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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.2021.747763</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>Comparison of Primary Production Using <italic>in situ</italic> and Satellite-Derived Values at the SEATS Station in the South China Sea</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name><surname>Shih</surname> <given-names>Yung-Yen</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="http://loop.frontiersin.org/people/844529/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Shiah</surname> <given-names>Fuh-Kwo</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1061978/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Lai</surname> <given-names>Chao-Chen</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/987644/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Chou</surname> <given-names>Wen-Chen</given-names></name>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/640010/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Tai</surname> <given-names>Jen-Hua</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/853195/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Wu</surname> <given-names>Yu-Shun</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1476027/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Lai</surname> <given-names>Cheng-Yang</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1444890/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Ko</surname> <given-names>Chia-Ying</given-names></name>
<xref ref-type="aff" rid="aff6"><sup>6</sup></xref>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name><surname>Hung</surname> <given-names>Chin-Chang</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/147929/overview"/>
</contrib>
</contrib-group>
<aff id="aff1"><sup>1</sup><institution>Department of Applied Science, R.O.C. Naval Academy</institution>, <addr-line>Kaohsiung</addr-line>, <country>Taiwan</country></aff>
<aff id="aff2"><sup>2</sup><institution>Department of Oceanography, National Sun Yat-sen University</institution>, <addr-line>Kaohsiung</addr-line>, <country>Taiwan</country></aff>
<aff id="aff3"><sup>3</sup><institution>Research Center for Environmental Changes, Academia Sinica</institution>, <addr-line>Taipei</addr-line>, <country>Taiwan</country></aff>
<aff id="aff4"><sup>4</sup><institution>Institute of Marine Environment and Ecology, National Taiwan Ocean University</institution>, <addr-line>Keelung</addr-line>, <country>Taiwan</country></aff>
<aff id="aff5"><sup>5</sup><institution>Center of Excellence for the Oceans, National Taiwan Ocean University</institution>, <addr-line>Keelung</addr-line>, <country>Taiwan</country></aff>
<aff id="aff6"><sup>6</sup><institution>Institute of Fisheries Science, National Taiwan University</institution>, <addr-line>Taipei</addr-line>, <country>Taiwan</country></aff>
<author-notes>
<fn fn-type="edited-by"><p>Edited by: Bernardo Antonio Perez Da Gama, Fluminense Federal University, Brazil</p></fn>
<fn fn-type="edited-by"><p>Reviewed by: Hartmut Schulz, University of Tuebingen, Germany; David Michael Karl, University of Hawaii, United States</p></fn>
<corresp id="c001">&#x002A;Correspondence: Chin-Chang Hung, <email>cchung@mail.nsysu.edu.tw</email></corresp>
<fn fn-type="other" id="fn004"><p>This article was submitted to Marine Ecosystem Ecology, a section of the journal Frontiers in Marine Science</p></fn>
</author-notes>
<pub-date pub-type="epub">
<day>04</day>
<month>10</month>
<year>2021</year>
</pub-date>
<pub-date pub-type="collection">
<year>2021</year>
</pub-date>
<volume>8</volume>
<elocation-id>747763</elocation-id>
<history>
<date date-type="received">
<day>26</day>
<month>07</month>
<year>2021</year>
</date>
<date date-type="accepted">
<day>02</day>
<month>09</month>
<year>2021</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2021 Shih, Shiah, Lai, Chou, Tai, Wu, Lai, Ko and Hung.</copyright-statement>
<copyright-year>2021</copyright-year>
<copyright-holder>Shih, Shiah, Lai, Chou, Tai, Wu, Lai, Ko and Hung</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>Satellite-based observations of primary production (PP) are broadly used to assess carbon fixation rate of phytoplankton in the global ocean with small spatiotemporal limitations. However, the remote sensing can only reach the ocean surface, the assumption of a PP vertically exponential decrease with increasing depth from the surface to the bottom of euphotic zone may cause a substantial and potential discrepancy between <italic>in situ</italic> measurements and satellite-based observations of PP. This study compared euphotic zone integrated PP derived from measurements based on ship-based <italic>in situ</italic> incubation (i.e., PP<sub><italic>in situ</italic></sub>) and those derived from the satellite-based vertically generalized production model (VGPM; PP<sub>VGPM</sub>) for the period 2003&#x223C;2016 at the South East Asian Time-series Study (SEATS) station. PP values obtained during the NE-monsoon (NEM: Nov&#x223C;Mar; PP<sub><italic>in situ</italic></sub> = 323 &#x00B1; 134; PP<sub>VGPM</sub> = 443 &#x00B1; 142 mg-C m<sup>&#x2013;2</sup> d<sup>&#x2013;1</sup>) were &#x223C;2-fold higher than those recorded during the SW-monsoon (SWM: Apr&#x223C;Oct; PP<sub><italic>in situ</italic></sub> = 159 &#x00B1; 58; PP<sub>VGPM</sub> = 250 &#x00B1; 36 mg-C m<sup>&#x2013;2</sup> d<sup>&#x2013;1</sup>), regardless of the method used for derivation. The main reason for the higher PP values during the NEM appears to have been a greater abundance of inorganic nutrients were made available by vertical advection. Note that on average, PP<sub><italic>in situ</italic></sub> estimates were &#x223C;50% lower than PP<sub>VGPM</sub> estimates, regardless of the monsoon. These discrepancies can be mainly attributed to differences from the euphotic zone depth between satellite-based and <italic>in situ</italic> measurements. The significantly negative relationship between PP measurements obtained <italic>in situ</italic> and sea surface temperatures observed throughout this study demonstrates that both methods are effective indicators in estimating PP. Overall, our PP<sub><italic>in situ</italic></sub> analysis indicates that a warming climate is unfavorable for primary production in low-latitude open ocean ecosystems.</p>
</abstract>
<kwd-group>
<kwd>carbon fixation rate</kwd>
<kwd>remote sensing</kwd>
<kwd>time-series study</kwd>
<kwd>global warming</kwd>
<kwd>low-latitude ocean</kwd>
<kwd>VGPM</kwd>
</kwd-group>
<contract-sponsor id="cn001">Ministry of Science and Technology<named-content content-type="fundref-id">10.13039/501100003711</named-content></contract-sponsor>
<counts>
<fig-count count="7"/>
<table-count count="2"/>
<equation-count count="3"/>
<ref-count count="85"/>
<page-count count="16"/>
<word-count count="13112"/>
</counts>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="S1">
<title>Introduction</title>
<p>Primary production at the bottom of the marine food web plays a key role in the ocean ecosystem (<xref ref-type="bibr" rid="B62">Pauly and Christensen, 1995</xref>) and represents a major pathway for sequestration and/or cycling of atmospheric CO<sub>2</sub> by the oceans. However, this process is highly susceptible to environmental and climatic changes (<xref ref-type="bibr" rid="B6">Buitenhuis et al., 2013</xref>; <xref ref-type="bibr" rid="B40">Hung et al., 2013</xref>, <xref ref-type="bibr" rid="B35">2016</xref>; <xref ref-type="bibr" rid="B55">Liu et al., 2021</xref>; <xref ref-type="bibr" rid="B84">Zhong et al., 2021</xref>). It is important for oceanographers to gain a comprehensive understanding of the spatiotemporal characteristics of primary production (<xref ref-type="bibr" rid="B24">Field et al., 1998</xref>; <xref ref-type="bibr" rid="B7">Campbell et al., 2002</xref>; <xref ref-type="bibr" rid="B76">Tang et al., 2008</xref>; <xref ref-type="bibr" rid="B78">Tilstone et al., 2015</xref>).</p>
<p>The quantification of oceanic primary production is generally based on <italic>in situ</italic> measurements pertaining to incubation wherein the euphotic zone integrated primary production (PP) is calculated <italic>via</italic> trapezoidal integration (i.e., the PP inventory is evaluated by integrating PP divided into small trapezoids from the surface to the bottom of the euphotic zone). Measurements of PP by the radiolabeled carbon (C-14) uptake method has been extensively used in marine environments since the introduction of this method to determine the carbon uptake rate of phytoplankton (<xref ref-type="bibr" rid="B58">Nielsen, 1952</xref>; <xref ref-type="bibr" rid="B30">Hama et al., 1983</xref>; <xref ref-type="bibr" rid="B26">Gong, 1993</xref>; <xref ref-type="bibr" rid="B65">Shiah et al., 2000</xref>). Despite considerable research in the western North Pacific (WNP), i.e., East China Sea, South China Sea (SCS), and Taiwan Strait, to establish phytoplankton carbon fixation rates, researchers continue to debate whether phytoplankton growth conditions at the surface are indicative of conditions in deeper waters. Obtaining measurements of temperature at depth is straightforward; however, the <italic>in situ</italic> observation of PP (PP<sub><italic>in situ</italic></sub>) requires considerable effort in terms of manpower and time. Specifically, seawater samples collected at discrete depths must be incubated on deck within an incubator using running surface water for cooling (<xref ref-type="bibr" rid="B65">Shiah et al., 2000</xref>, <xref ref-type="bibr" rid="B66">2003</xref>, <xref ref-type="bibr" rid="B67">2005</xref>; <xref ref-type="bibr" rid="B14">Chen, 2005</xref>; <xref ref-type="bibr" rid="B48">Lai et al., 2014</xref>; <xref ref-type="bibr" rid="B12">Chen et al., 2016</xref>). Note also that gaps in PP<sub><italic>in situ</italic></sub> coverage inevitably lead to discrepancies in corresponding estimates. The first empirical algorithm for PP predictions based on remote sensing was proposed by <xref ref-type="bibr" rid="B2">Balch et al. (1989)</xref>. <xref ref-type="bibr" rid="B4">Behrenfeld and Falkowski (1997)</xref> developed an attractive alternative approach to estimating global PP using a small number of inputs. Their vertically generalized production model (VGPM; PP<sub>VGPM</sub>) is widely regarded as the most highly optimized yet usable methods for PP estimation (<xref ref-type="bibr" rid="B43">Kameda and Ishizaka, 2005</xref>; <xref ref-type="bibr" rid="B81">Yamada et al., 2005</xref>; <xref ref-type="bibr" rid="B42">Ishizaka et al., 2007</xref>; <xref ref-type="bibr" rid="B33">Hill and Zimmerman, 2010</xref>).</p>
<p>The semi-analytical VGPM uses satellite-based data as an input to calculate PP. Thus, it is conceivable that data measured <italic>in situ</italic> could be used as an alternative to satellite-based input data, and vice versa (<xref ref-type="bibr" rid="B4">Behrenfeld and Falkowski, 1997</xref>; <xref ref-type="bibr" rid="B33">Hill and Zimmerman, 2010</xref>). The VGPM can be used to derive PP using satellite-based observations, such as remote passive ocean color, sea surface temperature (SST), surface optimal carbon fixation rate (<inline-formula><mml:math id="INEQ1"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula>) per unit chlorophyll <italic>a</italic> (Chl) of the euphotic zone (Z<sub>eu</sub>), surface Chl concentration (Chl<sub>s</sub>), surface light intensity (E<sub>0</sub>), and the surface light diffuse attenuation coefficient (K<sub>d</sub>) (<xref ref-type="bibr" rid="B81">Yamada et al., 2005</xref>; <xref ref-type="bibr" rid="B42">Ishizaka et al., 2007</xref>). Nonetheless, it is still problematic whether the assumption based on peak values of PP and Chl<sub>s</sub> occur at the surface and decrease exponentially with depth until reaching the bottom of the Z<sub>eu</sub> is true (<xref ref-type="bibr" rid="B33">Hill and Zimmerman, 2010</xref>; <xref ref-type="bibr" rid="B6">Buitenhuis et al., 2013</xref>). Furthermore, the reliability of VGPM in evaluating PP diminishes when geographic features, vertical hydrographic distributions, regional characteristics, extreme weather events, and climate change are taken into consideration (<xref ref-type="bibr" rid="B21">Dierssen, 2010</xref>; <xref ref-type="bibr" rid="B25">Friedland et al., 2012</xref>; <xref ref-type="bibr" rid="B40">Hung et al., 2013</xref>, <xref ref-type="bibr" rid="B35">2016</xref>; <xref ref-type="bibr" rid="B9">Chen et al., 2015</xref>; <xref ref-type="bibr" rid="B69">Shih et al., 2015</xref>, <xref ref-type="bibr" rid="B71">2020b</xref>). In the absence of a robust estimation method, the results derived from PP<sub>VGPM</sub> cannot be relied upon to reflect the actual situation throughout the oceans. Empirical models developed by <xref ref-type="bibr" rid="B23">Dunne et al. (2005)</xref>; <xref ref-type="bibr" rid="B49">Laws et al. (2011)</xref>, and <xref ref-type="bibr" rid="B32">Henson et al. (2011)</xref> are widely used by oceanographers to estimate particulate organic carbon (POC) export flux; however, reliance on PP<sub>VGPM</sub> also calls into question all corresponding estimates pertaining to global carbon export flux and oceanic carbon sequestration.</p>
