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  <front>
    <journal-meta>
<journal-id journal-id-type="publisher">BG</journal-id>
<journal-title-group>
<journal-title>Biogeosciences</journal-title>
<abbrev-journal-title abbrev-type="publisher">BG</abbrev-journal-title>
<abbrev-journal-title abbrev-type="nlm-ta">Biogeosciences</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1726-4189</issn>
<publisher><publisher-name>Copernicus Publications</publisher-name>
<publisher-loc>Göttingen, Germany</publisher-loc>
</publisher>
</journal-meta>

    <article-meta>
      <article-id pub-id-type="doi">10.5194/bg-13-5151-2016</article-id><title-group><article-title>Climate change impacts on net primary production (NPP) and export production
(EP) regulated by increasing stratification and phytoplankton community
structure in the CMIP5 models</article-title>
      </title-group><?xmltex \runningtitle{Climate change impacts on NPP and EP}?><?xmltex \runningauthor{W.~Fu et al.}?>
      <contrib-group>
        <contrib contrib-type="author" corresp="yes" rid="aff1">
          <name><surname>Fu</surname><given-names>Weiwei</given-names></name>
          <email>weiweif@uci.edu</email>
        <ext-link>https://orcid.org/0000-0003-4965-0832</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Randerson</surname><given-names>James T.</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Moore</surname><given-names>J. Keith</given-names></name>
          
        </contrib>
        <aff id="aff1"><institution>Department of Earth System Science, University of California, Irvine,
California, 92697, USA</institution>
        </aff>
      </contrib-group>
      <author-notes><corresp id="corr1">Weiwei Fu (weiweif@uci.edu)</corresp></author-notes><pub-date><day>16</day><month>September</month><year>2016</year></pub-date>
      
      <volume>13</volume>
      <issue>18</issue>
      <fpage>5151</fpage><lpage>5170</lpage>
      <history>
        <date date-type="received"><day>16</day><month>June</month><year>2015</year></date>
           <date date-type="rev-request"><day>12</day><month>August</month><year>2015</year></date>
           <date date-type="rev-recd"><day>10</day><month>July</month><year>2016</year></date>
           <date date-type="accepted"><day>3</day><month>August</month><year>2016</year></date>
      </history>
      <permissions>
<license license-type="open-access">
<license-p>This work is licensed under a Creative Commons Attribution 3.0 Unported License. To view a copy of this license, visit <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/3.0/">http://creativecommons.org/licenses/by/3.0/</ext-link></license-p>
</license>
</permissions><self-uri xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016.html">This article is available from https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016.html</self-uri>
<self-uri xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016.pdf">The full text article is available as a PDF file from https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016.pdf</self-uri>


      <abstract>
    <p>We
examine climate change impacts on net primary production (NPP) and export
production (sinking particulate flux; EP) with simulations from nine Earth
system models (ESMs) performed in the framework of the fifth phase of the Coupled Model
Intercomparison Project (CMIP5). Global NPP and EP are reduced by the end of
the century for the intense warming scenario of Representative Concentration
Pathway (RCP) 8.5. Relative to the 1990s, NPP in the 2090s is reduced by
2–16 % and EP by 7–18 %. The models with the largest increases in
stratification (and largest relative declines in NPP and EP) also show the
largest positive biases in stratification for the contemporary period,
suggesting overestimation of climate change impacts on NPP and EP. All of the
CMIP5 models show an increase in stratification in response to surface–ocean
warming and freshening, which is accompanied by decreases in surface
nutrients, NPP and EP.</p>
    <p>There is considerable variability across the models in the magnitudes of NPP,
EP, surface nutrient concentrations and their perturbations by climate
change. The negative response of NPP and EP to increasing stratification
reflects primarily a bottom-up control, as upward nutrient flux declines at
the global scale. Models with dynamic phytoplankton community structure show
larger declines in EP than in NPP. This pattern is driven by phytoplankton
community composition shifts, with reductions in productivity by large
phytoplankton as smaller phytoplankton (which export less efficiently) are
favored under the increasing nutrient stress. Thus, the projections of the
NPP response to climate change are critically dependent on the simulated
phytoplankton community structure, the efficiency of the biological pump and
the resulting levels of regenerated production, which vary widely across the
models. Community structure is represented simply in the CMIP5 models, and
should be expanded to better capture the spatial patterns and climate-driven
changes in export efficiency.</p>
  </abstract>
    </article-meta>
  </front>
<body>
      

<sec id="Ch1.S1" sec-type="intro">
  <title>Introduction</title>
      <p>Ocean net primary production (NPP) and particulate organic carbon export (EP)
are key elements of marine biogeochemistry that are vulnerable to ongoing
climate change from rising concentrations of atmospheric CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> and other
greenhouse gases. Ocean warming has increasing impacts on ocean ecosystems by
modifying the ecophysiology and distribution of marine organisms, and by
altering ocean circulation and stratification. Ocean ecosystems also are
important components of the climate system, influencing the atmospheric
abundance of radiative agents such as CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula>, N<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula>O, aerosols and the
bio-optical properties of seawater (Siegenthaler and Wenk, 1984; Goldstein et
al., 2003; Manizza et al., 2008; Schmittner et al., 2008; Bopp et al., 2013).
