<mml:math display="block" id="M3">
<mml:mrow>
<mml:msub>
<mml:mi>I</mml:mi>
<mml:mrow>
<mml:mtext>ED</mml:mtext>
</mml:mrow>
</mml:msub>
<mml:mo stretchy="false">/</mml:mo>
<mml:msub>
<mml:mi>I</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:mn>0.01</mml:mn>
</mml:mrow>
</mml:math>
</disp-formula>
</sec>
<sec id="s2_4_2">
<label>2.4.2</label>
<title>The mixed layer depth</title>
<p>Due to the convective overturning, the temperature, salinity and density are vertically uniform within the mixed layer (<xref ref-type="bibr" rid="B5">Bosc et&#xa0;al., 2009</xref>). The MLD is the main factor determining the heat content of the upper ocean and the vertical distribution of solar radiation between the mixed layer and the subsurface layer (<xref ref-type="bibr" rid="B8">de Boyer Mont&#xe9;gut, 2004</xref>). The MLD can be estimated based on the difference or gradient of potential temperature or density. Here, we define the density (<inline-formula>
<mml:math display="inline" id="im16">
<mml:mi>&#x3c1;</mml:mi>
</mml:math>
</inline-formula>)-based MLD following <xref ref-type="bibr" rid="B59">Sprintall and Tomczak (1992)</xref>, which considers both salinity and temperature effects:</p>
<disp-formula id="eq4">
<label>(4)</label>
<mml:math display="block" id="M4">
<mml:mrow>
<mml:msub>
<mml:mi>&#x3c1;</mml:mi>
<mml:mrow>
<mml:mi>M</mml:mi>
<mml:mi>L</mml:mi>
<mml:mi>D</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:mi>&#x3c1;</mml:mi>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mi>T</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
<mml:mo>+</mml:mo>
<mml:mi>&#x394;</mml:mi>
<mml:mi>T</mml:mi>
<mml:mo>,</mml:mo>
<mml:msub>
<mml:mi>S</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
<mml:mo>,</mml:mo>
<mml:msub>
<mml:mi>p</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <inline-formula>
<mml:math display="inline" id="im17">
<mml:mrow>
<mml:msub>
<mml:mi>T</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>, <inline-formula>
<mml:math display="inline" id="im18">
<mml:mrow>
<mml:msub>
<mml:mi>S</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> and <inline-formula>
<mml:math display="inline" id="im19">
<mml:mrow>
<mml:msub>
<mml:mi>p</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> are the temperature, salinity and pressure values at the sea surface, respectively; <inline-formula>
<mml:math display="inline" id="im20">
<mml:mrow>
<mml:mi>&#x394;</mml:mi>
<mml:mi>T</mml:mi>
</mml:mrow>
</mml:math>
</inline-formula> is the temperature threshold (<inline-formula>
<mml:math display="inline" id="im21">
<mml:mrow>
<mml:mi>&#x394;</mml:mi>
<mml:mi>T</mml:mi>
</mml:mrow>
</mml:math>
</inline-formula> =0.5&#xb0;C).</p>
</sec>
<sec id="s2_4_3">
<label>2.4.3</label>
<title>Ocean chlorophyll-induced heating terms (<inline-formula>
<mml:math display="inline" id="im22">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>, <inline-formula>
<mml:math display="inline" id="im23">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> and <inline-formula>
<mml:math display="inline" id="im24">
<mml:mrow>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>)</title>
<p>
<xref ref-type="bibr" rid="B29">Lewis et&#xa0;al. (1990)</xref> indicated that the amplitude of shortwave radiation flux reaching the bottom of the mixed layer is equivalent to that of the net surface heat flux, especially in the western Pacific. With the increase of chlorophyll concentration, the turbidity and absorption of shortwave radiation in the upper ocean increase, and more heat is retained in the mixed layer, further affecting SST. In this study, three chlorophyll-induced heating terms are analyzed according to <xref ref-type="bibr" rid="B67">Zhang (2015a)</xref> to quantify the direct OBH effects.</p>
<p>According to the radiation transfer function, the solar radiation that penetrates through the base of the surface mixed layer (denoted as <inline-formula>
<mml:math display="inline" id="im25">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>) can be expressed as follows:</p>
<disp-formula id="eq5">
<label>(5)</label>
<mml:math display="block" id="M5">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mtext>sr</mml:mtext>
</mml:mrow>
</mml:msub>
<mml:mo>&#xb7;</mml:mo>
<mml:mi>&#x3b3;</mml:mi>
<mml:mo>&#xb7;</mml:mo>
<mml:mi>exp</mml:mi>
<mml:mrow>
<mml:mo stretchy="false">[</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mi>m</mml:mi>
</mml:msub>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mi>K</mml:mi>
<mml:mi>w</mml:mi>
</mml:msub>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>K</mml:mi>
<mml:mrow>
<mml:mi>c</mml:mi>
<mml:mi>h</mml:mi>
<mml:mi>l</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mstyle displaystyle="true">
<mml:mrow>
<mml:msubsup>
<mml:mo>&#x222b;</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mi>m</mml:mi>
</mml:msub>
</mml:mrow>
<mml:mn>0</mml:mn>
</mml:msubsup>
<mml:mrow>
<mml:mi>c</mml:mi>
<mml:mi>h</mml:mi>
<mml:mi>l</mml:mi>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>&#x3b6;</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mtext>d</mml:mtext>
<mml:mi>&#x3b6;</mml:mi>
</mml:mrow>
</mml:mrow>
</mml:mstyle>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mrow>
<mml:mo stretchy="false">]</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <inline-formula>
<mml:math display="inline" id="im26">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mtext>sr</mml:mtext>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> is the incident solar radiation flux at the sea surface, and <inline-formula>
<mml:math display="inline" id="im27">
<mml:mrow>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mi>m</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> is the MLD.</p>
<p>The absorbed solar radiation flux within the mixed layer (denoted as <inline-formula>
<mml:math display="inline" id="im28">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>) is the difference between the solar radiation flux at the surface and at the MLD:</p>
<disp-formula id="eq6">
<label>(6)</label>