<p>The South East Asian Time-series Study (SEATS) conducted in the South China Sea (SCS) was the lowest latitude time-series program implemented during the Joint Global Ocean Flux Study era (<xref ref-type="bibr" rid="B45">Karl et al., 2003</xref>; <xref ref-type="bibr" rid="B80">Wong et al., 2007</xref>). Numerous studies have characterized the upper ocean of the SCS as stratified and oligotrophic (<xref ref-type="bibr" rid="B16">Chen et al., 2004</xref>; <xref ref-type="bibr" rid="B14">Chen, 2005</xref>; <xref ref-type="bibr" rid="B80">Wong et al., 2007</xref>). Thus, biological activity and regulation of biogeochemical responses depend heavily on dynamic perturbations, including the yearly monsoon, typhoons, storms, internal waves, Kuroshio intrusion, and atmospheric deposition as well as nutrient supply in the form of phytoplankton nitrogen fixation (<xref ref-type="bibr" rid="B54">Liu et al., 2002</xref>; <xref ref-type="bibr" rid="B19">Chou et al., 2006</xref>; <xref ref-type="bibr" rid="B18">Chen et al., 2008</xref>, <xref ref-type="bibr" rid="B10">2020</xref>; <xref ref-type="bibr" rid="B22">Du et al., 2013</xref>; <xref ref-type="bibr" rid="B82">Yang et al., 2014</xref>; <xref ref-type="bibr" rid="B51">Li et al., 2018</xref>; <xref ref-type="bibr" rid="B70">Shih et al., 2020a</xref>).</p>
<p>Throughout the SCS basin, the annual modeled PP ranges from 280 to 343 and PP<sub>VGPM</sub> ranges from 308 to 354 mg-C m <sup>&#x2013;2</sup> d<sup>&#x2013;1</sup> (<xref ref-type="table" rid="T1">Table 1</xref>; <xref ref-type="bibr" rid="B54">Liu et al., 2002</xref>; <xref ref-type="bibr" rid="B74">Tan and Shi, 2009</xref>; <xref ref-type="bibr" rid="B56">Ma et al., 2014</xref>). High PP values are generally associated with the strong NE-monsoon (NEM) system during the cold season, whereas low PP values are associated with a relative weak SW-monsoon (SWM) during the warm season (<xref ref-type="table" rid="T1">Table 1</xref>). Based on long-term satellite-based SST records, <xref ref-type="bibr" rid="B10">Chen et al. (2020)</xref> reported that declining PP can be attributed at least in part to rising SST (0.012&#x00B0;C y<sup>&#x2013;</sup><sup>1</sup>). Their and several selected researches in our study went a long way toward establishing a connection between monsoons and PP; however, all of the suppositions are based on indirect measurements; i.e., remote sensing (<xref ref-type="bibr" rid="B54">Liu et al., 2002</xref>; <xref ref-type="bibr" rid="B31">Hao et al., 2007</xref>; <xref ref-type="bibr" rid="B83">Zhao et al., 2008</xref>; <xref ref-type="bibr" rid="B74">Tan and Shi, 2009</xref>; <xref ref-type="bibr" rid="B60">Pan et al., 2012</xref>; <xref ref-type="bibr" rid="B56">Ma et al., 2014</xref>; <xref ref-type="bibr" rid="B10">Chen et al., 2020</xref>), rather than direct <italic>in situ</italic> incubation, such as the C<sup>13</sup> and C<sup>14</sup> methods (<xref ref-type="bibr" rid="B54">Liu et al., 2002</xref>; <xref ref-type="bibr" rid="B16">Chen et al., 2004</xref>, <xref ref-type="bibr" rid="B17">2007</xref>; <xref ref-type="bibr" rid="B59">Ning et al., 2004</xref>). Note also that even PP research based on <italic>in situ</italic> oceanographic analysis is limited to short-term observations rather than long-term time-series (<xref ref-type="bibr" rid="B14">Chen, 2005</xref>; <xref ref-type="bibr" rid="B11">Chen et al., 1998</xref>, <xref ref-type="bibr" rid="B8">2006</xref>; <xref ref-type="bibr" rid="B54">Liu et al., 2002</xref>; <xref ref-type="bibr" rid="B79">Tseng et al., 2005</xref>).</p>
<table-wrap position="float" id="T1">
<label>TABLE 1</label>
<caption><p>Integrated primary production in the euphotic zone (PP) based on selected studies conducted in the South China Sea.</p></caption>
<table cellspacing="5" cellpadding="5" frame="hsides" rules="groups">
<thead>
<tr>
<td valign="top" align="left">Region</td>
<td valign="top" align="center" colspan="2">Location<hr/></td>
<td valign="top" align="center">Month/period</td>
<td valign="top" align="center">PP<hr/></td>
<td valign="top" align="center">Method</td>
<td valign="top" align="center">References</td>
</tr>
<tr>
<td/>
<td valign="top" align="center">(&#x00B0;N)</td>
<td valign="top" align="center">(&#x00B0;E)</td>
<td/>
<td valign="top" align="center">(mg-C m<sup>&#x2013;2</sup> d<sup>&#x2013;1</sup>)</td>
<td/>
<td/>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.5</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Nov&#x2013;Feb</td>
<td valign="top" align="center">207</td>
<td valign="top" align="center">POC flux-estimate<xref ref-type="table-fn" rid="t1fn1">&#x002A;</xref></td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B11">Chen et al., 1998</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.5</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Mar&#x2013;May, Oct</td>
<td valign="top" align="center">59</td>
<td valign="top" align="center">POC flux-estimate<xref ref-type="table-fn" rid="t1fn1">&#x002A;</xref></td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B11">Chen et al., 1998</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.5</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Jun&#x2013;Sep</td>
<td valign="top" align="center">149</td>
<td valign="top" align="center">POC flux-estimate<xref ref-type="table-fn" rid="t1fn1">&#x002A;</xref></td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B11">Chen et al., 1998</xref></td>
</tr>
<tr>
<td valign="top" align="left">CSCS</td>
<td valign="top" align="center">14.6</td>
<td valign="top" align="center">115.1</td>
<td valign="top" align="center">Nov&#x2013;Feb</td>
<td valign="top" align="center">196</td>
<td valign="top" align="center">POC flux-estimate<xref ref-type="table-fn" rid="t1fn1">&#x002A;</xref></td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B11">Chen et al., 1998</xref></td>
</tr>
<tr>
<td valign="top" align="left">CSCS</td>
<td valign="top" align="center">14.6</td>
<td valign="top" align="center">115.1</td>
<td valign="top" align="center">Mar&#x2013;May, Oct</td>
<td valign="top" align="center">159</td>
<td valign="top" align="center">POC flux-estimate<xref ref-type="table-fn" rid="t1fn1">&#x002A;</xref></td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B11">Chen et al., 1998</xref></td>
</tr>
<tr>
<td valign="top" align="left">CSCS</td>
<td valign="top" align="center">14.6</td>
<td valign="top" align="center">115.1</td>
<td valign="top" align="center">Jun&#x2013;Sep</td>
<td valign="top" align="center">203</td>
<td valign="top" align="center">POC flux-estimate<xref ref-type="table-fn" rid="t1fn1">&#x002A;</xref></td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B11">Chen et al., 1998</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">17.9&#x2013;22.3</td>
<td valign="top" align="center">115.5&#x2013;119.8</td>
<td valign="top" align="center">Mar</td>
<td valign="top" align="center">313</td>
<td valign="top" align="center">C<sup>14</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B54">Liu et al., 2002</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">17.9&#x2013;22.3</td>
<td valign="top" align="center">115.5&#x2013;119.8</td>
<td valign="top" align="center">Mar</td>
<td valign="top" align="center">363</td>
<td valign="top" align="center">Model</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B54">Liu et al., 2002</xref></td>
</tr>
<tr>
<td valign="top" align="left">SCS</td>
<td valign="top" align="center">2.0&#x2013;24.8</td>
<td valign="top" align="center">99.0&#x2013;124.6</td>
<td valign="top" align="center">Jan&#x2013;Dec</td>
<td valign="top" align="center">354</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B54">Liu et al., 2002</xref></td>
</tr>
<tr>
<td valign="top" align="left">SCS</td>
<td valign="top" align="center">2.0&#x2013;24.8</td>
<td valign="top" align="center">99.0&#x2013;124.6</td>
<td valign="top" align="center">Jan&#x2013;Dec</td>
<td valign="top" align="center">280</td>
<td valign="top" align="center">Model</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B54">Liu et al., 2002</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">115.5</td>
<td valign="top" align="center">Mar</td>
<td valign="top" align="center">180&#x2013;330</td>
<td valign="top" align="center">C<sup>13</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B16">Chen et al., 2004</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">115.0</td>
<td valign="top" align="center">Jun&#x2013;Jul</td>
<td valign="top" align="center">228</td>
<td valign="top" align="center">C<sup>14</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B59">Ning et al., 2004</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">17.5</td>
<td valign="top" align="center">114.5</td>
<td valign="top" align="center">Nov&#x2013;Dec</td>
<td valign="top" align="center">509</td>
<td valign="top" align="center">C<sup>14</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B59">Ning et al., 2004</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">115.5</td>
<td valign="top" align="center">Mar</td>
<td valign="top" align="center">190&#x2013;350</td>
<td valign="top" align="center">C<sup>13</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B14">Chen, 2005</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">115.5</td>
<td valign="top" align="center">Jul</td>
<td valign="top" align="center">190</td>
<td valign="top" align="center">C<sup>13</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B14">Chen, 2005</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">115.5</td>
<td valign="top" align="center">Oct</td>
<td valign="top" align="center">270&#x2013;280</td>
<td valign="top" align="center">C<sup>13</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B14">Chen, 2005</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">115.5</td>
<td valign="top" align="center">Jan</td>
<td valign="top" align="center">550</td>
<td valign="top" align="center">C<sup>13</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B14">Chen, 2005</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Jan</td>
<td valign="top" align="center">300</td>
<td valign="top" align="center">C<sup>14</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B79">Tseng et al., 2005</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Feb&#x2013;Nov</td>
<td valign="top" align="center">110</td>
<td valign="top" align="center">C<sup>14</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B79">Tseng et al., 2005</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">19.0</td>
<td valign="top" align="center">118.5</td>
<td valign="top" align="center">Jan</td>
<td valign="top" align="center">684</td>
<td valign="top" align="center">Chl function</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B8">Chen et al., 2006</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">19.0</td>
<td valign="top" align="center">118.5</td>
<td valign="top" align="center">Mar</td>
<td valign="top" align="center">148</td>
<td valign="top" align="center">Chl function</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B8">Chen et al., 2006</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">19.0</td>
<td valign="top" align="center">118.5</td>
<td valign="top" align="center">May</td>
<td valign="top" align="center">275</td>
<td valign="top" align="center">Chl function</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B8">Chen et al., 2006</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">19.0</td>
<td valign="top" align="center">118.5</td>
<td valign="top" align="center">Jul</td>
<td valign="top" align="center">86</td>
<td valign="top" align="center">Chl function</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B8">Chen et al., 2006</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Mar&#x2013;May</td>
<td valign="top" align="center">310</td>
<td valign="top" align="center">C<sup>13</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B17">Chen et al., 2007</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Jul&#x2013;Aug</td>