Therefore, understanding the mechanisms controlling NPP and EP is essential
for understanding the global cycles of carbon and other bioactive elements,
and their links to climate (Passow and Carlson, 2012).</p>
      <p>Upper ocean stratification plays a key role in ocean biogeochemical
processes. In particular, mixed layer depth (MLD) regulates the interplay
between light availability for photosynthesis (Hannon et al., 2001) and
nutrient supply to the upper ocean (Pollard et al., 2009). Upper ocean
stratification is defined here as the density difference between the surface
and 200 m depth (Capotondi et al., 2012), which is indicative of the degree
of coupling and nutrient fluxes between the euphotic zone and the ocean
interior. The density gradient at the base of the mixed layer affects
entrainment processes, which play a crucial role in mixed layer deepening and
in particle sinking/export from the euphotic zone. Stratification can also
influence ocean ventilation (Luo et al., 2009), which has important
consequences for oceanic uptake of carbon and oxygen. Thus, changes in
stratification over the remainder of the 21st century have the potential to
influence NPP and EP across marine ecosystems.</p>
      <p>Stratification tends to increase in response to ocean surface warming and
freshening in 21st century climate change simulations. Increased
stratification reduces the input of subsurface nutrients to the euphotic
zone and can lead to decreasing NPP and EP through increasing nutrient
limitation. Many studies have suggested decreases in global NPP and EP over
the 21st century using models with varying degrees of complexity (Bopp et al.,
2001; Plattner et al., 2001; Fung et al., 2005; Schmittner et al., 2008;
Steinacher et al., 2009; Dutkiewicz et al., 2013; Cabré et al., 2015).
For the RCP8.5 scenario, CMIP5 ESM estimates of changes in export production
range from <inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>7 to <inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>18 % relative to the 1990s, and for NPP these
changes range from <inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>2 to <inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>16 % (Bopp et al., 2013).</p>
      <p>The relative importance of different ecological controls on NPP and EP
depends, in part, on an individual model's capacity to represent plankton
functional types (PFTs) (Le Quéré et al., 2005; Jin et al., 2006) and
their unique physiological and ecological characteristics, which determine
how efficiently they are exported from surface waters. Increasing nutrient
stress can shift phytoplankton community composition, favoring smaller
phytoplankton, which are more efficient at nutrient uptake, over larger
phytoplankton (Bopp et al., 2001; Steinacher et al., 2010; Vichi et al.,
2011; Moore et al., 2013). These community shifts can modify the efficiency
of carbon export. However, treatment of plankton communities is relatively
simple in the CMIP5 models, with 1–3 phytoplankton functional types and
typically one zooplankton group (Bopp et al., 2013).</p>
      <p>Several previous studies examined the biogeochemical response to climate
change in the CMIP5 models. Bopp et al. (2013) examined output from 10 CMIP5
models emphasizing model mean biogeochemical responses to multiple stressors
and trends over the 21st century relative to 1990s means for each model.
Cabré et al. (2015) analyzed the CMIP5 models examining changes between
model output averaged over the period 1980–1999 with years 2080–2099. This
study broke down the global output into different ocean biomes for analysis.