<mml:math display="block" id="M6">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
<mml:mo>&#x2212;</mml:mo>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
<mml:mo>&#xb7;</mml:mo>
<mml:mrow>
<mml:mo>{</mml:mo>
<mml:mrow>
<mml:mn>1</mml:mn>
<mml:mo>&#x2212;</mml:mo>
<mml:mi>&#x3b3;</mml:mi>
<mml:mo>&#xb7;</mml:mo>
<mml:mi>exp</mml:mi>
<mml:mrow>
<mml:mo stretchy="false">[</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mi>m</mml:mi>
</mml:msub>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mi>K</mml:mi>
<mml:mi>w</mml:mi>
</mml:msub>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>K</mml:mi>
<mml:mrow>
<mml:mi>c</mml:mi>
<mml:mi>h</mml:mi>
<mml:mi>l</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mstyle displaystyle="true">
<mml:mrow>
<mml:msubsup>
<mml:mo>&#x222b;</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mi>m</mml:mi>
</mml:msub>
</mml:mrow>
<mml:mn>0</mml:mn>
</mml:msubsup>
<mml:mrow>
<mml:mi>c</mml:mi>
<mml:mi>h</mml:mi>
<mml:mi>l</mml:mi>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>&#x3b6;</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mtext>d</mml:mtext>
<mml:mi>&#x3b6;</mml:mi>
</mml:mrow>
</mml:mrow>
</mml:mstyle>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mrow>
<mml:mo stretchy="false">]</mml:mo>
</mml:mrow>
</mml:mrow>
<mml:mo>}</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:math>
</disp-formula>
<p>Furthermore, the rate of temperature variation resulting from the <inline-formula>
<mml:math display="inline" id="im29">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> effect in the mixed layer (denoted as <inline-formula>
<mml:math display="inline" id="im30">
<mml:mrow>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>) is directly related to the <inline-formula>
<mml:math display="inline" id="im31">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> and <inline-formula>
<mml:math display="inline" id="im32">
<mml:mrow>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mi>m</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>:</p>
<disp-formula id="eq7">
<label>(7)</label>
<mml:math display="block" id="M7">
<mml:mrow>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo stretchy="false">/</mml:mo>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mi>&#x3c1;</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
<mml:msub>
<mml:mi>c</mml:mi>
<mml:mi>p</mml:mi>
</mml:msub>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mi>m</mml:mi>
</mml:msub>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <inline-formula>
<mml:math display="inline" id="im33">
<mml:mrow>
<mml:msub>
<mml:mi>&#x3c1;</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> and <inline-formula>
<mml:math display="inline" id="im34">
<mml:mrow>
<mml:msub>
<mml:mi>c</mml:mi>
<mml:mi>p</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> are respectively the mean density and specific heat capacity of seawater, whose typical values are set to <inline-formula>
<mml:math display="inline" id="im35">
<mml:mrow>
<mml:msub>
<mml:mi>&#x3c1;</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>=1025 kg m<sup>&#x2212;3</sup> and <inline-formula>
<mml:math display="inline" id="im36">
<mml:mrow>
<mml:msub>
<mml:mi>c</mml:mi>
<mml:mi>p</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>=4000 J K<sup>&#x2212;1</sup> kg<sup>&#x2212;1</sup>. Since the temperature in the mixed layer is almost identical to that at the sea surface, the temperature variation resulting from the <inline-formula>
<mml:math display="inline" id="im37">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> effect in the mixed layer (i.e., <inline-formula>
<mml:math display="inline" id="im38">
<mml:mrow>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>) can help diagnose the direct bio-effects on SST.</p>
</sec>
</sec>
</sec>
<sec id="s3">
<label>3</label>
<title>The simulated results</title>
<sec id="s3_1">
<label>3.1</label>
<title>Annual mean</title>
<p>The horizontal distributions of annual mean physical and ecological fields in the tropical Pacific are displayed in <xref ref-type="fig" rid="f1">
<bold>Figures&#xa0;1&#x2013;3</bold>
</xref>. The spatial pattern of the simulated SST (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1A</bold>
</xref>) is similar to that of observation (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1B</bold>
</xref>), and the Indo-Pacific warm pool and the eastern Pacific cold tongue can be well simulated. A detailed comparison between simulation and observation reveals that the simulated SST is slightly lower in the eastern equatorial Pacific; the differences in these simulated and observed fields can be more convincingly seen (figures not shown). Due to the effects of easterly wind above the equator which transports seawater to the west, the thermocline (the 20&#xb0;C isothermal depth) is shallow in the eastern equatorial Pacific, with the shoaled mixed layer (&lt;25 m) (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3A</bold>
</xref>). The mixed layer is deep (&gt;40 m) in the mid-basin and shallow over the western equatorial Pacific. Near 10&#xb0;N in the central-eastern Pacific, the SST is higher (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1A</bold>
</xref>) and the MLD is smaller (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3A</bold>
</xref>). This is because there is an intertropical convergence zone (<xref ref-type="bibr" rid="B6">Byrne et&#xa0;al., 2018</xref>), which is maintained by several positive feedbacks, such as the convection-shortwave flux-SST feedback and the convection-wind-evaporation-SST feedback (<xref ref-type="bibr" rid="B57">Song and Zhang, 2009</xref>). The simulated MLD (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3A</bold>
</xref>) is consistent with the corresponding observations (shown in the <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figure 1A</bold>