<td valign="top" align="center">250</td>
<td valign="top" align="center">C<sup>13</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B17">Chen et al., 2007</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Oct&#x2013;Nov</td>
<td valign="top" align="center">400</td>
<td valign="top" align="center">C<sup>13</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B17">Chen et al., 2007</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Dec&#x2013;Feb</td>
<td valign="top" align="center">550</td>
<td valign="top" align="center">C<sup>13</sup> incubation</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B17">Chen et al., 2007</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0&#x2013;23.0</td>
<td valign="top" align="center">110.0&#x2013;117.0</td>
<td valign="top" align="center">Apr&#x2013;Oct</td>
<td valign="top" align="center">340</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B31">Hao et al., 2007</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0&#x2013;23.0</td>
<td valign="top" align="center">110.0&#x2013;117.0</td>
<td valign="top" align="center">Nov&#x2013;Mar</td>
<td valign="top" align="center">573</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B31">Hao et al., 2007</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.3</td>
<td valign="top" align="center">115.5</td>
<td valign="top" align="center">Jan&#x2013;Dec</td>
<td valign="top" align="center">329&#x2013;466</td>
<td valign="top" align="center">Overview</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B80">Wong et al., 2007</xref></td>
</tr>
<tr>
<td valign="top" align="left">CSCS</td>
<td valign="top" align="center">12.5&#x2013;16.5</td>
<td valign="top" align="center">112.5&#x2013;116.0</td>
<td valign="top" align="center">Nov</td>
<td valign="top" align="center">274</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B83">Zhao et al., 2008</xref></td>
</tr>
<tr>
<td valign="top" align="left">WSCS</td>
<td valign="top" align="center">13.0&#x2013;16.0</td>
<td valign="top" align="center">110.0&#x2013;113.0</td>
<td valign="top" align="center">Oct&#x2013;Nov</td>
<td valign="top" align="center">312</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B83">Zhao et al., 2008</xref></td>
</tr>
<tr>
<td valign="top" align="left">SCS</td>
<td valign="top" align="center">5.0&#x2013;18.0</td>
<td valign="top" align="center">103.0&#x2013;120.0</td>
<td valign="top" align="center">Mar&#x2013;May</td>
<td valign="top" align="center">281</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B74">Tan and Shi, 2009</xref></td>
</tr>
<tr>
<td valign="top" align="left">SCS</td>
<td valign="top" align="center">5.0&#x2013;18.0</td>
<td valign="top" align="center">103.0&#x2013;120.0</td>
<td valign="top" align="center">Jun&#x2013;Aug</td>
<td valign="top" align="center">267</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B74">Tan and Shi, 2009</xref></td>
</tr>
<tr>
<td valign="top" align="left">SCS</td>
<td valign="top" align="center">5.0&#x2013;18.0</td>
<td valign="top" align="center">103.0&#x2013;120.0</td>
<td valign="top" align="center">Sep&#x2013;Nov</td>
<td valign="top" align="center">280</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B74">Tan and Shi, 2009</xref></td>
</tr>
<tr>
<td valign="top" align="left">SCS</td>
<td valign="top" align="center">5.0&#x2013;18.0</td>
<td valign="top" align="center">103.0&#x2013;120.0</td>
<td valign="top" align="center">Dec&#x2013;Feb</td>
<td valign="top" align="center">374</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B74">Tan and Shi, 2009</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">17.5&#x2013;18.5</td>
<td valign="top" align="center">115.5&#x2013;116.5</td>
<td valign="top" align="center">May&#x2013;Oct</td>
<td valign="top" align="center">214</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B60">Pan et al., 2012</xref></td>
</tr>
<tr>
<td valign="top" align="left">SCS</td>
<td valign="top" align="center">9.0&#x2013;16.5</td>
<td valign="top" align="center">110.0&#x2013;119.0</td>
<td valign="top" align="center">Jan&#x2013;Dec</td>
<td valign="top" align="center">343</td>
<td valign="top" align="center">Model</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B56">Ma et al., 2014</xref></td>
</tr>
<tr>
<td valign="top" align="left">SCS</td>
<td valign="top" align="center">9.0&#x2013;16.5</td>
<td valign="top" align="center">110.0&#x2013;119.0</td>
<td valign="top" align="center">Jan&#x2013;Dec</td>
<td valign="top" align="center">351</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center"><xref ref-type="bibr" rid="B56">Ma et al., 2014</xref></td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Apr&#x2013;Oct</td>
<td valign="top" align="center">159 (76&#x2013;247)</td>
<td valign="top" align="center">C<sup>14</sup> incubation</td>
<td valign="top" align="center">This study</td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Nov&#x2013;Mar</td>
<td valign="top" align="center">323 (117&#x2013;528)</td>
<td valign="top" align="center">C<sup>14</sup> incubation</td>
<td valign="top" align="center">This study</td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Apr&#x2013;Oct</td>
<td valign="top" align="center">250 (154&#x2013;771)</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center">This study</td>
</tr>
<tr>
<td valign="top" align="left">NSCS</td>
<td valign="top" align="center">18.0</td>
<td valign="top" align="center">116.0</td>
<td valign="top" align="center">Nov&#x2013;Mar</td>
<td valign="top" align="center">443 (190&#x2013;1193)</td>
<td valign="top" align="center">VGPM</td>
<td valign="top" align="center">This study</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn id="t1fn1"><p><italic>NSCS, northern South China Sea; CSCS, central South China Sea; WSCS, western South China Sea. &#x002A;PP = POC flux / (17/Z + 1/100), where Z is the specific depth of the POC flux, units of POC flux and Z are (g-C m<sup>&#x2013;</sup><sup>2</sup> y<sup>&#x2013;</sup><sup>1</sup>) and (m) (<xref ref-type="bibr" rid="B11">Chen et al., 1998</xref>).</italic></p></fn>
</table-wrap-foot>
</table-wrap>
<p>Remote sensing has made it possible to conduct large-scale PP monitoring over extended durations; however, most existing sensing technology is limited to the upper ocean and there is little evidence supporting the assumption of an exponential decrease in PP as a function of depth (<xref ref-type="bibr" rid="B6">Buitenhuis et al., 2013</xref>; <xref ref-type="bibr" rid="B71">Shih et al., 2020b</xref>). In the current study, we employed data obtained in the SEATS time-series study to elucidate long-term variations in PP<sub><italic>in situ</italic></sub> under the prevailing monsoon system. We also looked for discrepancies between satellite-based and <italic>in situ</italic> observations, which could potentially influence PP estimates derived using the VGPM algorithm.</p>
</sec>
<sec id="S2" sec-type="materials|methods">
<title>Materials and Methods</title>
<sec id="S2.SS1">
<title><italic>In situ</italic> Observation</title>
<p>A total of 44 <italic>in situ</italic> observations were conducted at the SEATS site (18 &#x00B0;N, 116 &#x00B0;E) at regular intervals between 1998 and 2016. These expeditions involved hydrographic seawater sampling and biogeochemical field experiments using research vessels (<italic>RV</italic>; <italic>Ocean Researcher I</italic>, <italic>Ocean Researcher III</italic>, and <italic>Fishery Researcher I</italic>) (<xref ref-type="bibr" rid="B45">Karl et al., 2003</xref>; <xref ref-type="bibr" rid="B80">Wong et al., 2007</xref>). On-deck incubation of PP in accordance with <sup>14</sup>C protocols was also conducted during 20 of 25 expeditions between 2003 and 2016 (<xref ref-type="fig" rid="F1">Figure 1</xref>). Conductivity&#x2013;temperature&#x2013;depth (CTD) (SBE9/11, SeaBird) and quantum scalar irradiance (QSP-200L, Biospherical) were, respectively, used to obtain vertical profiles of temperature and underwater photosynthetically available radiation (PAR). A Biospherical instrument (QSR-240) was used to obtain daily PAR measurements (i.e., daily surface light intensity). In that study, Z<sub>eu</sub> was defined as the depth at which underwater PAR reached 1%. The mixed layer depth (MLD) was defined as the deepest depth at which the temperature was 0.8&#x00B0;C lower than at the surface (<xref ref-type="bibr" rid="B44">Kara et al., 2000</xref>; <xref ref-type="bibr" rid="B19">Chou et al., 2006</xref>).</p>
<fig id="F1" position="float">
<label>FIGURE 1</label>
<caption><p>Location of the South East Asian Time-series Study (SEATS) site. Inner box represents the date of cruises conducted at the SEATS site. Major and minor ticks, respectively, exhibit intervals of years and months. WNP refers to the western North Pacific.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-08-747763-g001.tif"/>
</fig>
<p>Seawater samples were collected at discrete depths to determine Chl concentrations and PP values. Briefly, 2L seawater samples were filtered using 47-mm GF/F filters before obtaining Chl concentrations (<italic>via</italic> acidification) using a Turner 10-AU-005 fluorometer (<xref ref-type="bibr" rid="B28">Gong et al., 2003</xref>; <xref ref-type="bibr" rid="B66">Shiah et al., 2003</xref>). PP was measured using the <sup>14</sup>C assimilation method (<xref ref-type="bibr" rid="B61">Parsons et al., 1984</xref>) following incubation under an artificial light source (2,000 &#x03BC;E m<sup>&#x2013;</sup><sup>2</sup> s<sup>&#x2013;</sup><sup>1</sup> with full spectrum from 350 to 2450 nm, similar to sunlight) for &#x223C;3 h in a proprietary isolation container using flowing surface seawater for cooling. Following incubation and acidification (0.5 N HCl), radioactive substances collected in 25-mm GF/F filters were transported to a lab for quantification using a scintillation counter (Packard 2200) (<xref ref-type="bibr" rid="B28">Gong et al., 2003</xref>; <xref ref-type="bibr" rid="B66">Shiah et al., 2003</xref>; <xref ref-type="bibr" rid="B48">Lai et al., 2014</xref>). Finally, PP was calculated <italic>via</italic> trapezoidal integration (i.e. integrated from surface to 1% PAR). Note that SST, Chl and PP measurements from the Ocean Data Bank (Ministry of Science and Technology, Taiwan<sup><xref ref-type="fn" rid="footnote1">1</xref></sup>) were used to compensate for deficiencies in <italic>in situ</italic> observations. Some of the hydrographic records (i.e., SST) and biogeochemical parameters (i.e., PP, Chl<sub>s</sub>) of May&#x2013;October from 2003 to 2006 have been present by <xref ref-type="bibr" rid="B60">Pan et al. (2012)</xref>.</p>
</sec>
<sec id="S2.SS2">
<title>Remote Sensing Observation</title>
<p>Between 2003 and 2016, the daily data of level-3 products were obtained using a passive ocean color MODerate resolution Imaging Spectroradiometer (Aqua sensor; MODIS-Aqua) with spatial resolution of 4 km for the region covering 17.5&#x2013;18.5&#x00B0;N and 115.5&#x2013;116.5&#x00B0;E. We also collected PP<sub>VGPM</sub>, SST, E<sub>0</sub>, K<sub>d</sub>, and Chl<sub>s</sub> values from the Environmental Research Division&#x2019;s Data Access Program (ERDDAP), National Oceanic and Atmospheric Administration, Department of Commerce, U. S.<sup><xref ref-type="fn" rid="footnote2">2</xref></sup>. Note that 8-day composite data were applied in situations where daily observations were missing. Note also that PP<sub>VGPM</sub> values obtained from ERDDAP were estimated using VGPM developed by <xref ref-type="bibr" rid="B4">Behrenfeld and Falkowski (1997)</xref>.</p>
<p>Plausible deviations between PP<sub>VGPM</sub> and PP<sub><italic>in situ</italic></sub> were examined by comparing data obtained from satellites vs. data obtained <italic>in situ</italic>, including i.e., SST<sub>sate</sub>, (<inline-formula><mml:math id="INEQ10"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula>)<sub>sate</sub>, E<sub>0&#x2013;sate</sub>, [E<sub>0</sub>/(E<sub>0</sub>+4.1)]<sub>sate</sub>, K<sub>d&#x2013;sate</sub>, Z<sub>eu&#x2013;sate</sub>, Chl<sub>s&#x2013;sate</sub>; SST<sub><italic>in situ</italic></sub>, (<inline-formula><mml:math id="INEQ11"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula>)<sub><italic>in situ</italic></sub>, E<sub>0&#x2013;</sub><sub><italic>in situ</italic></sub>, [E<sub>0</sub>/(E<sub>0</sub>+4.1)]<sub><italic>in situ</italic></sub>, K<sub>d&#x2013;</sub><sub><italic>in situ</italic></sub>, Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub>, Chl<sub>s&#x2013;</sub><sub><italic>in situ</italic></sub>. Note that this analysis was conducted using the VGPM algorithm proposed by <xref ref-type="bibr" rid="B4">Behrenfeld and Falkowski (1997)</xref> (with far fewer input variables), as Equation 2&#x2013;1:</p>