Laufkötter et al. (2015) also analyzed output from nine coupled
climate–carbon ESMs, including many of the CMIP5 models to study how climate
change processes impact NPP, comparing two 20-year periods (2012–2031
and 2081–2100). They suggested strong roles for temperature and top-down
grazing control in driving the NPP response, particularly at lower latitudes.
Both Cabré et al. (2015) and Laufkötter et al. (2015) conclude that
changing light levels were not a primary driver of changes in NPP except at
the highest latitudes where there were large decreases in sea ice cover.
Thus, we do not consider light effects in this work, where our focus is on
global-scale trends. More detailed regional studies of the CMIP5 model output
have been carried out for the Arctic Ocean (Vancoppenolle et al., 2013) and
the Southern Ocean (Hauck and Volker, 2015; Ito et al., 2015; Leung et al.,
2015).</p>
      <p>We analyzed centennial-scale changes in NPP and EP in response to increasing
surface stratification and other physical factors. We use historical
(1850–2005) and Representative Concentration Pathway (RCP) 8.5 (2006–2100)
ESM simulations from the fifth phase of the Coupled Model Intercomparison
Project (CMIP5) to study long-term trends in NPP and EP and to identify the
mechanisms behind these changes, including the physical factors that regulate
nutrient availability. We also examined variability in NPP, EP and surface
nutrient concentrations across the models, to highlight some of the large
differences and uncertainties in projections of climate change impacts on
marine biogeochemistry in current-generation ESMs.</p>
</sec>
<sec id="Ch1.S2">
  <title>Methods</title>
      <p>We analyzed simulations from a set of nine ESMs that contributed output to the
Earth System Grid Federation as a part of CMIP5 (Taylor et al., 2012).
Required physical ocean variables were temperature, salinity and potential
density; required biogeochemistry variables were macronutrients (nitrate,
phosphate and silicic acid), iron, chlorophyll, NPP and EP. The selection of
the nine models investigated here (Table 1) was based on the availability of
these variables.</p>

<?xmltex \floatpos{t}?><table-wrap id="Ch1.T1" specific-use="star"><caption><p>A brief description of components of the ESMs used in this
study. For atmosphere and ocean components, the number of levels in
the vertical is indicated by “lev”  and the horizontal resolution
is indicated in degrees; vertical coordinates of the ocean and biogeochemical
components are indicated by <inline-formula><mml:math display="inline"><mml:mi>Z</mml:mi></mml:math></inline-formula> (geopotential) or <inline-formula><mml:math display="inline"><mml:mi>I</mml:mi></mml:math></inline-formula> (isopycnal).</p></caption><oasis:table frame="topbot"><?xmltex \begin{scaleboxenv}{.87}[.87]?><oasis:tgroup cols="8">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="left"/>
     <oasis:colspec colnum="3" colname="col3" align="left"/>
     <oasis:colspec colnum="4" colname="col4" align="left"/>
     <oasis:colspec colnum="5" colname="col5" align="left"/>
     <oasis:colspec colnum="6" colname="col6" align="left"/>
     <oasis:colspec colnum="7" colname="col7" align="left"/>
     <oasis:colspec colnum="8" colname="col8" align="left"/>
     <oasis:thead>
       <oasis:row>  
         <oasis:entry colname="col1">Model</oasis:entry>  
         <oasis:entry rowsep="1" namest="col2" nameend="col3" align="center">Resolution </oasis:entry>  
         <oasis:entry colname="col4">Vertical</oasis:entry>  
         <oasis:entry colname="col5">Reference</oasis:entry>  
         <oasis:entry colname="col6">Biogeochemical</oasis:entry>  
         <oasis:entry colname="col7">References</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">Atmosphere</oasis:entry>  
         <oasis:entry colname="col3">Ocean</oasis:entry>  
         <oasis:entry colname="col4">coordinate</oasis:entry>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6">component</oasis:entry>  
         <oasis:entry colname="col7"/>  
         <oasis:entry colname="col8"/>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>  
         <oasis:entry colname="col1">GFDL-ES2M</oasis:entry>  