</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Horizontal distributions of annual-mean SST fields for <bold>(A)</bold> the model simulation and <bold>(B)</bold> the observation from OISST V2 data averaged over the years 1995-2018. The contour interval is 1 &#xb0;C.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1473208-g001.tif"/>
</fig>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Horizontal distributions of annual-mean <bold>(A)</bold> simulated and <bold>(B)</bold> satellite SeaWiFS surface chlorophyll concentrations averaged over the years 1998-2009. Note that the color mapping and the tick values along the colorbar are displayed on a logarithmic scale. The contour interval is 0.02 <italic>mg Chl m</italic>
<sup>&#x2013;3</sup>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1473208-g002.tif"/>
</fig>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Horizontal distributions of simulated annual-mean <bold>(A)</bold> mixed layer depth and <bold>(B)</bold> euphotic depth averaged over the years 1995-2018. The contour interval is 5 m.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1473208-g003.tif"/>
</fig>
<p>As shown in <xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>, by comparing it with satellite data, it is seen that the model can well capture main features of the annual mean chlorophyll concentration. For example, the chlorophyll concentration is higher in the eastern equatorial Pacific, where strong upwelling brings plenty of nutrients. In contrast, the chlorophyll concentration is low in the subtropical gyres. Note that the simulated chlorophyll concentration in the eastern equatorial Pacific is somehow overestimated. The simulated maximum value can reach 0.5 mgChl m<sup>&#x2212;3</sup>, while the maximum observation is only about 0.25 mgChl m<sup>&#x2212;3</sup>. The reason for this bias may be that the simulated upwelling is too vigorous, which results in a high nitrate concentration. The ED can be estimated from the chlorophyll concentration in the upper ocean (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2A</bold>
</xref>). The spatial pattern of the ED in <xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3B</bold>
</xref> shows a spatial structure that is opposite to the chlorophyll concentration. Shortwave radiation decays faster in the eastern Pacific because of the higher chlorophyll concentration, whereas it can penetrate more than 60 m in the subtropical gyres with little chlorophyll. The ED is less than 15 m in most areas of the equatorial Pacific, and the MLD is greater than this value except for the region to the east of 120&#xb0;W in the eastern equatorial Pacific (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3A</bold>
</xref>).</p>
<p>The aforementioned mathematical expressions in <xref ref-type="disp-formula" rid="eq5">Equations 5&#x2013;7</xref> indicate that the three heating terms are functions of shortwave radiation, the MLD and chlorophyll concentration. As can be seen in <xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>, the horizontal distribution of the annual mean <inline-formula>
<mml:math display="inline" id="im39">
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:math>
</inline-formula> is similar to that of shortwave radiation (figure omitted). <inline-formula>
<mml:math display="inline" id="im40">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> has an order of 200 W m<sup>&#x2212;2</sup>, while <inline-formula>
<mml:math display="inline" id="im41">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> is less than 30 W m<sup>&#x2212;2</sup>. This indicates that most of the incident shortwave radiation is absorbed within the mixed layer, while a little can penetrate into the subsurface. For <inline-formula>
<mml:math display="inline" id="im42">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>, regions with large values (&gt;15 W m<sup>&#x2212;2</sup>) are mainly located in the subtropical gyres where the euphotic zone is deep (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3B</bold>
</xref>), and around 10&#xb0;N where the mixed layer is shallow (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3A</bold>
</xref>). <inline-formula>
<mml:math display="inline" id="im43">
<mml:mrow>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> exhibits high values in the eastern basin near the equator and 10&#xb0;N, and its spatial pattern is similar to that of the MLD (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3A</bold>
</xref>).</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Horizontal distributions of simulated annual-mean <bold>(A)</bold> <italic>Q<sub>abs</sub>
</italic>, <bold>(B)</bold> <italic>Q<sub>pen</sub>
</italic>, and <bold>(C)</bold> <italic>R<sub>sr</sub>
</italic> averaged over the years 1995-2018. The contour interval is 10 <italic>W m</italic>
<sup>&#x2013;2</sup> in <bold>(A)</bold>, 2 <italic>W m</italic>
<sup>&#x2013;2</sup> in <bold>(B)</bold>, and 1 &#xb0;C <italic>mon</italic>
<sup>&#x2013;1</sup> in <bold>(C)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1473208-g004.tif"/>
</fig>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Seasonal variability</title>
<p>
<xref ref-type="fig" rid="f5">
<bold>Figures&#xa0;5A, B</bold>
</xref> demonstrate the annual variations (i.e., the climatological annual-mean are subtracted from climatological monthly means) of simulated and observed SST over the tropical Pacific. There is an obvious seasonal variation in the eastern equatorial Pacific, with the warmest in spring and the coldest in autumn. The simulation exhibits a phase difference by one month compared with the corresponding observation. The warming occurs in February in simulation, while it is in March in observation. Correspondingly, the simulated cooling precedes the observed cooling also for about one month. <xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6A</bold>