<disp-formula id="S2.Ex1"><label>(2-1)</label><mml:math id="M1" display="block"><mml:mtable><mml:mtr><mml:mtd columnalign="left"><mml:mrow><mml:mtext>PP</mml:mtext><mml:mo>=</mml:mo><mml:mpadded width="+3.3pt"><mml:mn>0.66125</mml:mn></mml:mpadded><mml:mo rspace="5.8pt">&#x00D7;</mml:mo><mml:mpadded width="+3.3pt"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mrow><mml:mtext>B</mml:mtext></mml:mrow></mml:msubsup></mml:mpadded><mml:mo rspace="5.8pt">&#x00D7;</mml:mo><mml:mrow><mml:mo stretchy="false">[</mml:mo><mml:msub><mml:mtext>E</mml:mtext><mml:mn>0</mml:mn></mml:msub><mml:mo>/</mml:mo><mml:mrow><mml:mo stretchy="false">(</mml:mo><mml:msub><mml:mtext>E</mml:mtext><mml:mn>0</mml:mn></mml:msub><mml:mo>+</mml:mo><mml:mn>4.1</mml:mn><mml:mo stretchy="false">)</mml:mo></mml:mrow><mml:mo rspace="5.8pt" stretchy="false">]</mml:mo></mml:mrow><mml:mo rspace="5.8pt">&#x00D7;</mml:mo><mml:mpadded width="+3.3pt"><mml:msub><mml:mtext>Z</mml:mtext><mml:mrow><mml:mtext>eu</mml:mtext></mml:mrow></mml:msub></mml:mpadded><mml:mo>&#x00D7;</mml:mo></mml:mrow></mml:mtd></mml:mtr><mml:mtr><mml:mtd columnalign="left"><mml:mspace width="2.5em"/><mml:mrow><mml:mpadded width="+3.3pt"><mml:msub><mml:mtext>Chl</mml:mtext><mml:mrow><mml:mtext>s</mml:mtext></mml:mrow></mml:msub></mml:mpadded><mml:mo rspace="5.8pt">&#x00D7;</mml:mo><mml:msub><mml:mtext>D</mml:mtext><mml:mrow><mml:mtext>irr</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:mtd></mml:mtr></mml:mtable></mml:math></disp-formula>
<p>where the D<sub>irr</sub> refers to the photoperiod. Z<sub>eu</sub> was estimated from the average K<sub>d</sub> of the water column (<xref ref-type="bibr" rid="B4">Behrenfeld and Falkowski, 1997</xref> and references therein).</p>
<disp-formula id="S2.Ex2"><label>(2-2)</label><mml:math id="M2" display="block"><mml:mtable><mml:mtr><mml:mtd columnalign="left"><mml:mrow><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mrow><mml:mtext>B</mml:mtext></mml:mrow></mml:msubsup><mml:mo>=</mml:mo><mml:mo>-</mml:mo><mml:mpadded width="+3.3pt"><mml:mn>3.27</mml:mn></mml:mpadded><mml:mo rspace="5.8pt">&#x00D7;</mml:mo><mml:msup><mml:mn>10</mml:mn><mml:mrow><mml:mo>-</mml:mo><mml:mn>8</mml:mn></mml:mrow></mml:msup><mml:msup><mml:mtext>T</mml:mtext><mml:mn>7</mml:mn></mml:msup><mml:mo>+</mml:mo><mml:mpadded width="+3.3pt"><mml:mn>3.4132</mml:mn></mml:mpadded><mml:mo rspace="5.8pt">&#x00D7;</mml:mo><mml:msup><mml:mn>10</mml:mn><mml:mrow><mml:mo>-</mml:mo><mml:mn>6</mml:mn></mml:mrow></mml:msup><mml:msup><mml:mtext>T</mml:mtext><mml:mn>6</mml:mn></mml:msup><mml:mo>-</mml:mo><mml:mpadded width="+3.3pt"><mml:mn>1.348</mml:mn></mml:mpadded><mml:mo>&#x00D7;</mml:mo></mml:mrow></mml:mtd></mml:mtr><mml:mtr><mml:mtd columnalign="left"><mml:mspace width="3em"/><mml:mrow><mml:mrow><mml:mrow><mml:mrow><mml:msup><mml:mn>10</mml:mn><mml:mrow><mml:mo>-</mml:mo><mml:mn>4</mml:mn></mml:mrow></mml:msup><mml:msup><mml:mtext>T</mml:mtext><mml:mn>5</mml:mn></mml:msup></mml:mrow><mml:mo>+</mml:mo><mml:mrow><mml:mrow><mml:mpadded width="+3.3pt"><mml:mn>2.462</mml:mn></mml:mpadded><mml:mo rspace="5.8pt">&#x00D7;</mml:mo><mml:msup><mml:mn>10</mml:mn><mml:mrow><mml:mo>-</mml:mo><mml:mn>3</mml:mn></mml:mrow></mml:msup></mml:mrow><mml:msup><mml:mtext>T</mml:mtext><mml:mn>4</mml:mn></mml:msup></mml:mrow></mml:mrow><mml:mo>-</mml:mo><mml:mrow><mml:mn>0.0205</mml:mn><mml:msup><mml:mtext>T</mml:mtext><mml:mn>3</mml:mn></mml:msup></mml:mrow></mml:mrow><mml:mo>+</mml:mo><mml:mrow><mml:mrow><mml:mn>0.0617</mml:mn><mml:msup><mml:mtext>T</mml:mtext><mml:mn>2</mml:mn></mml:msup></mml:mrow><mml:mo>+</mml:mo></mml:mrow></mml:mrow></mml:mtd></mml:mtr><mml:mtr><mml:mtd columnalign="left"><mml:mspace width="3em"/><mml:mrow><mml:mrow><mml:mn>0.2749</mml:mn><mml:mi mathvariant="normal">T</mml:mi></mml:mrow><mml:mo>+</mml:mo><mml:mn>1.2956</mml:mn></mml:mrow></mml:mtd></mml:mtr></mml:mtable></mml:math></disp-formula>
<p>To evaluate the impact of monsoonal force on time-series variations, all data sets were divided into two groups: SW-monsoon (SWM, Apr. to Oct., including inter-monsoons of Apr and Oct) and NE-monsoon (NEM, Nov. to Mar). Variations between satellite-based observations and <italic>in situ</italic> measurements were examined by averaging all relevant values and reporting them as mean &#x00B1; standard deviation (SD). Linear regression was used to assess relationships between any two of the variables. The <italic>t</italic>-tests were used to compare sets of observations with the significance level set at <italic>p</italic> = 0.10.</p>
</sec>
</sec>
<sec sec-type="results" id="S3">
<title>Results</title>
<sec id="S3.SS1">
<title>Time-Series Distributions: PP, SST, E<sub>0</sub>, K<sub>d</sub>, and Chl<sub>s</sub></title>
<p>As shown in <xref ref-type="fig" rid="F2">Figure 2A</xref>, PP<sub><italic>in situ</italic></sub> was &#x223C;2.0 times higher during the NEM (117&#x2013;528, average = 323 &#x00B1; 134 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup>) than during the SWM (76&#x2013;247, average = 159 &#x00B1; 58 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>) (<xref ref-type="table" rid="T2">Table 2</xref>). PP<sub>VGPM</sub> values between 2003 and 2016 were as follows: SWM was 250 &#x00B1; 36; 154&#x2013;771 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup> and NEM was 443 &#x00B1; 142; 190&#x2013;1,153 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup> (<xref ref-type="fig" rid="F2">Figure 2A</xref>). Note that PP<sub>VGPM</sub> during the NEM was &#x223C;1.8 times higher than during the SWM (<italic>p</italic> &#x003C; 0.01) (<xref ref-type="table" rid="T2">Table 2</xref>). The differences in PP<sub><italic>in situ</italic></sub> and PP<sub>VGPM</sub> between the NEM and SWM are in line with those reported by <xref ref-type="bibr" rid="B59">Ning et al. (2004)</xref>; <xref ref-type="bibr" rid="B14">Chen (2005)</xref>, and <xref ref-type="bibr" rid="B31">Hao et al. (2007)</xref>, all of which were obtained from the same area of the SCS using different methods (<xref ref-type="table" rid="T1">Table 1</xref>). PP<sub>VGPM</sub> exhibited monsoonal variations resembling the curve derived from PP<sub><italic>in situ</italic></sub>; however, the magnitudes were 37% higher during the NEM and 57% higher during the SWM. On average, PP<sub><italic>in situ</italic></sub> estimates were &#x223C;50% lower than PP<sub>VGPM</sub> estimates, regardless of monsoon.</p>
<fig id="F2" position="float">
<label>FIGURE 2</label>
<caption><p>Time-series distributions of satellite-based (circles) and <italic>in situ</italic> measurements obtained at the SEATS site (crosses): <bold>(A)</bold> Euphotic zone integrated primary production, PP; <bold>(B)</bold> sea surface temperature, SST; <bold>(C)</bold> optimal carbon fixation rate, <inline-formula><mml:math id="INEQ64"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mrow><mml:mtext>B</mml:mtext></mml:mrow></mml:msubsup></mml:math></inline-formula>; <bold>(D)</bold> sea surface daily light intensity, E<sub>0</sub>; <bold>(E)</bold> E<sub>0</sub>/(E<sub>0</sub>+4.1) ratio; <bold>(F)</bold> light diffuse attenuation coefficient, K<sub>d</sub>; <bold>(G)</bold> depth of euphotic zone, Z<sub>eu</sub>; <bold>(H)</bold> sea surface Chl, Chl<sub>s</sub>. Red and blue colors on the horizontal axis, respectively, indicate SWM (Apr&#x2013;Oct) and NEM (Nov&#x2013;Mar).</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-08-747763-g002.tif"/>
</fig>
<table-wrap position="float" id="T2">
<label>TABLE 2</label>
<caption><p>Summary of euphotic zone PP (PP) and its relevant variables in the current study conducted at the SEATS: SST, <inline-formula><mml:math id="INEQ15"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula>, E<sub>0</sub>, E<sub>0</sub>/(E<sub>0</sub>+4.1), K<sub>d</sub>, Z<sub>eu</sub>, Chl<sub>s</sub> and Z<sub>ML</sub>.</p></caption>
<table cellspacing="5" cellpadding="5" frame="hsides" rules="groups">
<thead>
<tr>
<td valign="top" align="left">Variables</td>
<td valign="top" align="center">NE-monsoon (n)</td>
<td valign="top" align="center">SW-monsoon (n)</td>
<td valign="top" align="center"><italic>p</italic> value</td>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">PP<sub><italic>in situ</italic></sub> (mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>)</td>
<td valign="top" align="center">323 &#x00B1; 134(10)</td>
<td valign="top" align="center">159 &#x00B1; 58(10)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">PP<sub>VGPM</sub> (mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>)</td>
<td valign="top" align="center">443 &#x00B1; 142(66)</td>
<td valign="top" align="center">250 &#x00B1; 36(96)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">SST<sub><italic>in situ</italic></sub> (&#x00B0;C)</td>
<td valign="top" align="center">25.0 &#x00B1; 1.4(12)</td>
<td valign="top" align="center">28.9 &#x00B1; 0.9(13)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">SST<sub>sate</sub> (&#x00B0;C)</td>
<td valign="top" align="center">25.1 &#x00B1; 1.2(70)</td>
<td valign="top" align="center">28.8 &#x00B1; 1.0(98)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">(<inline-formula><mml:math id="INEQ26"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula>)<sub><italic>in situ</italic></sub> (mg-C mg-Chl<sup>&#x2013;</sup><sup>1</sup> h<sup>&#x2013;</sup><sup>1</sup>)</td>
<td valign="top" align="center">5.4 &#x00B1; 0.5(12)</td>
<td valign="top" align="center">4.1 &#x00B1; 0.2(13)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">(<inline-formula><mml:math id="INEQ29"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula>)<sub>sate</sub> (mg-C mg-Chl<sup>&#x2013;</sup><sup>1</sup> h<sup>&#x2013;</sup><sup>1</sup>)</td>
<td valign="top" align="center">5.4 &#x00B1; 0.5(70)</td>
<td valign="top" align="center">4.2 &#x00B1; 0.2(98)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">E<sub>0&#x2013;</sub><sub><italic>in situ</italic></sub> (Eins m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>)</td>
<td valign="top" align="center">33 &#x00B1; 14(5)</td>
<td valign="top" align="center">50 &#x00B1; 11(11)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">E<sub>0&#x2013;sate</sub> (Eins m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>)</td>
<td valign="top" align="center">36 &#x00B1; 8(70)</td>
<td valign="top" align="center">47 &#x00B1; 6(98)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">(E<sub>0</sub>/(E<sub>0</sub>+4.1))<sub><italic>in situ</italic></sub></td>
<td valign="top" align="center">0.88 &#x00B1; 0.05(5)</td>
<td valign="top" align="center">0.92 &#x00B1; 0.02(11)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.10</td>
</tr>
<tr>
<td valign="top" align="left">(E<sub>0</sub>/(E<sub>0</sub>+4.1))<sub>sate</sub></td>
<td valign="top" align="center">0.89 &#x00B1; 0.02(70)</td>
<td valign="top" align="center">0.92 &#x00B1; 0.01(98)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">K<sub>d&#x2013;</sub><sub><italic>in situ</italic></sub> (&#x00D7;10<sup>&#x2013;</sup><sup>3</sup> m<sup>&#x2013;</sup><sup>1</sup>)</td>
<td valign="top" align="center">63 &#x00B1; 11(12)</td>
<td valign="top" align="center">54 &#x00B1; 6(13)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.05</td>
</tr>
<tr>
<td valign="top" align="left">K<sub>d&#x2013;sate</sub> (&#x00D7;10<sup>&#x2013;</sup><sup>3</sup> m<sup>&#x2013;</sup><sup>1</sup>)</td>
<td valign="top" align="center">40 &#x00B1; 10(69)</td>
<td valign="top" align="center">30 &#x00B1; 3(94)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> (m)</td>
<td valign="top" align="center">75 &#x00B1; 14(12)</td>
<td valign="top" align="center">87 &#x00B1; 10(13)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.05</td>
</tr>
<tr>
<td valign="top" align="left">Z<sub>eu&#x2013;sate</sub> (m)</td>
<td valign="top" align="center">123 &#x00B1; 30(69)</td>
<td valign="top" align="center">156 &#x00B1; 17(94)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">Chl<sub>s&#x2013;</sub><sub><italic>in situ</italic></sub> (mg m<sup>&#x2013;</sup><sup>3</sup>)</td>
<td valign="top" align="center">0.24 &#x00B1; 0.16(12)</td>
<td valign="top" align="center">0.08 &#x00B1; 0.02(13)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">Chl<sub>s&#x2013;sate</sub> (mg m<sup>&#x2013;</sup><sup>3</sup>)</td>
<td valign="top" align="center">0.20 &#x00B1; 0.08(68)</td>
<td valign="top" align="center">0.10 &#x00B1; 0.02(96)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
<tr>