         <oasis:entry colname="col2">24 lev, <inline-formula><mml:math display="inline"><mml:mrow><mml:mn>2.5</mml:mn><mml:mo>/</mml:mo><mml:mn>2.0</mml:mn></mml:mrow></mml:math></inline-formula><inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">50 lev, 0,3–1<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col4"><inline-formula><mml:math display="inline"><mml:mi>Z</mml:mi></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col5">Dunne et al. (2013a)</oasis:entry>  
         <oasis:entry colname="col6">TOPAZ2</oasis:entry>  
         <oasis:entry colname="col7">Dunne et al. (2013b)</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">GFDL-ES2G</oasis:entry>  
         <oasis:entry colname="col2">24 lev, <inline-formula><mml:math display="inline"><mml:mrow><mml:mn>2.5</mml:mn><mml:mo>/</mml:mo><mml:mn>2.0</mml:mn></mml:mrow></mml:math></inline-formula><inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">50 lev, 0,3–1<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col4"><inline-formula><mml:math display="inline"><mml:mrow><mml:mi>Z</mml:mi><mml:mo>+</mml:mo><mml:mi>I</mml:mi></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col5">Dunne et al. (2013a)</oasis:entry>  
         <oasis:entry colname="col6">TOPAZ2</oasis:entry>  
         <oasis:entry colname="col7">Dunne et al. (2013b)</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">MPI-ESM-LR</oasis:entry>  
         <oasis:entry colname="col2">47 lev, 1.9<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">40 lev, 1.5<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col4"><inline-formula><mml:math display="inline"><mml:mi>Z</mml:mi></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col5">Giorgetta et al. (2013)</oasis:entry>  
         <oasis:entry colname="col6">HAMOCC5.2</oasis:entry>  
         <oasis:entry colname="col7">Ilyina et al. (2013)</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">MPI-ESM-MR</oasis:entry>  
         <oasis:entry colname="col2">47 lev, 1.9<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">40 lev, 0.4<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col4"><inline-formula><mml:math display="inline"><mml:mi>Z</mml:mi></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col5">Giorgetta et al. (2013)</oasis:entry>  
         <oasis:entry colname="col6">HAMOCC5.2</oasis:entry>  
         <oasis:entry colname="col7">Ilyina et al. (2013)</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">IPSL-CM5A-LR</oasis:entry>  
         <oasis:entry colname="col2">39 lev, <inline-formula><mml:math display="inline"><mml:mrow><mml:mn>1.9</mml:mn><mml:mo>/</mml:mo><mml:mn>3.8</mml:mn></mml:mrow></mml:math></inline-formula><inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">31 lev, 0.5–2<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col4"><inline-formula><mml:math display="inline"><mml:mi>Z</mml:mi></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col5">Dufresne et al. (2013)</oasis:entry>  
         <oasis:entry colname="col6">PISCES</oasis:entry>  
         <oasis:entry colname="col7">Aumont and Bopp (2006)</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">IPSL-CM5A-MR</oasis:entry>  
         <oasis:entry colname="col2">39 lev, <inline-formula><mml:math display="inline"><mml:mrow><mml:mn>1.2</mml:mn><mml:mo>/</mml:mo><mml:mn>1.9</mml:mn></mml:mrow></mml:math></inline-formula><inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">31 lev, 0.5–2<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col4"><inline-formula><mml:math display="inline"><mml:mi>Z</mml:mi></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col5">Dufresne et al. (2013)</oasis:entry>  
         <oasis:entry colname="col6">PISCES</oasis:entry>  
         <oasis:entry colname="col7">Aumont and Bopp (2006)</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>  
         <oasis:entry colname="col7">Seferian et al. (2013)</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">HadGEM2-ES</oasis:entry>  
         <oasis:entry colname="col2">38 lev, <inline-formula><mml:math display="inline"><mml:mrow><mml:mn>1.2</mml:mn><mml:mo>/</mml:mo><mml:mn>1.9</mml:mn></mml:mrow></mml:math></inline-formula><inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">40 lev, 0.3–1<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col4"><inline-formula><mml:math display="inline"><mml:mi>Z</mml:mi></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col5">Jones et al. (2011)</oasis:entry>  