</xref> shows the seasonal evolution of the MLD along the equator. The simulated seasonal cycle of the MLD is in good agreement with the corresponding observation (<xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Figure 1B</bold></xref>). The mixed layer is the deepest in the central Pacific basin in summer and winter. In the eastern equatorial Pacific, the MLD simulation presents notable seasonal variations. The mixed layer is shallow in spring but deep in autumn.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Annual variations (relative to their annual mean) along the equator (5&#xb0;S&#x2013;5&#xb0;N) for simulated (left panels) and observed (right panels) <bold>(A, B)</bold> SST and <bold>(C, D)</bold> surface chlorophyll concentrations in the Pacific Ocean. The contour interval is 0.5 &#xb0;C in <bold>(A, B)</bold>, and 0.02 <italic>mg Chl m</italic>
<sup>&#x2013;3</sup> in <bold>(C, D)</bold>. The observed SST is from OISST V2 data, and the observed surface chlorophyll concentration is from SeaWiFS.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1473208-g005.tif"/>
</fig>
<fig id="f6" position="float">
<label>Figure&#xa0;6</label>
<caption>
<p>Seasonal cycles along the equator (5&#xb0;S&#x2013;5&#xb0;N) for simulated <bold>(A)</bold> mixed layer depth and <bold>(B)</bold> euphotic depth in the Pacific Ocean. The contour interval is 5 m, and the green dots indicate that the depth of the euphotic layer is greater than that of the mixed layer.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1473208-g006.tif"/>
</fig>
<p>
<xref ref-type="fig" rid="f5">
<bold>Figures&#xa0;5C, D</bold>
</xref> demonstrate the seasonal cycle of sea surface chlorophyll concentration over the tropical Pacific for simulation and observation, respectively. The annual mean is removed to avoid the influence of the overestimated chlorophyll concentration. The simulated surface chlorophyll concentration differs from observation in the eastern equatorial Pacific; only some broad features can be captured well, such as a decrease in spring and summer and an increase in autumn and winter, respectively. The seasonally varying signal starts from the South American coast and then propagates westward. Its seasonality and propagation characteristics resemble those of the SST in the eastern Pacific, suggesting that the seasonal variation in chlorophyll may be dominated by the upwelling. The ED is calculated based on the chlorophyll concentration. <xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6B</bold>
</xref> exhibits that the euphotic zone is generally shallow in the equatorial Pacific (&lt;20 m); it deepens in spring and shallows in autumn. In February, March and April, the euphotic layer over the eastern equatorial Pacific is deeper than the mixed layer (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6B</bold>
</xref>), with a reduced chlorophyll concentration (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5C</bold>
</xref>) and a shallower mixed layer (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6A</bold>
</xref>), indicating that the incident shortwave radiation penetrates through the mixed layer and reaches the deeper ocean.</p>
<p>Seasonal variations in the three biology-induced heating terms along the equator (averaged from 5&#xb0;S&#x2013;5&#xb0;N) are shown in <xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7</bold>
</xref>. The variation of <inline-formula>
<mml:math display="inline" id="im44">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> closely resembles that of shortwave radiation (figure omitted), except in the east of 120&#xb0;W from January to May when the mixed layer is shallow and the euphotic layer is deep. The shortwave radiation can penetrate below the mixed layer, resulting in <inline-formula>
<mml:math display="inline" id="im45">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> that exceeds 15 W m<sup>&#x2212;2</sup>. The seasonal variation of <inline-formula>
<mml:math display="inline" id="im46">
<mml:mrow>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> is basically opposite to that of the MLD. In the eastern equatorial Pacific, <inline-formula>
<mml:math display="inline" id="im47">
<mml:mrow>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> is greater than 10&#xb0;C month<sup>&#x2212;1</sup> throughout the year, with a notable increase from January to May, corresponding to a shallower mixed layer (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6A</bold>
</xref>). These findings indicate that these terms are predominately determined by the MLD. Nevertheless, <inline-formula>
<mml:math display="inline" id="im48">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> is also modulated by the ED, which is characterized by strong regional and seasonal dependences. For instance, <inline-formula>
<mml:math display="inline" id="im49">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> rises markedly in the eastern equatorial Pacific during late winter and early spring, coinciding with the euphotic layer that lies beneath the mixed layer (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref>). In other regions or other months, the mixed layers are so deep that the incident shortwave radiation is almost absorbed within the mixed layer, leading to <inline-formula>
<mml:math display="inline" id="im50">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> that is extremely small.</p>
<fig id="f7" position="float">
<label>Figure&#xa0;7</label>
<caption>
<p>Seasonal cycles along the equator (5&#xb0;S&#x2013;5&#xb0;N) for simulated <bold>(A)</bold> <italic>Q<sub>abs</sub>
</italic>, <bold>(B)</bold> <italic>Q<sub>pen</sub>
</italic>, and <bold>(C)</bold> <italic>R<sub>sr</sub>
</italic> fields in the Pacific Ocean. The contour interval is 10 <italic>W m</italic>
<sup>&#x2013;2</sup> in <bold>(A)</bold>, 2 <italic>W m</italic>
<sup>&#x2013;2</sup> in <bold>(B)</bold>, and 1 &#xb0;C <italic>mon</italic>
<sup>&#x2013;1</sup> in <bold>(C)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1473208-g007.tif"/>
</fig>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Interannual variability</title>
<p>
<xref ref-type="fig" rid="f8">
<bold>Figure&#xa0;8A</bold>
</xref> displays the temporal evolution of simulated SST anomalies along the equator. The simulated