<td valign="top" align="left">Z<sub>ML</sub> (m)</td>
<td valign="top" align="center">66 &#x00B1; 20(31)</td>
<td valign="top" align="center">35 &#x00B1; 11(29)</td>
<td valign="top" align="center"><italic>p</italic> &#x003C; 0.01</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn><p><italic>Values obtained during the NEM and SWM are presented as means and SD, where n indicates the number of observations and p values indicate the level of significance in the t test. In situ, in situ measured; VGPM, VGPM estimate; sate, satellite-base.</italic></p></fn>
</table-wrap-foot>
</table-wrap>
<p>The lowest SST<sub><italic>in situ</italic></sub> (25.0 &#x00B1; 1.4&#x00B0;C) and SST<sub>sate</sub> (25.1 &#x00B1; 1.2&#x00B0;C) values were obtained under the prevailing NEM during the cold season. The highest SST<sub><italic>in situ</italic></sub> (28.9 &#x00B1; 0.9&#x00B0;C) and SST<sub>sate</sub> (28.8 &#x00B1; 1.0&#x00B0;C) values were obtained under the prevailing SWM during the warm season (<xref ref-type="table" rid="T2">Table 2</xref>). The low SST values recorded during the NEM can be attributed to the combined dynamics of NEM-driven vertical mixing of seawater and reduced solar radiation (<xref ref-type="bibr" rid="B79">Tseng et al., 2005</xref>; <xref ref-type="bibr" rid="B85">Zhou et al., 2020</xref>). By contrast, the <inline-formula><mml:math id="INEQ58"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mrow><mml:mtext>B</mml:mtext></mml:mrow></mml:msubsup></mml:math></inline-formula> showed a synchronous but opposite distribution (<xref ref-type="fig" rid="F2">Figures 2B,C</xref>). The (<inline-formula><mml:math id="INEQ59"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mrow><mml:mtext>B</mml:mtext></mml:mrow></mml:msubsup></mml:math></inline-formula>)<sub><italic>in situ</italic></sub> and (<inline-formula><mml:math id="INEQ60"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mrow><mml:mtext>B</mml:mtext></mml:mrow></mml:msubsup></mml:math></inline-formula>)<sub>sate</sub> values computed from SST for the NEM, respectively, ranged from 4.4 to 6.1 and 4.0 to 6.6 mg-C mg-Chl<sup>&#x2013;</sup><sup>1</sup> h<sup>&#x2013;</sup><sup>1</sup>, and the average value was the same (5.4 &#x00B1; 0.5 mg-C mg-Chl<sup>&#x2013;</sup><sup>1</sup> h<sup>&#x2013;</sup><sup>1</sup>). The (<inline-formula><mml:math id="INEQ61"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mrow><mml:mtext>B</mml:mtext></mml:mrow></mml:msubsup></mml:math></inline-formula>)<sub><italic>in situ</italic></sub> and (<inline-formula><mml:math id="INEQ62"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mrow><mml:mtext>B</mml:mtext></mml:mrow></mml:msubsup></mml:math></inline-formula>)<sub>sate</sub> values for the SWM, respectively, ranged from 4.0 to 4.4 and 4.0 to 6.0 mg-C mg-Chl<sup>&#x2013;</sup><sup>1</sup> h<sup>&#x2013;</sup><sup>1</sup>, with mean <inline-formula><mml:math id="INEQ63"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mrow><mml:mtext>B</mml:mtext></mml:mrow></mml:msubsup></mml:math></inline-formula> values of 4.1 &#x00B1; 0.2 and 4.2 &#x00B1; 0.2 mg-C mg-Chl<sup>&#x2013;</sup><sup>1</sup> h<sup>&#x2013;</sup><sup>1</sup> (<xref ref-type="table" rid="T2">Table 2</xref>).</p>
<p>E<sub>0&#x2013;</sub><sub><italic>in situ</italic></sub> and E<sub>0&#x2013;sate</sub> values in the SWM (50 &#x00B1; 11 and 47 &#x00B1; 6 Eins m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>) were higher than those in the NEM (33 &#x00B1; 14 and 36 &#x00B1; 8 Eins m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>) (see <xref ref-type="fig" rid="F2">Figures 2D,E</xref> and <xref ref-type="table" rid="T2">Table 2</xref>). From the perspective of variables in the VGPM algorithm, E<sub>0</sub>/(E<sub>0</sub>+4.1) ratios varied little between the <italic>in situ</italic> measurement and satellite-based observation. The variable of [E<sub>0</sub>/(E<sub>0</sub>+4.1)] thereby seemed not an important parameter to influence the PP level. Z<sub>eu</sub> was estimated using the expression of &#x2013;ln(0.01)/K<sub>d</sub> (<xref ref-type="bibr" rid="B46">Kirk, 1994</xref>; <xref ref-type="bibr" rid="B4">Behrenfeld and Falkowski, 1997</xref>), which led to a reversal of monsoonal K<sub>d</sub> and Z<sub>eu</sub> distributions (see <xref ref-type="fig" rid="F2">Figures 2F,G</xref>). The mean Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> was deeper during the SWM than during the NEM (87 &#x00B1; 10 and 75 &#x00B1; 14 m, respectively). The Z<sub>eu&#x2013;sate</sub> as well as the Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> presented similar trends between SWM and NEM (156 &#x00B1; 17 and 120 &#x00B1; 30 m, respectively) (<xref ref-type="table" rid="T2">Table 2</xref>). Z<sub>eu</sub> values were shallower in the NEM than in the SWM, due perhaps to reduced light penetration resulting from more abundant biomass in the water column (<xref ref-type="bibr" rid="B14">Chen, 2005</xref>; <xref ref-type="bibr" rid="B79">Tseng et al., 2005</xref>; <xref ref-type="bibr" rid="B18">Chen et al., 2008</xref>).</p>
<p><xref ref-type="fig" rid="F2">Figure 2H</xref> presents long-term temporal variations in Chl<sub>s</sub>. <italic>In situ</italic> measurements and satellite-based observations revealed a distinct NEM maximum (0.24 &#x00B1; 0.16 and 0.20 &#x00B1; 0.08 mg m<sup>&#x2013;</sup><sup>3</sup>, respectively) and a SWM minimum (0.08 &#x00B1; 0.02 and 0.10 &#x00B1; 0.02 mg m<sup>&#x2013;</sup><sup>3</sup>, respectively). Chl<sub>s&#x2013;</sub><sub><italic>in situ</italic></sub> and Chl<sub>s&#x2013;sate</sub> concentrations were, respectively, 0.07 to 0.58 and 0.09 to 0.71 mg m<sup>&#x2013;</sup><sup>3</sup> during the NEM, and were, respectively, 0.04 to 0.11 and 0.05 to 0.43 mg m<sup>&#x2013;</sup><sup>3</sup> during the SWM. For the same location, our observations were close to the data reported by <xref ref-type="bibr" rid="B14">Chen (2005)</xref> and <xref ref-type="bibr" rid="B79">Tseng et al. (2005)</xref>. The SWM minimum Chl<sub>s</sub> values in the current study were nearly the same as those reported in the oligotrophic ocean time-series studies at HOT (Hawaii Ocean Time-Series) and BATS (Bermuda Atlantic Time-Series Study) during the summer (0.05 mg m<sup>&#x2013;</sup><sup>3</sup>); however, NEM values at SEATS exceeded the winter maximum at HOT and BATS (0.1 and 0.3 mg m<sup>&#x2013;</sup><sup>3</sup>, respectively) (<xref ref-type="bibr" rid="B45">Karl et al., 2003</xref>). The high NEM maximum at SEATS may perhaps be explained by an increase in phytoplankton biomass (particularly larger sizes of &#x003E;3 &#x03BC;m), which was stimulated by the deepening Z<sub>ML</sub> (<xref ref-type="table" rid="T2">Table 2</xref>). The deeper nutrient-rich water was then efficiently transported to the upper surface water under the influence of NEM given that the nutrient-cline depth was shallower in SCS than in other oceans (<xref ref-type="bibr" rid="B14">Chen, 2005</xref>; <xref ref-type="bibr" rid="B79">Tseng et al., 2005</xref>).</p>
</sec>
<sec id="S3.SS2">
<title>Monthly Variations in PP, SST, E<sub>0</sub>, K<sub>d</sub> and Chl<sub>s</sub></title>
<p>The PP<sub>VGPM</sub> values in this study are in good agreement with PP<sub><italic>in situ</italic></sub> in terms of amplitude as well as phase. Overall, we observed a maximum PP<sub><italic>in situ</italic></sub> of 394 &#x00B1; 190 in January (NEM) and a minimum of 143 &#x00B1; 70 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup> in August (SWM) (<xref ref-type="fig" rid="F3">Figure 3A</xref>). The magnitude of PP<sub>VGPM</sub> was higher than the values obtained using <italic>in situ</italic> measurements; however, the trend over a span of 12 months was similar, with a pronounced NEM peak 619 &#x00B1; 113 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup> in January and the lowest SWM value of 230 &#x00B1; 28 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup> in September. These results are similar to those reported by other researchers for same area using different methods, such as the particulate organic carbon flux re-calculation (NEM: 207, SWM: 149 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>) (<xref ref-type="bibr" rid="B11">Chen et al., 1998</xref>); on-deck C<sup>14</sup> incubation (NEM: 300&#x2013;509, SWM: 110&#x2013;228 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>) (<xref ref-type="bibr" rid="B59">Ning et al., 2004</xref>; <xref ref-type="bibr" rid="B79">Tseng et al., 2005</xref>); on-deck C<sup>13</sup> incubation (NEM: 190&#x2013;550, SWM: 190&#x2013;280 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>) (<xref ref-type="bibr" rid="B14">Chen, 2005</xref>; <xref ref-type="bibr" rid="B17">Chen et al., 2007</xref>), and Chl empirical function (NEM: 148&#x2013;684, SWM: 86&#x2013;275 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>) (<xref ref-type="bibr" rid="B8">Chen et al., 2006</xref>; <xref ref-type="table" rid="T1">Table 1</xref>).</p>
<fig id="F3" position="float">
<label>FIGURE 3</label>
<caption><p>Monthly averages (&#x00B1;1 SD) of <bold>(A)</bold> PP, <bold>(B)</bold> SST, <bold>(C)</bold> <inline-formula><mml:math id="INEQ65"><mml:mrow><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mrow><mml:mtext>B</mml:mtext></mml:mrow></mml:msubsup><mml:mo>,</mml:mo></mml:mrow></mml:math></inline-formula> <bold>(D)</bold> E<sub>0</sub>, <bold>(E)</bold> E<sub>0</sub>/(E<sub>0</sub>+4.1), <bold>(F)</bold> K<sub>d</sub>, <bold>(G)</bold> Z<sub>eu</sub>, <bold>(H)</bold> Chl<sub>s</sub> at the SEATS site. Circles and triangles, respectively, represent <italic>in situ</italic> measurements and satellite-based observations. Red and blue colors on the horizontal axis, respectively, indicate SWM (Apr&#x2013;Oct) and NEM (Nov&#x2013;Mar).</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-08-747763-g003.tif"/>
</fig>
<p>We observed opposing trends between monthly SST and the corresponding <inline-formula><mml:math id="INEQ66"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula>. The highest (<inline-formula><mml:math id="INEQ67"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula>)<sub><italic>in situ</italic></sub> values (5.9 mg-C mg-Chl<sup>&#x2013;1</sup> h<sup>&#x2013;1</sup>) were observed in January and the lowest (4.0 mg-C mg-Chl<sup>&#x2013;1</sup> h<sup>&#x2013;1</sup>) were observed in August. The (<inline-formula><mml:math id="INEQ68"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula>)<sub>sate</sub> values converted from SST<sub>sate</sub> (5.8 and 4.0 mg-C mg-Chl<sup>&#x2013;1</sup> h<sup>&#x2013;1</sup> in January and June, respectively) were nearly identical to those for the same time slots obtained <italic>via in situ</italic> measurements (<xref ref-type="fig" rid="F3">Figures 3B,C</xref>). The highest mean monthly E<sub>0&#x2013;</sub><sub><italic>in situ</italic></sub> in April (63 &#x00B1; 0 Eins m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>) was higher than the highest mean monthly E<sub>0&#x2013;sate</sub> (53 &#x00B1; 3 Eins m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>). Overall, E<sub>0&#x2013;</sub><sub><italic>in situ</italic></sub> and E<sub>0&#x2013;sate</sub> levels in December were very similar (&#x223C;26 and &#x223C;27 Eins m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>, respectively). Based on E<sub>0&#x2013;</sub><sub><italic>in situ</italic></sub>, the highest E<sub>0</sub>/(E<sub>0</sub>+4.1)<sub><italic>in situ</italic></sub> ratio (0.94) occurred in April and the lowest E<sub>0</sub>/(E<sub>0</sub>+4.1)<sub><italic>in situ</italic></sub> ratio (0.85) occurred in December. E<sub>0</sub>/(E<sub>0</sub>+4.1)<sub>sate</sub> values of 0.93 in April and 0.86 in December were similar with the E<sub>0</sub>/(E<sub>0</sub>+4.1)<sub><italic>in situ</italic></sub> values (<xref ref-type="fig" rid="F3">Figures 3D,E</xref>). As for parameterization, it appears that <inline-formula><mml:math id="INEQ69"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> and E<sub>0</sub>/(E<sub>0</sub>+4.1) values had little influence on VGPM results (PP), regardless of whether the values were obtained from <italic>in situ</italic> measurements or satellite-based observation.</p>