         <oasis:entry colname="col6">Diat-HadOCC</oasis:entry>  
         <oasis:entry colname="col7">Palmer and Totterdell (2000)</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">Collins et al. (2011)</oasis:entry>  
         <oasis:entry colname="col6"/>  
         <oasis:entry colname="col7"/>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">CESM1(BGC)</oasis:entry>  
         <oasis:entry colname="col2">26 lev, <inline-formula><mml:math display="inline"><mml:mrow><mml:mn>1.25</mml:mn><mml:mo>/</mml:mo><mml:mn>0.94</mml:mn></mml:mrow></mml:math></inline-formula><inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">60 lev, 1.125<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col4"><inline-formula><mml:math display="inline"><mml:mi>Z</mml:mi></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col5">Gent et al. (2011)</oasis:entry>  
         <oasis:entry colname="col6">BEC</oasis:entry>  
         <oasis:entry colname="col7">Moore et al. (2004)</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"><inline-formula><mml:math display="inline"><mml:mrow><mml:mo>/</mml:mo><mml:mn>0.27</mml:mn></mml:mrow></mml:math></inline-formula>–0.53<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">Lindsay et al. (2014)</oasis:entry>  
         <oasis:entry colname="col6"/>  
         <oasis:entry colname="col7">Doney et al. (2009)</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">NorESM1-ME</oasis:entry>  
         <oasis:entry colname="col2">26 lev, 1.9<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">70 lev, 1.5<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col4"><inline-formula><mml:math display="inline"><mml:mi>I</mml:mi></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col5">Bentsen et al. (2013)</oasis:entry>  
         <oasis:entry colname="col6">HAMOCC5.1</oasis:entry>  
         <oasis:entry colname="col7">Tjiputra et al. (2013)</oasis:entry>  
         <oasis:entry colname="col8"/>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup><?xmltex \end{scaleboxenv}?></oasis:table></table-wrap>

      <p>The historical and RCP8.5 simulations we analyzed had prescribed atmospheric
CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> mole fractions and forcing from other greenhouse gases and aerosols,
anthropogenic land use and solar variability. Volcanic forcing also was
included during the historical period. The RCP8.5 is a strong warming
scenario with an increase in radiative forcing of 8.5 W m<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula> by 2100 as
atmospheric CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> mole fractions reach 936 ppm (Moss et al., 2010; van
Vuuren et al., 2011). In the case where several ensemble members were
available from an individual ESM, we analyzed only the first member.</p>
      <p>A simple description of the nine ESMs is presented in Tables 1 and 2.
Atmospheric and ocean resolutions vary across the models (Table 1). Typical
atmospheric horizontal grid resolution is <inline-formula><mml:math display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 2<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>, but it ranges
from 0.94 to 3.8<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>. Typical ocean horizontal resolution is
<inline-formula><mml:math display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 1<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>, ranging from 0.3 to 2<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>. In the vertical, there
are 24–95 levels in the atmosphere and 31–63 levels in the ocean. All
marine biogeochemical components are
nutrient–phytoplankton–zooplankton–detritus (NPZD) models, but with
varying degrees of complexity illustrated, for instance, by the number of
phytoplankton functional groups (from 1 to 3) or limiting nutrients (from 3
to 5) that are explicitly represented (Table 2).</p>
      <p>In our analysis, we used the CMIP5 variable denoting the vertical integration
of NPP (intpp) and sinking export of organic particles at 100 m (EP;
epc100). We present global mean estimates as the area-weighted or
volume-weighted mean by the grid-cell area/volume from an individual model.