interannual SST anomaly is consistent with the observation (<xref ref-type="supplementary-material" rid="SM2">
<bold>Supplementary Figure 2A</bold>
</xref>). For example, there are large warm SST anomalies during 1997/98 and 2015/16 when extreme El Ni&#xf1;o events happened. Apart from the typical eastern Pacific (EP) El Ni&#xf1;o events (large warm SST anomalies occur both in the eastern and central Pacific, hereinafter referred to as EP-El Ni&#xf1;o) (<xref ref-type="bibr" rid="B49">Radenac et&#xa0;al., 2012</xref>), another type of El Ni&#xf1;o, termed the central Pacific (CP) El Ni&#xf1;o events (large warm SST anomalies occur in the central Pacific, hereinafter referred to as CP-El Ni&#xf1;o) (<xref ref-type="bibr" rid="B49">Radenac et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B73">Zhang et al., 2022</xref>), is also captured during 2002/03, 2004/05 and 2009/10, respectively. The large center of interannual SST variability is located in the central-eastern equatorial Pacific. Warm SST anomalies are accompanied by a weakened upwelling in the eastern Pacific and westerly wind anomalies in the western Pacific (<xref ref-type="bibr" rid="B51">Ren and Jin, 2013</xref>). The mixed layer (<xref ref-type="fig" rid="f9">
<bold>Figure&#xa0;9A</bold>
</xref>) is anomalously deep in the eastern Pacific and anomalously shallow in the central-western
Pacific during El Ni&#xf1;o events, and it is opposite during La Ni&#xf1;a events, e.g., 1998/2000, 2007/08, and 2010/11. The interannual variability of the MLD is characterized by a see-saw pattern in the zonal direction with a zero line near 160&#xb0;W along the equator, similar to observations (<xref ref-type="supplementary-material" rid="SM4">
<bold>Supplementary Figure 1C</bold>
</xref>).</p>
<fig id="f8" position="float">
<label>Figure&#xa0;8</label>
<caption>
<p>Longitude-time sections along the equator (5&#xb0;S&#x2013;5&#xb0;N) for simulated interannual anomalies of <bold>(A)</bold> SST and <bold>(B)</bold> surface chlorophyll concentrations in the Pacific Ocean. The contour interval is 1 &#xb0;C in <bold>(A)</bold>, and 0.05 <italic>mg Chl m</italic>
<sup>&#x2013;3</sup> in <bold>(B)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1473208-g008.tif"/>
</fig>
<fig id="f9" position="float">
<label>Figure&#xa0;9</label>
<caption>
<p>Longitude-time sections along the equator (5&#xb0;S&#x2013;5&#xb0;N) for simulated interannual anomalies of <bold>(A)</bold> mixed layer depth and <bold>(B)</bold> euphotic depth in the Pacific Ocean. The contour interval is 2 m in <bold>(A, B)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1473208-g009.tif"/>
</fig>
<p>With these anomalies in physical fields, the surface chlorophyll shows a basin-scale interannual variation dominated by ENSO events (<xref ref-type="fig" rid="f8">
<bold>Figure&#xa0;8B</bold>
</xref>), similar to observations (<xref ref-type="supplementary-material" rid="SM2">
<bold>Supplementary Figure 2B</bold>
</xref>). The chlorophyll concentration decreases in the central and eastern equatorial Pacific during the 1997/98 and 2015/16 El Ni&#xf1;o events due to the suppression of the equatorial upwelling, which carries rich nutrients from deep waters. The core of the maximum negative chlorophyll anomaly is near the dateline for EP-El Ni&#xf1;o events and around 160&#xb0;E for CP-El Ni&#xf1;o. The spatial pattern of surface chlorophyll concentration during La Ni&#xf1;a events is somewhat opposite to that during CP-El Ni&#xf1;o events. In general, a coherently opposite relationship exists between chlorophyll anomalies and SST anomalies over the equatorial Pacific (<xref ref-type="bibr" rid="B70">Zhang et&#xa0;al., 2011</xref>). The interannual variability of the ED along the equator (<xref ref-type="fig" rid="f9">
<bold>Figure&#xa0;9B</bold>
</xref>) is similar to that of the surface chlorophyll concentration (<xref ref-type="fig" rid="f8">
<bold>Figure&#xa0;8B</bold>
</xref>). For example, in the western-central equatorial Pacific, the ED exhibits positive anomalies during El Ni&#xf1;o events, while chlorophyll concentration presents negative anomalies. During La Ni&#xf1;a events, the ED presents negative anomalies, while the chlorophyll concentration presents positive anomalies. In this region, the amplitude of the interannual variability of the ED is large and comparable to that of the MLD; in the eastern equatorial Pacific, it is much smaller than that of the MLD (<xref ref-type="fig" rid="f9">
<bold>Figure&#xa0;9</bold>
</xref>).</p>
<p>
<xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10</bold>
</xref> displays interannual variations in <inline-formula>
<mml:math display="inline" id="im51">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>, <inline-formula>
<mml:math display="inline" id="im52">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> and <inline-formula>
<mml:math display="inline" id="im53">
<mml:mrow>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> along the equator. The interannual anomaly of <inline-formula>
<mml:math display="inline" id="im54">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> (<xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10A</bold>
</xref>) is consistent with that of the shortwave radiation (figure omitted) in most areas, with only minor differences to the east of 130&#xb0;W. <inline-formula>
<mml:math display="inline" id="im55">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> exhibits two variability centers (<xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10B</bold>
</xref>): one is located in the western-central equatorial Pacific, with variations similar to those of the ED (<xref ref-type="fig" rid="f9">
<bold>Figure&#xa0;9B</bold>
</xref>); the other is located in the eastern equatorial Pacific, with significant variations only in late winter and early spring. Focusing on the equatorial western-central Pacific, it is found that the maximum interannual variability of the ED is also located in this region, with its amplitude being comparable to that of the MLD (<xref ref-type="fig" rid="f9">