<p>We did not observe large monsoonal variations in K<sub>d&#x2013;</sub><sub><italic>in situ</italic></sub> and K<sub>d&#x2013;sate</sub>; however, the highest K<sub>d&#x2013;</sub><sub><italic>in situ</italic></sub> (0.072 &#x00B1; 0.009 m<sup>&#x2013;</sup><sup>1</sup>) was obtained in December and the highest K<sub>d&#x2013;sate</sub> (0.052 &#x00B1; 0.009 m<sup>&#x2013;</sup><sup>1</sup>) was obtained in January. These high K<sub>d</sub> values resulted in a shallower Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> (64 &#x00B1; 8 m) and Z<sub>eu&#x2013;sate</sub> (92 &#x00B1; 19 m), compared to the values converted from the K<sub>d&#x2013;</sub><sub><italic>in situ</italic></sub> (0.053 &#x00B1; 0.007 m<sup>&#x2013;</sup><sup>1</sup>) for August (Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> = 89 &#x00B1; 13 m) and the K<sub>d&#x2013;sate</sub> (0.029 &#x00B1; 0.003 m<sup>&#x2013;</sup><sup>1</sup>) for September (Z<sub>eu&#x2013;sate</sub> = 162 &#x00B1; 22 m) (<xref ref-type="fig" rid="F3">Figures 3F,G</xref>). The shallow Z<sub>eu</sub> observed in December and January can be attributed mainly to an increase in phytoplankton biomass that reduces the light penetration in that region during the NEM (<xref ref-type="bibr" rid="B14">Chen, 2005</xref>; <xref ref-type="bibr" rid="B79">Tseng et al., 2005</xref>; <xref ref-type="bibr" rid="B18">Chen et al., 2008</xref>).</p>
<p><xref ref-type="fig" rid="F3">Figure 3H</xref> presents temporal variations in Chl<sub>s&#x2013;</sub><sub><italic>in situ</italic></sub> and Chl<sub>s&#x2013;sate</sub>. Conspicuously high Chl<sub>s&#x2013;</sub><sub><italic>in situ</italic></sub> values were observed in December (0.40 &#x00B1; 0.15 mg m<sup>&#x2013;</sup><sup>3</sup>) and high Chl<sub>s&#x2013;sate</sub> values were observed in January (0.29 &#x00B1; 0.09 mg m<sup>&#x2013;</sup><sup>3</sup>). The high Chl<sub>s</sub> values observed throughout the NEM were triggered by the monsoonal force, which stimulated phytoplankton photosynthesis, thereby increasing the phytoplankton abundance or enriching the area with Chl<sub>s</sub> from subsurface waters (<xref ref-type="bibr" rid="B54">Liu et al., 2002</xref>; <xref ref-type="bibr" rid="B79">Tseng et al., 2005</xref>). The drop in Chl<sub>s&#x2013;</sub><sub><italic>in situ</italic></sub> and Chl<sub>s&#x2013;sate</sub> values to nearly &#x003C; 0.10 mg m<sup>&#x2013;</sup><sup>3</sup> during the SWM has previously been reported in studies focusing on the same region of the SCS (<xref ref-type="bibr" rid="B54">Liu et al., 2002</xref>; <xref ref-type="bibr" rid="B14">Chen, 2005</xref>; <xref ref-type="bibr" rid="B79">Tseng et al., 2005</xref>; <xref ref-type="bibr" rid="B70">Shih et al., 2020a</xref>). Similar findings were also recorded in oligotrophic time-series studies, such as HOT and BATS (<xref ref-type="bibr" rid="B45">Karl et al., 2003</xref>).</p>
</sec>
<sec id="S3.SS3">
<title>Relationships Among of PP, SST, E<sub>0</sub>, K<sub>d</sub> and Chl<sub>s</sub> in <italic>in situ</italic> Measurements and Satellite-Based Observations</title>
<p>PP<sub><italic>in situ</italic></sub> was generally lower than PP<sub>VGPM</sub>; however, we observed a significantly positive correlation between these two parameters; i.e., slope = 0.70, <italic>r</italic><sup>2</sup> = 0.42, <italic>p</italic> &#x003C; 0.01 (<xref ref-type="fig" rid="F4">Figure 4A</xref>). This clearly indicates the feasibility of the VGPM for predictions; however, tuning would be required for the study area. As shown in <xref ref-type="fig" rid="F4">Figure 4</xref>, we identified significantly positive correlations between PP<sub><italic>in situ</italic></sub> and PP<sub>VGPM</sub> as well as their respective variables SST, E<sub>0</sub>, K<sub>d</sub> and Chl<sub>s</sub>. As indicated by high <italic>r</italic><sup>2</sup> and low <italic>p</italic> values with slopes close to 1, the most significant correlations were found in SST and E<sub>0</sub> (<xref ref-type="fig" rid="F4">Figures 4C&#x2013;F</xref>) : SST (slope = 0.92, <italic>r</italic><sup>2</sup> = 0.92, <italic>p</italic> &#x003C; 0.01) and E<sub>0</sub> (slope = 0.62, <italic>r</italic><sup>2</sup> = 0.58, <italic>p</italic> &#x003C; 0.01). Taken together, these results indicate that SST<sub>sate</sub> and E<sub>0&#x2013;sate</sub> were the variables most strongly correlated with SST<sub><italic>in situ</italic></sub> and E<sub>0&#x2013;</sub><sub><italic>in situ</italic></sub>, providing the most accurate estimates of PP when using the VGPM. As indicated by the 1:1 lines in <xref ref-type="fig" rid="F4">Figures 4B,G,H</xref>, the parameters with the greatest variability in terms of slope were Chl<sub>s</sub> (slope = 0.41, <italic>r</italic><sup>2</sup> = 0.74, <italic>p</italic> &#x003C; 0.01) and K<sub>d</sub> (slope = 0.59, <italic>r</italic><sup>2</sup> = 0.50, <italic>p</italic> &#x003C; 0.01). This suggests Chl<sub>s&#x2013;sate</sub> and K<sub>d&#x2013;sate</sub> could potentially bias PP estimates obtained using the VGPM.</p>
<fig id="F4" position="float">
<label>FIGURE 4</label>
<caption><p>Linear relationships among satellite-based observations and <italic>in situ</italic> measurements. Panels <bold>(A&#x2013;H)</bold> are PP, Chl<sub>s</sub>, SST, <inline-formula><mml:math id="INEQ70"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mrow><mml:mtext>B</mml:mtext></mml:mrow></mml:msubsup></mml:math></inline-formula> (estimated from SST), E<sub>0</sub>, E<sub>0</sub>/(E<sub>0</sub>+4.1) ratio, K<sub>d</sub>, and Z<sub>eu</sub> (estimated from K<sub>d</sub>), respectively. Diagonals in panels <bold>(A&#x2013;H)</bold> are 1:1 lines.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-08-747763-g004.tif"/>
</fig>
<p>The fact that the regression line between PP<sub>VGPM</sub> and PP<sub><italic>in situ</italic></sub> lies above the 1:1 line indicates that PP<sub>VGPM</sub> estimates exceeded PP<sub><italic>in situ</italic></sub>. When using the VGPM to estimate PP, the two main variables are <inline-formula><mml:math id="INEQ71"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> and E<sub>0</sub>/(E<sub>0</sub>+4.1), derived from <italic>in situ</italic> measurements and satellite-based observations of SST (SST<sub><italic>in situ</italic></sub> and SST<sub>sate</sub>) and E<sub>0</sub> (E<sub>0&#x2013;</sub><sub><italic>in situ</italic></sub> and E<sub>0&#x2013;sate</sub>). Between SST<sub><italic>in situ</italic></sub> and SST<sub>sate</sub> as well as between E<sub>0&#x2013;</sub><sub><italic>in situ</italic></sub> and E<sub>0&#x2013;sate</sub>, we observed nearly linear relationships (i.e., close to the 1:1 diagonal). This suggests that <inline-formula><mml:math id="INEQ72"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> and E<sub>0</sub>/(E<sub>0</sub>+4.1) were not scaling variables governing the magnitude of PP in this study. Influences derived from these two variables [<inline-formula><mml:math id="INEQ73"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula>, E<sub>0</sub>/(E<sub>0</sub>+4.1)] of the both methods on calculated results were less than 1%. By contrast, the correlations between Chl<sub>s&#x2013;</sub><sub><italic>in situ</italic></sub> and Chl<sub>s&#x2013;sate</sub>, and Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> and Z<sub>eu&#x2013;sate</sub> (converted from K<sub>d&#x2013;</sub><sub><italic>in situ</italic></sub> and K<sub>d&#x2013;sate</sub>, respectively) had a pronounced impact on VGPM PP estimates. If we considered only the difference between Chl<sub>s&#x2013;</sub><sub><italic>in situ</italic></sub> and Chl<sub>s&#x2013;sate</sub>, then PP values estimated using VGPM would be slightly higher (&#x223C; 5%, depended on the given case) than those obtained using <italic>in situ</italic> measurements. If we considered only the difference between Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> and Z<sub>eu&#x2013;sate</sub>, then PP values estimated using VGPM would be apparently 1.72-fold higher than those obtained using <italic>in situ</italic> measurements.</p>
</sec>
</sec>
<sec sec-type="discussion" id="S4">
<title>Discussion</title>
<sec id="S4.SS1">
<title>Uncertainty in Estimating PP Due to Differences Between Chl<sub>s&#x2013;<italic>in situ</italic></sub> and Chl<sub>s&#x2013;sate</sub> as Well as Z<sub>eu&#x2013;<italic>in situ</italic></sub> and Z<sub>eu&#x2013;sate</sub>: Implications</title>
<p>Our analysis of revealed a number of potential uncertainties pertaining to PP estimation using the VGPM algorithm. Most of the discrepancies were due primarily to differences between Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> and Z<sub>eu&#x2013;sate</sub> as well as between Chl<sub>s&#x2013;</sub><sub><italic>in situ</italic></sub> and Chl<sub>s&#x2013;sate</sub>. Overall, the product of Z<sub>eu</sub> and Chl<sub>s</sub> (i.e., the phytoplankton inventory in the euphotic zone), suggests that the base assumption of vertically distributed standing stock biomass in low latitude waters (SCS) may perhaps be erroneous. If so, then it will be necessary to reformulate methods for the prediction of biomass standing stocks when implementing the VGPM algorithm (<xref ref-type="bibr" rid="B59">Ning et al., 2004</xref>; <xref ref-type="bibr" rid="B33">Hill and Zimmerman, 2010</xref>).</p>
<p>The fundamental structure of the VGPM is based on a relationship between integrated phytoplankton biomass in the euphotic zone and Chl<sub>s</sub> (<xref ref-type="bibr" rid="B4">Behrenfeld and Falkowski, 1997</xref>; <xref ref-type="bibr" rid="B33">Hill and Zimmerman, 2010</xref>). Thus, obtaining accurate estimates of PP by comparing PP<sub><italic>in situ</italic></sub> and PP<sub>VGPM</sub> results depends on reliable estimates of the integrated phytoplankton biomass in the euphotic zone. However, passive satellites recording the color of the ocean surface are unable to elucidate the situation at arbitrary depths below the surface (<xref ref-type="bibr" rid="B33">Hill and Zimmerman, 2010</xref>; <xref ref-type="bibr" rid="B71">Shih et al., 2020b</xref>). Contrary to the assumption that Chl decreases exponentially with depth, most observations in the SCS revealed that the subsurface Chl maximum (SCM) was often found at great depths (<xref ref-type="bibr" rid="B54">Liu et al., 2002</xref>; <xref ref-type="bibr" rid="B14">Chen, 2005</xref>; <xref ref-type="bibr" rid="B70">Shih et al., 2020a</xref>, <xref ref-type="bibr" rid="B71">b</xref>). This makes it very difficult or even impossible to estimate the integrated biomass in the euphotic zone simply as a product of Z<sub>eu</sub> and Chl<sub>s</sub>. Enhancing the reliability of the VGPM requires that we increase the number of PP<sub><italic>in situ</italic></sub> observations and the corresponding variables in order to improve the correlation between our assumptions pertaining to phytoplankton integrated biomass and actual measurements obtained in the field. This is particularly important in phytoplankton populations, dynamics and assemblages in specific locations under specific conditions (<xref ref-type="bibr" rid="B33">Hill and Zimmerman, 2010</xref>; <xref ref-type="bibr" rid="B69">Shih et al., 2015</xref>, <xref ref-type="bibr" rid="B71">2020b</xref>). Only by increasing the number of observations and enhancing our analysis of water composition will it be possible to reduce the uncertainty associated with Z<sub>eu</sub> and Chl<sub>s</sub> in estimating PP using the VGPM.</p>
<p>The mean PP in the euphotic zone (PP/Z<sub>eu</sub>, mg-C m<sup>&#x2013;</sup><sup>3</sup> d<sup>&#x2013;</sup><sup>1</sup>) presented a positive linear relationship between (PP<sub>VGPM</sub>/Z<sub>eu&#x2013;sate</sub>) and (PP<sub><italic>in situ</italic></sub>/Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub>); i.e., slope: 0.51, <italic>r</italic><sup>2</sup> = 0.39, <italic>p</italic> &#x003C; 0.01 (<xref ref-type="fig" rid="F5">Figure 5A</xref>). Higher values were observed during the NEM (4.7 &#x00B1; 2.4 and 3.8 &#x00B1; 2.2 mg-C m<sup>&#x2013;</sup><sup>3</sup> d<sup>&#x2013;</sup><sup>1</sup> of PP<sub><italic>in situ</italic></sub>/Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> and PP<sub>VGPM</sub>/Z<sub>eu&#x2013;sate</sub>, respectively) and lower values were observed during the SWM (1.9 &#x00B1; 0.8 and 1.7 &#x00B1; 0.4 mg-C m<sup>&#x2013;</sup><sup>3</sup> d<sup>&#x2013;</sup><sup>1</sup> of PP<sub><italic>in situ</italic></sub>/Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> and PP<sub>VGPM</sub>/Z<sub>eu&#x2013;sate</sub>, respectively). The slope of 0.51 for PP<sub>VGPM</sub>/Z<sub>eu&#x2013;sate</sub> and PP<sub><italic>in situ</italic></sub>/Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> was lower than that of PP<sub>VGPM</sub> and PP<sub><italic>in situ</italic></sub> (slope = 0.70), such that most data fell on the right side of the 1:1 line. Ratios of PP<sub>VGPM</sub>/Z<sub>eu&#x2013;sate</sub> were nearly 20% lower than those of PP<sub><italic>in situ</italic></sub>/Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub>, indicating that Z<sub>eu&#x2013;sate</sub> was deeper than Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub>, particularly during the SWM. This also indicates that the VGPM parameter Z<sub>eu&#x2013;sate</sub> indeed substantially affected PP estimates in this study. It has been proposed that satellite-based K<sub>d</sub> values have to be calibrated against the zenith solar angle during the data processing in accordance with the methods outlined by <xref ref-type="bibr" rid="B50">Lee et al. (2005)</xref> and <xref ref-type="bibr" rid="B52">Li et al. (2015)</xref>. However, users may not to confirm the processing from the downloaded or retrieved satellite-based products of PP and its relevant variables.</p>