Monthly mean data are averaged to obtain annual means and the annual mean
data are interpolated onto a common 1<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> <inline-formula><mml:math display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 1<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> regular
grid for the comparison of the 2-D fields.</p>

<?xmltex \floatpos{t}?><table-wrap id="Ch1.T2" specific-use="star"><caption><p>A brief description of the marine biogeochemical components
included in the
ESMs. Nutrients limiting phytoplankton growth, the number of explicit phytoplankton groups, the
number of explicit zooplankton groups, representation of heterotrophic bacteria, the use of
fixed (Redfield: <inline-formula><mml:math display="inline"><mml:mi>R</mml:mi></mml:math></inline-formula>) or variable (<inline-formula><mml:math display="inline"><mml:mi>V</mml:mi></mml:math></inline-formula>) ratios for organic matter production, and <inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi>Q</mml:mi><mml:mn>10</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> for
temperature dependency of biogeochemical processes (autotrophic/heterotrophic) are indicated.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="6">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="left"/>
     <oasis:colspec colnum="3" colname="col3" align="left"/>
     <oasis:colspec colnum="4" colname="col4" align="center"/>
     <oasis:colspec colnum="5" colname="col5" align="left"/>
     <oasis:colspec colnum="6" colname="col6" align="center"/>
     <oasis:thead>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">Model</oasis:entry>  
         <oasis:entry colname="col2">Nutrients</oasis:entry>  
         <oasis:entry colname="col3">Phytoplankton</oasis:entry>  
         <oasis:entry colname="col4">Zooplankton</oasis:entry>  
         <oasis:entry colname="col5">Organic material ratio</oasis:entry>  
         <oasis:entry colname="col6"><inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi>Q</mml:mi><mml:mn>10</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>  
         <oasis:entry colname="col1">TOPAZ2</oasis:entry>  
         <oasis:entry colname="col2">5 (<inline-formula><mml:math display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">NO</mml:mi><mml:mn mathvariant="normal">3</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">NH</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">PO</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>,</mml:mo></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">3 (diatom, eukaryotes,</oasis:entry>  
         <oasis:entry colname="col4">1</oasis:entry>  
         <oasis:entry colname="col5"><inline-formula><mml:math display="inline"><mml:mi>R</mml:mi></mml:math></inline-formula>(C : N)</oasis:entry>  
         <oasis:entry colname="col6">1.88</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"><inline-formula><mml:math display="inline"><mml:mrow><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">SiO</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">F</mml:mi><mml:mi mathvariant="normal">e</mml:mi></mml:msub></mml:mrow><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">small diazotrophs)</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"><inline-formula><mml:math display="inline"><mml:mrow><mml:mi>V</mml:mi><mml:mrow class="chem"><mml:mo>(</mml:mo><mml:mi mathvariant="normal">P</mml:mi><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">S</mml:mi><mml:mi mathvariant="normal">i</mml:mi></mml:msub><mml:mo>,</mml:mo><mml:mi mathvariant="normal">Chl</mml:mi><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">F</mml:mi><mml:mi mathvariant="normal">e</mml:mi></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">HAMOCC5.2</oasis:entry>  
         <oasis:entry colname="col2">3 (<inline-formula><mml:math display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">NO</mml:mi><mml:mn mathvariant="normal">3</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">F</mml:mi><mml:mi mathvariant="normal">e</mml:mi></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">PO</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">1 (separated into</oasis:entry>  
         <oasis:entry colname="col4">1</oasis:entry>  
         <oasis:entry colname="col5"><inline-formula><mml:math display="inline"><mml:mi>R</mml:mi></mml:math></inline-formula>(C : N : P : Fe)</oasis:entry>  
         <oasis:entry colname="col6">1.88</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">diatoms and calcifiers)</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">HAMOCC5.1</oasis:entry>  
         <oasis:entry colname="col2">3 (<inline-formula><mml:math display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">NO</mml:mi><mml:mn mathvariant="normal">3</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">F</mml:mi><mml:mi mathvariant="normal">e</mml:mi></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">PO</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">1 (separated into</oasis:entry>  
         <oasis:entry colname="col4">1</oasis:entry>  
         <oasis:entry colname="col5"><inline-formula><mml:math display="inline"><mml:mi>R</mml:mi></mml:math></inline-formula>(C:N:P:Fe)</oasis:entry>  