<bold>Figure&#xa0;9</bold>
</xref>). In this region, the interannual variability of the ED (chlorophyll) exerts an asymmetric modulating effect on <inline-formula>
<mml:math display="inline" id="im56">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>, which is out-of-phase (in-phase) with that of the mixed layer depth. During El Ni&#xf1;o events, the mixed layer becomes shallow, which leads to a decrease in <inline-formula>
<mml:math display="inline" id="im57">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> and an increase in <inline-formula>
<mml:math display="inline" id="im58">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> (<xref ref-type="fig" rid="f9">
<bold>Figures&#xa0;9</bold>
</xref>, <xref ref-type="fig" rid="f10">
<bold>10</bold>
</xref>). Meanwhile, the ED increases as the chlorophyll decreases, and can even exceed the MLD
(<xref ref-type="supplementary-material" rid="SM3">
<bold>Supplementary Figure 3</bold>
</xref>). In this case, <inline-formula>
<mml:math display="inline" id="im59">
<mml:mrow>
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:mrow>
</mml:math>
</inline-formula> becomes pronouncedly larger, and the positive ED anomaly (the negative chlorophyll anomaly) acts to enhance a positive <inline-formula>
<mml:math display="inline" id="im60">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> anomaly. During La Ni&#xf1;a events, the depth variations of the mixed layer and the euphotic layer are opposite. The negative ED anomaly (the positive chlorophyll anomaly) should theoretically enhance the negative <inline-formula>
<mml:math display="inline" id="im61">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> anomaly. However, since the depth of the euphotic layer here is smaller than that of the mixed layer, which means that almost all of the shortwave radiation is absorbed within the euphotic layer, and very little shortwave radiation penetrates below the mixed layer (i.e., the <inline-formula>
<mml:math display="inline" id="im62">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> is extremely small) (<xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10B</bold>
</xref>). Therefore, the changes in <inline-formula>
<mml:math display="inline" id="im63">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> are not significant, and the variations in the ED (chlorophyll) have little effects on <inline-formula>
<mml:math display="inline" id="im64">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>. <inline-formula>
<mml:math display="inline" id="im65">
<mml:mrow>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> is also a function of both MLD and <inline-formula>
<mml:math display="inline" id="im66">
<mml:mrow>
<mml:msub>
<mml:mi>Q</mml:mi>
<mml:mrow>
<mml:mstyle>
<mml:mi>a</mml:mi>
<mml:mi>b</mml:mi>
<mml:mi>s</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>, with the latter being also related to the MLD. Comparing <xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10C</bold>
</xref> with <xref ref-type="fig" rid="f9">
<bold>Figure&#xa0;9</bold>
</xref>, it is evident that the interannual variations of <inline-formula>
<mml:math display="inline" id="im67">
<mml:mrow>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> are more effectively affected by the MLD.</p>
<fig id="f10" position="float">
<label>Figure&#xa0;10</label>
<caption>
<p>Longitude-time sections along the equator (5&#xb0;S&#x2013;5&#xb0;N) for simulated interannual anomalies of <bold>(A)</bold> <italic>Q<sub>abs</sub>
</italic>, <bold>(B)</bold> <italic>Q<sub>pen</sub>
</italic>, and <bold>(C)</bold> <italic>R<sub>sr</sub>
</italic> fields in the Pacific Ocean. The contour interval is 5 <italic>W m</italic>
<sup>&#x2013;2</sup> in <bold>(A, B)</bold> and 1 &#xb0;C <italic>mon</italic>
<sup>&#x2013;1</sup> in <bold>(C)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1473208-g010.tif"/>
</fig>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<p>The ROMS employs a stretched terrain-following coordinate system, which not only has the benefit of a &#x3c3;-coordinate system, but also enables further refinement near the surface or the bottom boundary regions. The ROMS, as a state-of-the-art ocean dynamical model that allows higher resolution and better adaptability to coastal boundaries, has been extensively utilized by the scientific community, but it is often limited to regional-scale applications. Here, we apply the ROMS to a basinwide tropical Indo-Pacific Ocean to achieve a two-way coupling between its physical and ecological components, which can represent interactive impacts of the marine ecological fields on the physical fields.</p>
<p>As stated in the introduction, bio-modeling in the tropical Pacific with differently formulated OGCMs is clearly needed to explore interactions between ocean biogeochemistry and the climate system in the tropical Pacific. The ROMS supplies several ecological modules, including the NPZD-IRON ecological model, which can be activated in modeling experiments. Due to the simple variable selections of the ecological model, it is economical in terms of computations. It should be noted that the coupling between the ROMS and ecological model in its default setting is still one-way from physical side to ecological side, not the other way around. Thus, in most previous studies, ecological models based on the ROMS are often used to study marine ecological dynamics. In this study, the coupling between the ROMS and the NPZD-IRON has been expanded from one-way to two-way by modifying the default shortwave penetration parameterization to reflect the influences of the ocean ecological fields on the physical fields.</p>