<fig id="F5" position="float">
<label>FIGURE 5</label>
<caption><p><bold>(A)</bold> Relationship between satellite-based observations of (PP<sub>VGPM</sub> / Z<sub>eu&#x2013;sate</sub>) and <italic>in situ</italic> measurements of (PP<sub><italic>in situ</italic></sub> / Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub>) ratios; <bold>(B)</bold> relationship between PP<sub><italic>in situ</italic></sub> and SST<sub><italic>in situ</italic></sub>. Red and blue symbols, respectively, indicate observations made during the SW-monsoon and NE-monsoon. The diagonal in panel <bold>(A)</bold> is 1:1 line.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-08-747763-g005.tif"/>
</fig>
<p>The significant linear correlation between (<inline-formula><mml:math id="INEQ74"><mml:mrow><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:mrow></mml:math></inline-formula>)<sub>sate</sub> and (<inline-formula><mml:math id="INEQ75"><mml:mrow><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:mrow></mml:math></inline-formula>)<sub><italic>in situ</italic></sub> (slope = 0.97, <italic>r</italic><sup>2</sup> = 0.94, <italic>p</italic> &#x003C; 0.01; <xref ref-type="fig" rid="F4">Figure 4D</xref>) is simply a reflection of the relationship between the two SST records, resulting from the fact that <inline-formula><mml:math id="INEQ76"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> is computed as a function of SST (i.e., a seventh-order polynomial, Equation 2&#x2013;2) (<xref ref-type="bibr" rid="B4">Behrenfeld and Falkowski, 1997</xref>). Thus, any potential deviations between SST and <inline-formula><mml:math id="INEQ77"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> had only a negligible influence on PP estimates obtained using VGPM. Many studies have nevertheless concluded that the accuracy of VGPM-based estimates of PP are poor, when using the 7th order polynomial of SST (Equation 2&#x2013;2) to calculate the input of <inline-formula><mml:math id="INEQ78"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> (<xref ref-type="bibr" rid="B57">Mizobata and Saitoh, 2004</xref>; <xref ref-type="bibr" rid="B43">Kameda and Ishizaka, 2005</xref>; <xref ref-type="bibr" rid="B81">Yamada et al., 2005</xref>; <xref ref-type="bibr" rid="B73">Siswanto et al., 2006</xref>; <xref ref-type="bibr" rid="B42">Ishizaka et al., 2007</xref>; <xref ref-type="bibr" rid="B76">Tang et al., 2008</xref>; <xref ref-type="bibr" rid="B75">Tang and Chen, 2016</xref>).</p>
<p>Researchers have reported that much of the uncertainty in estimating PP<sub>VGPM</sub> is related to the computation of <inline-formula><mml:math id="INEQ79"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> under the effects of phytoplankton physiology, growth conditions, abundance, size, and productivity (<xref ref-type="bibr" rid="B27">Gong and Liu, 2003</xref>; <xref ref-type="bibr" rid="B43">Kameda and Ishizaka, 2005</xref>; <xref ref-type="bibr" rid="B81">Yamada et al., 2005</xref>). It has been suggested that <inline-formula><mml:math id="INEQ80"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> is influenced by SST as well as E<sub>0</sub> and various biological parameters, such as Chl concentration. <inline-formula><mml:math id="INEQ81"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> represents an optimal daily carbon fixation rate in the water column previously described as a 7th order polynomial of SST, however, when SST exceeds 28.5&#x00B0;C, <inline-formula><mml:math id="INEQ83"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> remains fixed at a constant 4 mg-C mg-Chl<sup>&#x2013;1</sup> h<sup>&#x2013;1</sup> (<xref ref-type="bibr" rid="B4">Behrenfeld and Falkowski, 1997</xref>; <xref ref-type="bibr" rid="B57">Mizobata and Saitoh, 2004</xref>), indicating that <inline-formula><mml:math id="INEQ84"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> tends to be underestimated when SST exceeds 28.5&#x00B0;C. In low-latitude regions of the SCS, the constant <inline-formula><mml:math id="INEQ86"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> mentioned above usually occurred in late spring, summer, and early fall, during which SST<sub><italic>in situ</italic></sub> and SST<sub>sate</sub> both exceeded 28.5&#x00B0;C. This increased the margin between actual PP values and the estimates obtained by inputting <inline-formula><mml:math id="INEQ88"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> derived using SST<sub><italic>in situ</italic></sub> or SST<sub>sate</sub>. The reliability of <inline-formula><mml:math id="INEQ89"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula> estimates obtained using the VGPM seventh-order polynomial SST algorithm is not universally applicable (temporally or spatially). This has prompted oceanographers to tune existing methods or devise new methods for the precise estimation of <inline-formula><mml:math id="INEQ90"><mml:msubsup><mml:mtext>P</mml:mtext><mml:mrow><mml:mtext>opt</mml:mtext></mml:mrow><mml:mi>B</mml:mi></mml:msubsup></mml:math></inline-formula>, especially for ocean water at low latitudes.</p>
<p>Our results reveal that the satellite-derived primary production (e.g., PP<sub>VGPM</sub>) may significantly affect global carbon export flux to deep waters, but what is the overall significance and impact of these PP values on POC fluxes? For example, <xref ref-type="bibr" rid="B23">Dunne et al. (2005)</xref> used the empirical model expression (Equation 4&#x2013;1) to estimate carbon flux (or sequestration) in oceans. PP (mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>), Z<sub>eu</sub> (m) and SST (&#x00B0;C) are three major factors affecting the estimated values of POC flux which is almost linearly proportional to PP.</p>
<disp-formula id="S4.Ex7"><label>(4-1)</label><mml:math id="M3" display="block"><mml:mtable><mml:mtr><mml:mtd columnalign="left"><mml:mrow><mml:mpadded width="+5pt"><mml:mi>POC</mml:mi></mml:mpadded><mml:mi>flux</mml:mi><mml:mo>=</mml:mo><mml:mi>P</mml:mi><mml:mpadded width="+3.3pt"><mml:mi>P</mml:mi></mml:mpadded><mml:mo rspace="5.8pt">&#x00D7;</mml:mo><mml:mrow><mml:mo maxsize="210%" minsize="210%">[</mml:mo><mml:mo>-</mml:mo><mml:msup><mml:mn>0.0101</mml:mn><mml:mo>&#x2218;</mml:mo></mml:msup><mml:mpadded width="+3.3pt"><mml:msup><mml:mtext>C</mml:mtext><mml:mrow><mml:mo>-</mml:mo><mml:mn>1</mml:mn></mml:mrow></mml:msup></mml:mpadded><mml:mo rspace="5.8pt">&#x00D7;</mml:mo><mml:mi>S</mml:mi><mml:mi>S</mml:mi><mml:mpadded width="+3.3pt"><mml:mi>T</mml:mi></mml:mpadded><mml:mo rspace="5.8pt">+</mml:mo><mml:mpadded width="+3.3pt"><mml:mn>0.0582</mml:mn></mml:mpadded><mml:mo rspace="5.8pt">&#x00D7;</mml:mo></mml:mrow></mml:mrow></mml:mtd></mml:mtr><mml:mtr><mml:mtd columnalign="left"><mml:mspace width="5em"/><mml:mrow><mml:mi>ln</mml:mi><mml:mrow><mml:mo>(</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac><mml:mrow><mml:mi>P</mml:mi><mml:mi>P</mml:mi></mml:mrow><mml:msub><mml:mi>Z</mml:mi><mml:mrow><mml:mtext>eu</mml:mtext></mml:mrow></mml:msub></mml:mfrac></mml:mstyle><mml:mo>)</mml:mo></mml:mrow><mml:mo>+</mml:mo><mml:mn>0.419</mml:mn><mml:mo maxsize="210%" minsize="210%">]</mml:mo></mml:mrow></mml:mtd></mml:mtr></mml:mtable></mml:math></disp-formula>
<p>According to the expression 4&#x2013;1, the POC fluxes estimated from <italic>in situ</italic> measurements (PP<sub><italic>in situ</italic></sub>, SST<sub><italic>in situ</italic></sub> and Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub>) of <xref ref-type="bibr" rid="B41">Hung et al. (2000)</xref>; <xref ref-type="bibr" rid="B36">Hung and Gong (2007)</xref>, and <xref ref-type="bibr" rid="B69">Shih et al. (2015)</xref> were from 12 to 319 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>, an average of 20% less than the trap POC fluxes (25&#x2013;274 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>) (<xref ref-type="fig" rid="F6">Figure 6</xref>). As described above, the inputs of SST<sub><italic>in situ</italic></sub> and Z<sub>eu&#x2013;</sub><sub><italic>in situ</italic></sub> to the expression were fixed, the PP<sub><italic>in situ</italic></sub> was replaced by the PP<sub>VGPM</sub>, an average difference between estimated and trap POC fluxes (estimated POC flux: 16&#x2013;781 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>) was a factor of 2. It has been suggested that the uncertainty of these POC fluxes is quite large if satellite-based PP is used to estimate carbon sequestrations in oceans. If the discrepancy between <italic>in situ</italic> measurements and satellite-based observations of PP can be diminished and the reliance on them (e.g., PP<sub>VGPM</sub>) can be increased, it is to dedicate the potential importance and goal of the present study.</p>
<fig id="F6" position="float">
<label>FIGURE 6</label>
<caption><p>A comparison of the difference between <italic>in situ</italic> trap measured and estimated POC fluxes. The estimated POC fluxes were computed according to the empirical model expression revealed by <xref ref-type="bibr" rid="B23">Dunne et al. (2005)</xref>. The dashed line represented the proportional relationship of estimated POC flux and PP. The PP<sub><italic>in situ</italic></sub> and PP<sub>VGPM</sub> were also exerted on the expression to estimate POC fluxes (red square and blue triangle, respectively). <italic>In situ</italic> measurements of PP<sub><italic>in situ</italic></sub> and trap measured POC fluxes (green circle) were based on <xref ref-type="bibr" rid="B41">Hung et al. (2000)</xref>; <xref ref-type="bibr" rid="B36">Hung and Gong (2007)</xref>, and <xref ref-type="bibr" rid="B69">Shih et al. (2015)</xref>.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-08-747763-g006.tif"/>
</fig>
</sec>
<sec id="S4.SS2">
<title>Impact of Global Warming on <italic>in situ</italic> Observations</title>
<p>In both the Atlantic and Pacific oceans, increased phytoplankton biomass is observed at low latitudes during the boreal warm season. Other than equatorial upwelling, there are no other conspicuous seasonal trends in biogeochemical activities (<xref ref-type="bibr" rid="B20">Dandonneau et al., 2004</xref>). In the Arabian Sea, enhanced biogeochemical responses are also observed at low latitudes during the SWM (<xref ref-type="bibr" rid="B3">Banse and English, 2000</xref>). In this study, long-term and monthly variations in PP<sub><italic>in situ</italic></sub> and PP<sub>VGPM</sub> demonstrated influential monsoonal system at the SEATS (<xref ref-type="fig" rid="F2">Figures 2</xref>, <xref ref-type="fig" rid="F3">3</xref> and <xref ref-type="table" rid="T2">Table 2</xref>). The concurrence of low SST, deep Z<sub>ML</sub>, high Chl<sub>s</sub>, shallow Z<sub>eu</sub>, and increased PP suggest that increased phytoplankton biomass or specific phytoplankton communities dominating were triggered during the NEM. It has been reported that the average nitrate+nitrite (N+N) concentration, one of the key nutrients for phytoplankton growth in the SCS, in the MLD in the seasons of NEM (0.1&#x2013;0.3 &#x03BC;M) is &#x223C;10 times higher than of SWM (&#x223C;0.03 &#x03BC;M) (<xref ref-type="bibr" rid="B79">Tseng et al., 2005</xref>). Moreover, the inventory of N+N and the depth of nitracline in the NEM (30 &#x00B1; 19 mmol m<sup>&#x2013;</sup><sup>2</sup> and 28&#x2013;62 m, respectively) have been observed a &#x223C; 4.5 fold higher and a &#x223C; 25&#x2013;50% shallower than those reported in the SWM (7 &#x00B1; 4 mmol m<sup>&#x2013;</sup><sup>2</sup> and 52&#x2013;82 m, respectively) (<xref ref-type="bibr" rid="B14">Chen, 2005</xref>; <xref ref-type="bibr" rid="B70">Shih et al., 2020a</xref>). Evidences of abundant nutrient (e.g., N+N) and shallow nitracline depth favoring biological activities, imply that the growth of phytoplankton communities and associated photosynthesis were affected mainly by vertical advection providing inorganic nutrients from deeper waters, under the NEM system prevailing in the SCS (<xref ref-type="bibr" rid="B54">Liu et al., 2002</xref>; <xref ref-type="bibr" rid="B1">Bai et al., 2018</xref>; <xref ref-type="bibr" rid="B10">Chen et al., 2020</xref>; <xref ref-type="bibr" rid="B85">Zhou et al., 2020</xref>).</p>