         <oasis:entry colname="col6">1.88</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">diatoms and calcifiers)</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">PISCES</oasis:entry>  
         <oasis:entry colname="col2">5 (<inline-formula><mml:math display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">NO</mml:mi><mml:mn mathvariant="normal">3</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">F</mml:mi><mml:mi mathvariant="normal">e</mml:mi></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">PO</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>,</mml:mo></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">2 (diatoms and</oasis:entry>  
         <oasis:entry colname="col4">2 (micro and</oasis:entry>  
         <oasis:entry colname="col5"><inline-formula><mml:math display="inline"><mml:mi>R</mml:mi></mml:math></inline-formula>(C : N : P)</oasis:entry>  
         <oasis:entry colname="col6"><inline-formula><mml:math display="inline"><mml:mrow><mml:mn>1.88</mml:mn><mml:mo>/</mml:mo><mml:mn>2.14</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"><inline-formula><mml:math display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">NH</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">SiO</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">nanophytoplankton)</oasis:entry>  
         <oasis:entry colname="col4">meso)</oasis:entry>  
         <oasis:entry colname="col5"><inline-formula><mml:math display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">V</mml:mi><mml:mo>(</mml:mo><mml:msub><mml:mi mathvariant="normal">S</mml:mi><mml:mi mathvariant="normal">i</mml:mi></mml:msub><mml:mo>,</mml:mo><mml:mi mathvariant="normal">Chl</mml:mi><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">F</mml:mi><mml:mi mathvariant="normal">e</mml:mi></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Diat-HadOCC</oasis:entry>  
         <oasis:entry colname="col2">4 (<inline-formula><mml:math display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">NO</mml:mi><mml:mn mathvariant="normal">3</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">F</mml:mi><mml:mi mathvariant="normal">e</mml:mi></mml:msub><mml:mo>,</mml:mo></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">2 (diatoms and</oasis:entry>  
         <oasis:entry colname="col4">1</oasis:entry>  
         <oasis:entry colname="col5"><inline-formula><mml:math display="inline"><mml:mi>R</mml:mi></mml:math></inline-formula>(C : N)</oasis:entry>  
         <oasis:entry colname="col6">none</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"><inline-formula><mml:math display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">NH</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">SiO</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">nondiatom)</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"><inline-formula><mml:math display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">V</mml:mi><mml:mo>(</mml:mo><mml:msub><mml:mi mathvariant="normal">S</mml:mi><mml:mi mathvariant="normal">i</mml:mi></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">F</mml:mi><mml:mi mathvariant="normal">e</mml:mi></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">BEC</oasis:entry>  
         <oasis:entry colname="col2">5 (<inline-formula><mml:math display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">NO</mml:mi><mml:mn mathvariant="normal">3</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">NH</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">PO</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>,</mml:mo></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">3 (diatom, nano-,</oasis:entry>  
         <oasis:entry colname="col4">1</oasis:entry>  
         <oasis:entry colname="col5"><inline-formula><mml:math display="inline"><mml:mi>R</mml:mi></mml:math></inline-formula>(C : N : P)</oasis:entry>  
         <oasis:entry colname="col6">2.0</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"><inline-formula><mml:math display="inline"><mml:mrow><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">SiO</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">F</mml:mi><mml:mi mathvariant="normal">e</mml:mi></mml:msub></mml:mrow><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col3">phyto, diazotrophs)</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"><inline-formula><mml:math display="inline"><mml:mrow><mml:mi>V</mml:mi><mml:mrow class="chem"><mml:mo>(</mml:mo><mml:msub><mml:mi mathvariant="normal">S</mml:mi><mml:mi mathvariant="normal">i</mml:mi></mml:msub><mml:mo>,</mml:mo><mml:mi mathvariant="normal">Chl</mml:mi><mml:mo>,</mml:mo><mml:msub><mml:mi mathvariant="normal">F</mml:mi><mml:mi mathvariant="normal">e</mml:mi></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

</sec>
<sec id="Ch1.S3">
  <title>Results</title>
<sec id="Ch1.S3.SS1">
  <title>Stratification changes</title>
      <p>Stratification, defined here as the density difference between the depth of
200 m and the surface, is a useful indicator of change in the upper ocean,
as it integrates changes in both temperature and salinity. In Fig. 1a, we
present the time series of global mean stratification changes for the
historical period and the RCP8.5 projection. All the models project an
increase in stratification (ranging from 6 to 30 % by the 2090s).