<p>In order to simplify the conversion of phytoplankton biomass into chlorophyll concentration, this study still uses a fixed C:Chl ratio; the NPZD-IRON model used in this paper is still at a lower trophic level with only two limiting nutrients and one size class for phytoplankton and zooplankton (<xref ref-type="bibr" rid="B11">Fiechter et&#xa0;al., 2009</xref>). As noted by <xref ref-type="bibr" rid="B64">Wang et&#xa0;al. (2009)</xref>, the C:Chl ratio can vary with the irradiance, temperature, nitrate and iron concentrations. Since the ROMS provides a variety of ecological modules, other more complex ecological models can be selected and tested in the future according to actual needs, including a way to represent a varying C: Chl ratio. Nevertheless, results from this modeling activity indicate that the model is sufficiently realistic for qualitative investigation since it can well reproduce the spatial pattern of chlorophyll, such as the triangular high-value region in the eastern Pacific. The ROMS-based ocean physical-ecological coupled model provides an alternative flexible vertical layering method for successful applications, and its results can be cross-validated with other ocean models (<xref ref-type="bibr" rid="B43">Nakamoto et&#xa0;al., 2001</xref>; <xref ref-type="bibr" rid="B42">Murtugudde et&#xa0;al., 2002</xref>; <xref ref-type="bibr" rid="B36">Manizza, 2005</xref>; <xref ref-type="bibr" rid="B32">Lin et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B35">L&#xf6;ptien et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B68">Zhang, 2015b</xref>).</p>
<p>Additionally, this study analyzes three heating terms that are related to chlorophyll and the MLD. It is generally recognized that chlorophyll absorbs solar radiation in seawater, but its impact on the ocean remains intricate and is not fully comprehended. For instance, the chlorophyll absorbs solar radiation and heats seawater directly. Nonetheless, the SST variation induced by the chlorophyll does not always increase (<xref ref-type="bibr" rid="B43">Nakamoto et&#xa0;al., 2001</xref>; <xref ref-type="bibr" rid="B36">Manizza, 2005</xref>; <xref ref-type="bibr" rid="B2">Anderson et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B32">Lin et&#xa0;al., 2007</xref>, <xref ref-type="bibr" rid="B33">2008</xref>; <xref ref-type="bibr" rid="B3">Anderson et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B17">Gnanadesikan and Anderson, 2009</xref>; <xref ref-type="bibr" rid="B35">L&#xf6;ptien et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B23">Jochum et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B44">Park et&#xa0;al., 2014</xref>). To better understand the CHL effects on the upper ocean, <xref ref-type="bibr" rid="B4">Ballabrera-Poy et&#xa0;al. (2007)</xref> performed a Taylor expansion of the equation for the absorbed radiation heat budget in the mixed layer; when considering only the first term, they pointed out that the sign of <italic>Q<sub>abs</sub>
</italic> is determined by the relative amplitude of the direct term (variation in the transparency) and indirect term (variation in the MLD), respectively. By analyzing the heating terms associated with the MLD, it is feasible to isolate and quantitatively diagnose the direct OBH effects on marine physical fields.</p>
<p>Moreover, the ED is analyzed in this study to represent the bio-effects compared with the penetration depth (H<sub>p</sub>; <xref ref-type="bibr" rid="B70">Zhang et&#xa0;al., 2011</xref>). The physical meaning of the ED is more apparent than the penetration depth (also called e-fold depth, a mathematically derived concept). Below the euphotic layer, there is little shortwave radiation, and phytoplankton is difficult to flourish. The relative positions of the euphotic layer and the mixed layer in the vertical direction are important factor affecting the variations in the related bio-heating terms and their effects on ocean thermodynamics and dynamics.</p>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusion</title>
<p>In this study, a two-way coupling between ocean physical-ecological models is constructed within the ROMS, providing a useful numerical modeling tool to represent chlorophyll-induced heating effects and the interactions between ocean physical and biogeochemistry processes in the tropical Indo-Pacific region at the basin scale. The comparison between the model output and observation shows that the model is capable of reproducing the spatial structure and temporal variations in ocean physical and ecological fields. For example, the simulated variations of SST and the MLD are consistent with observations. In addition, sea surface chlorophyll anomalies are out-of-phase with SST anomalies during the ENSO events, which reflects a coherent relationship between ecological and physical fields. These outcomes indicate that the model can well reproduce the spatial pattern of chlorophyll in the eastern Pacific.</p>
<p>Processes underlying the effects are analyzed. Three heating terms are calculated to quantitatively diagnose the direct heating effects induced by ocean biology. The results show that the mixed layer depth plays a dominant role in the allocation of solar radiation between the mixed layer and subsurface. In addition, the ED also has a modulation effect on penetrative solar radiation, especially in the central-western equatorial Pacific, where the interannual variability of the ED exhibits a comparable amplitude to that of the MLD. This modulation is asymmetric and more pronounced during El Ni&#xf1;o events relative to La Ni&#xf1;a events. As the chlorophyll decreases, the euphotic layer deepens and its maximum depth lies below the mixed layer. The shortwave radiation penetrating below the mixed layer increases significantly. In contrast, the structure and variation of <inline-formula>
<mml:math display="inline" id="im69">
<mml:mrow>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>r</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> are less affected by the ED, and are dominated by the MLD. These findings are consistent with those based on observational analyses (<xref ref-type="bibr" rid="B77">Zhang et&#xa0;al., 2019b</xref>).</p>