<p>The global decrease in PP is particularly pronounced in high-latitude waters, which lose roughly 2,000 Mt-C y<sup>&#x2013;</sup><sup>1</sup> (Mt = 10<sup>12</sup> g), accounting for a 70% decline in carbon fixation <italic>via</italic> photosynthesis (<xref ref-type="bibr" rid="B29">Gregg et al., 2003</xref>). To compare annually reductions in PP<sub><italic>in situ</italic></sub> (<bold>&#x2212;</bold>11 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup> y<sup>&#x2013;</sup><sup>1</sup>) vs. the annual PP<sub><italic>in situ</italic></sub> (241 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>; mean PP<sub><italic>in situ</italic></sub> of NEM and SWM; <xref ref-type="table" rid="T1">Table 1</xref>), it indicated that the annual reduction in carbon fixation <italic>via</italic> photosynthesis was &#x223C; -5% y<sup>&#x2013;</sup><sup>1</sup>. Based on the 200 m isobath boundary of oligotrophic waters in the SCS (2.76 &#x00D7; 10<sup>12</sup> m<sup>2</sup>; <xref ref-type="bibr" rid="B53">Lin et al., 2003</xref>), the e-ratios were 5&#x2013;16% in the SCS (<xref ref-type="bibr" rid="B37">Hung and Gong, 2010</xref>; <xref ref-type="bibr" rid="B72">Shih et al., 2019</xref>) and the decreased in carbon fixation was roughly 11 Mt-C, thereby accounting for 30&#x2013;90% of the export production. This suggests a gradual decrease in the efficiency of photosynthetic carbon fixation by phytoplankton. Nevertheless, satellite-based observations do not show the signs of global warming on carbon fixation and sequestration.</p>
<p><xref ref-type="fig" rid="F5">Figure 5B</xref> illustrates the significantly negative relationship between PP<sub><italic>in situ</italic></sub> and SST<sub><italic>in situ</italic></sub> at the SEATS site. The slope of &#x2212;37 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>&#x00B0;C <sup>&#x2013;</sup><sup>1</sup> was exceptionally close to the &#x2212;36 mg-C m<sup>&#x2013;</sup><sup>2</sup> d<sup>&#x2013;</sup><sup>1</sup>&#x00B0;C <sup>&#x2013;</sup><sup>1</sup> reported in previous studies (<xref ref-type="bibr" rid="B15">Chen and Chen, 2006</xref>; <xref ref-type="bibr" rid="B17">Chen et al., 2007</xref>, <xref ref-type="bibr" rid="B18">2008</xref>). Scaling factors related to the increase in SST<sub><italic>in situ</italic></sub> caused by global changes are extremely complex (<xref ref-type="bibr" rid="B63">Sarmiento et al., 1998</xref>; <xref ref-type="bibr" rid="B38">Hung et al., 2010</xref>; <xref ref-type="bibr" rid="B1">Bai et al., 2018</xref>). Notwithstanding the complexity of factors governing SST<sub><italic>in situ</italic></sub>, they can still be used to estimate PP values. The differences between daytime and nighttime SST<sub><italic>in situ</italic></sub> values were statistically insignificant, during the NEM as well as the SWM (one-tail <italic>t</italic> test: <italic>p</italic> = 0.15 and 0.24, respectively (<xref ref-type="fig" rid="F6">Figure 6</xref>). We therefore surmise that SST sampling schedules had no effect on the overall results. Furthermore, we observed a strong statistically significant correlation between SST<sub>sate</sub> and SST<sub><italic>in situ</italic></sub> (<xref ref-type="fig" rid="F4">Figure 4C</xref>), indicating the efficacy of SST in estimating PP distributions over a broad horizontal area, regardless of the method used for derivation.</p>
<p>The straightforward relationship between PP<sub><italic>in situ</italic></sub> and SST<sub><italic>in situ</italic></sub> is important when seeking to predict PP values and estimate new or export production in euphotic zones. Under the environmental conditions described above, the reduction in carbon fixation due to photosynthesis would be &#x223C;&#x2212;15% &#x00B0;C<sup>&#x2013;</sup><sup>1</sup>, and the decrease in carbon fixation would be roughly 37 Mt-C, thereby accounting for a 1- to 3-fold quantity of export production. The negative correlation between PP<sub><italic>in situ</italic></sub> and SST<sub><italic>in situ</italic></sub> in the current study matched the findings observed in mid-latitude tropical/subtropical regions of the Pacific and Atlantic oceans (<xref ref-type="bibr" rid="B13">Chen, 2000</xref>; <xref ref-type="bibr" rid="B77">Tilstone et al., 2009</xref>), but differed drastically from those reported in high-latitude regions (<xref ref-type="bibr" rid="B47">Kudryavtseva et al., 2018</xref>).</p>
<p>Generally speaking, the sampling resolution of <italic>in situ</italic> time-series is too low to eliminate temporal uncertainty over all timespans. Fortunately, we can use SST<sub>sate</sub> to compensate for deficiencies in SST<sub><italic>in situ</italic></sub> coverage (<xref ref-type="bibr" rid="B1">Bai et al., 2018</xref>; <xref ref-type="bibr" rid="B10">Chen et al., 2020</xref>). Only one daily SST<sub>sate</sub> reading can be obtained at any given location; however, it would be perfectly reasonable to substitute that value with one obtained SST<sub><italic>in situ</italic></sub>. We observed a statistically significant linear relationship between SST<sub>sate</sub> with SST<sub><italic>in situ</italic></sub> (<xref ref-type="fig" rid="F4">Figure 4C</xref>); however, differences between daytime and nighttime SST<sub><italic>in situ</italic></sub> measurements did not reach the level of significance (<xref ref-type="fig" rid="F7">Figure 7</xref>). This suggests that SST estimates obtained using either method could be used to assess the influence of temperature on biogeochemical phenomena, such as PP.</p>
<fig id="F7" position="float">
<label>FIGURE 7</label>
<caption><p>Averaged diel records (&#x00B1;1 SD) of SST during the <bold>(A)</bold> NE-monsoon and <bold>(B)</bold> SW-monsoon. Horizontal dashed and solid lines indicate the mean daytime measurements (GMT+8: 06&#x2013;18 and 05&#x2013;19 h, NE- and SW-monsoons, respectively) and nighttime measurements (GMT+8: 19&#x2013;05 and 20&#x2013;04 h, NE- and SW-monsoons, respectively).</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-08-747763-g007.tif"/>
</fig>
<p>Asynchronous variations between PP<sub><italic>in situ</italic></sub> and PP<sub>VGPM</sub> and their related variables in the VGPM algorithm indicate the following: (1) Satellite-based evaluations depend primarily on the assumption of an exponential decrease in the vertical distribution of PP from the surface to the euphotic zone base. Nonetheless, remote sensing cannot penetrate beyond the surface, and therefore cannot reflect the true vertical distribution of PP at depth (<xref ref-type="bibr" rid="B38">Hung et al., 2010</xref>; <xref ref-type="bibr" rid="B6">Buitenhuis et al., 2013</xref>; <xref ref-type="bibr" rid="B68">Shih et al., 2013</xref>, <xref ref-type="bibr" rid="B71">2020b</xref>). This assumption is the primary cause for uncertainty between PP values obtained from satellite-based observations (i.e., PP<sub>VGPM</sub>) and those derived from <italic>in situ</italic> measurements (i.e., PP<sub><italic>in situ</italic></sub>). (2) When using satellite-based observations to estimate horizontal PP distributions, mathematical extrapolation/interpolation is commonly used to compensate for gaps in data coverage resulting from cloud coverage, heavy rains, rough seas, extreme weather events, natural episodes, suspended particles, and/or chromophoric dissolved organic matter. Thus, this approach cannot reflect &#x201C;true&#x201D; or &#x201C;<italic>in situ</italic>&#x201D; biogeochemistry responses to PP in the oceans. (<xref ref-type="bibr" rid="B5">Boyd and Trull, 2007</xref>; <xref ref-type="bibr" rid="B64">Shang et al., 2008</xref>; <xref ref-type="bibr" rid="B76">Tang et al., 2008</xref>; <xref ref-type="bibr" rid="B39">Hung et al., 2009</xref>, <xref ref-type="bibr" rid="B38">2010</xref>; <xref ref-type="bibr" rid="B70">Shih et al., 2020a</xref>). For decades, SEATS has been used as a natural laboratory for studies of prolonged environmental changes and reciprocal biogeochemical responses. It is time to tune existing models or develop more reliable models if we are to gain meaningful estimates of biogeochemical phenomena in the oceans. The proposed calibration method aimed at improving PP estimates for the VGPM is expected to enhance our understanding of changes in the SCS.</p>
</sec>
</sec>
<sec id="S5">
<title>Summary</title>
<p>Our time-series study (2003 &#x223C; 2016) at SEATS compared PP estimates based on <italic>in situ</italic> measurements and those based on the VGPM in the SCS during the NEM and SWM. PP values obtained during the NEM exceeded those obtained during the SWM, which appears to indicate that weather conditions during the cold season are conducive to high PP values. PP<sub><italic>in situ</italic></sub> values were roughly 50% lower than PP<sub>VGPM</sub> values, regardless of the season (NEM or SWM). These discrepancies can be attributed to the satellite-based integrated phytoplankton biomass in the euphotic zone. The discrepancies can be derived as the product of Z<sub>eu</sub> and Chl<sub>s</sub>, which are two main variables in the VGPM algorithm, especially the impact of difference between <italic>in situ</italic> and satellite-based Z<sub>eu</sub> on the magnitude of PP.</p>
<p>The observed overall decrease in PP<sub><italic>in situ</italic></sub> can be partially explained by an increase in SST<sub><italic>in situ</italic></sub>. Our results also showed that SST<sub>sate</sub> could be used to predict horizontal PP distributions over extended time scales, based on our observation of a statistically significant relationship between SST<sub>sate</sub> with SST<sub><italic>in situ</italic></sub>. A significantly negative relationship between PP<sub><italic>in situ</italic></sub> and SST<sub><italic>in situ</italic></sub> appears to indicate that global changes, such as oceanic warming, could have a negative impact on ocean biogeochemistry in low-latitude regions of the SCS. Nonetheless, further research will be required to assess the influence of global changes on biogeochemical phenomena, particularly in low-latitude waters. The SEATS has been used for decades to assess the sensitivity and resilience of low-latitude oceans to environmental fluctuations. Our analysis of discrepancies between <italic>in situ</italic> measurements and satellite-based observations could help to guide revisions aimed at enhancing the robustness and reliability of the VGPM in estimating biogeochemical responses. Satellite-based data could be used to expand the spatiotemporal scale of observations and thereby shed light on the actual biogeochemical effects of global environmental changes in low-latitude regions of the SCS.</p>
</sec>
<sec sec-type="data-availability" id="S6">
<title>Data Availability Statement</title>
<p>The original contributions presented in the study are included in the article/supplementary material, further inquiries can be directed to the corresponding author.</p>
</sec>
<sec id="S7">
<title>Author Contributions</title>
<p>Y-YS, F-KS, and C-CH wrote the manuscript with contributions from W-CC. C-YL, J-HT, Y-SW, and C-CL performed the experiments and created the tables and figures. Y-YS, F-KS, C-CH, W-CC, C-YL, J-HT, Y-SW, C-CL and C-YK reviewed and revised the manuscript. All authors listed have made substantial, direct, and intellectual contribution to the work and approved it for publication.</p>
</sec>
<sec sec-type="COI-statement" id="conf1">
<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 sec-type="disclaimer" id="S8">
<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>
</body>
<back>
<sec sec-type="funding-information" id="S9">
<title>Funding</title>
<p>This research was supported by the MOST (Ministry of Sciences and Technology, Taiwan) under grant numbers 108-2611-M-012-001, 108-2611-M-110-019-MY3, 109-2611-M-012-001, 109-2740-M-110-001, 110-2611-M-012-002, 110-2740-M-110-001, and 110-2611-M-019-005.</p>
</sec>
<ack>
<p>We would like to thank the assistance given us by the crews of R/V <italic>Ocean Researcher I</italic> and R/V <italic>Fishery Researcher I</italic>. We would also like to thank the unsung heroes and contributors who have contributed to the SEATS program.</p>
</ack>
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