<p>There are obvious discrepancies between the simulated and observed chlorophyll fields in the tropical Pacific. For example, the simulated annual-mean chlorophyll is relatively higher than the observed in the eastern equatorial Pacific, which is due to the overestimation of upwelling and nutrients. This can also result from the simplicity of the components and marine food web in the ecological model used in this study. Since the ROMS also provides a variety of ecological modules, other more complex ecological submodels can be selected and tested to better represent CHL fields and explore its effects on the climate system in the tropical Pacific.</p>
<p>More numerical experiments based on this model are under way to better represent and understand how chlorophyll influences the physical environment, including the climate system over the tropical Pacific. For example, it has been planned to integrate this ROMS-based ocean model with a statistical atmospheric model to form a hybrid coupled model (HCM), which can achieve a multi-way coupling not only between its physical and ecological model components, but also between the atmosphere and ocean, representing interactive impacts of the marine ecological field on the physical field; such a constructed HCM can have a variety of applications. For instance, previous research suggested that the ocean-atmosphere interactions tend to increase the uncertainty in representing these bio-effects. So, the model can be used to examine the modulating impacts of chlorophyll interannual variability on ENSO in the tropical Pacific; it can be used to quantify how the ocean-atmosphere coupling intensity influences the chlorophyll-induced SST variation in the tropical Pacific, and so on. Additionally, the model covers both the Indian Ocean and the Pacific Ocean, and further research should be undertaken to find out the remote inter-basin effects from the Indian Ocean on conditions in the Pacific basin. Also, comparisons with other modeling results can be made with different model configurations (<xref ref-type="bibr" rid="B66">Zeng et al., 1991</xref>; <xref ref-type="bibr" rid="B71">Zhang and Endoh,1992</xref>; <xref ref-type="bibr" rid="B62">Timmerman and Jin, 2002</xref>; <xref ref-type="bibr" rid="B31">Lin et al., 2011</xref>).</p>
</sec>
</body>
<back>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation.</p>
</sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>WZ: Formal analysis, Investigation, Methodology, Software, Validation, Visualization, Writing &#x2013; original draft. CG: Funding acquisition, Methodology, Writing &#x2013; review &amp; editing. FT: Investigation, Methodology, Writing &#x2013; review &amp; editing. YY: Methodology, Writing &#x2013; review &amp; editing. HW: Funding acquisition, Supervision, Writing &#x2013; review &amp; editing. R-HZ: Conceptualization, Funding acquisition, Methodology, Supervision, Writing &#x2013; review &amp; editing.</p>
</sec>
<sec id="s8" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research, authorship, and/or publication of this article. This work is supported by the Strategic Priority Research Program of the Chinese Academy of Sciences (Grant No. XDB42000000), Laoshan Laboratory (No. LSKJ202202404), the National Natural Science Foundation of China (NSFC; Grant Nos. 42176032, 42030410), the Startup Foundation for Introducing Talent of NUIST, and Jiangsu Innovation Research Group (JSSCTD 202346).</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>The authors wish to thank the three reviewers for their comments that helped to improve the original manuscript.</p>
</ack>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s10" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s11" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fmars.2024.1473208/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fmars.2024.1473208/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="DataSheet1.docx" id="SM1" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document">
<label>Supplementary Figure&#xa0;1</label>
<caption>
<p>
<bold>(A)</bold> Horizontal distributions of observed annual-mean MLD in the tropical Pacific Ocean averaged over the years 2001-2018; <bold>(B)</bold> Seasonal cycles along the equator (5&#xb0;S&#x2013;5&#xb0;N) for observed MLD; <bold>(C)</bold> Longitude-time sections along the equator for observed interannual anomalies of MLD. The data are obtained from the Argo. The contour interval is 2 m in <bold>(A)</bold>, 5 m in <bold>(B)</bold>, and 2 m in <bold>(C)</bold>.</p>
</caption>
</supplementary-material>
<supplementary-material xlink:href="DataSheet1.docx" id="SM2" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document">
<label>Supplementary Figure&#xa0;2</label>
<caption>
<p>
<bold>(A)</bold> Longitude-time sections along the equator (5&#xb0;S&#x2013;5&#xb0;N) for observed interannual anomalies of SST; <bold>(B)</bold> Longitude-time sections along the equator for observed interannual anomalies of surface chlorophyll concentrations. The observed SST data are from the OISST V2 data and the observed surface chlorophyll concentration data are from the SeaWiFS. The contour interval is 1 &#xb0;C in <bold>(A)</bold> and 0.05 <italic>mg Chl m</italic>
<sup>&#x2013;3</sup> in <bold>(B)</bold>.</p>
</caption>
</supplementary-material>
<supplementary-material xlink:href="DataSheet1.docx" id="SM3" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document">
<label>Supplementary Figure&#xa0;3</label>
<caption>
<p>Longitude-time sections along the equator for simulated interannual <bold>(A)</bold> mixed layer depth and <bold>(B)</bold> euphotic depth. The contour interval is 5 m, and the green dots indicate that the depth of the euphotic layer is greater than that of the mixed layer.</p>
</caption>
</supplementary-material>
<supplementary-material xlink:href="DataSheet1.docx" id="SM4" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document">
<label>Supplementary Figure&#xa0;4</label>
<caption>
<p>Longitude-time sections along the equator for observed interannual anomalies of mixed layer depth in the Pacific Ocean. The data are obtained from the Argo. The contour interval is 2 m.</p>
</caption>
</supplementary-material>
</sec>
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