However, the amplitude of stratification differs considerably across the
models. The GFDL-ESM2M and MPI models are relatively close to the observed
mean stratification in the WOA09 data set (red square, 1.81 kg m<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> for
the present era. NorESM1-ME shows the weakest stratification
(1.74 kg m<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> while the stratification in HadGEM2-ES is strongest
(2.45 kg m<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>. Long-term trends are in general agreement across
models, but the rate of stratification increase varies, with IPSL-CM5A-MR
showing the most rapid increase and NorESM1-ME the slowest increase.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F1"><caption><p>Time series of global mean surface stratification, SST and SSS for
historical run and RCP8.5 over 1850–2100. Surface stratification is defined
as the density difference between 200 m and the surface. Red square
indicates observations from the WOA2009 data.</p></caption>
          <?xmltex \igopts{width=241.848425pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f01.png"/>

        </fig>

      <p>Surface processes that decrease density can largely explain the
stratification increase in the RCP8.5 projections. Global mean sea surface
temperature (SST) warms by 2.6–3.5 <inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C, accompanied by sea surface
salinity (SSS) decreases of 0.05–0.25 psu over the 21st century (Fig. 1).
By 2100, the global mean SST ranges from 20.4 <inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C (HadGEM2-ES) to
21.8 <inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C (NorESM1-ME). Model spread decreases in the RCP8.5
projections in response to strong anthropogenic forcing (Fig. 1b). SSS shows
a clear declining tendency from 1850 to 2100 (Fig. 1). Compared to the WOA09
observational data, most of the models are too fresh at the surface in the
1990s, especially the HadGEM2-ES, which has the lowest global mean SSS. The
model spread is partly due to internal variability simulated by the climate
models. Model differences in physics, but also in spin-up procedures, the way
RCP scenarios are set up and model climate sensitivities all likely
contribute to the model spread (Knutti and Hegerl, 2008; Szopa et al., 2013).</p>
      <p>Vertical density profiles help to further explain the changes in
stratification. Mean vertical profiles of density in the 1990s and the
density change between the 1990s and the 2090s show that all the models
become more buoyant at the surface as a consequence of heating and/or
freshening of the upper ocean (Fig. S1 in the Supplement). The density
changes at the surface vary by almost a factor of 2 among models, from
<inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>1.1 kg m<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula> (HadGEM-ES2) to <inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>0.6 kg m<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula> (GFDL-ES2M), but
converge to a relatively narrow range (approximately <inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>0.2 kg m<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> at
500 m depth. Most of the density change occurs between the surface and
200 m. Below 200 m, the density change in most of the models varies
linearly with depth. Thus, our definition of the stratification index, as the
density difference between the surface and 200 m, is reasonable. The
converging reductions in density among models at about 500 m agrees with
some previous studies based on observations and CMIP3 models (Bindoff et al.,
2007; Lyman et al., 2010; Capotondi et al., 2012). Compared to WOA09 data,
the models generally underestimate the density of the upper ocean in the top
150 m and most models overestimate the density below 350 m (resulting in a
positive stratification bias) (Fig. S1a).</p>
      <p>Vertical profiles of temperature and salinity from each model are also shown
in Fig. S1. The surface ocean exhibits strong warming of 1.7–3.5 <inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C
by the 2090s and the warming magnitude declines quickly with depth, which is
associated with the heat uptake capacity of individual models. For instance,
the GFDL models seem to be more efficient in transporting heat downward than
the IPSL models. Above 300 m, the temperature changes vary widely among the
models. Temperature changes as a function of depth are complex, and
model-to-model differences may be related to a number of factors including
rates of vertical mixing and the seasonal thermocline dynamics. At the depth
of 500 m, the mean temperature change converges at about 1.2 <inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C.
The ocean heat uptake capacity is linked to ocean diapycnal mixing and other
processes, such as mixing by mesoscale eddies. The weak temperature gradients
in the GFDL models suggest high rate of heat uptake, and are consistent with
ocean heat uptake estimates by Kuhlbrodt and Gregory (2012). The large model
spread in temperature profiles suggests considerable differences and
uncertainties in the parameterizations of these physical processes across the
models. Vertical profiles of salinity are more scattered than for temperature
(Fig. S1c). In the 1990s, most of the models underestimate salinity from the
surface down to 550 m. Surface salinity is generally biased low by
0.05–0.25 psu. Most of the freshening with climate change takes place above
100 m, which also acts to increase stratification. Note that the salinity
increases at 100–300 m in some models (IPSL, GFDL-ESM2M, HadGEM2-ES)
partially compensates the impact of rising temperatures on density.</p>
      <p>The percentage contribution of temperature change to the stratification
change from the 1990s to the 2090s is shown in Fig. S2. Previous studies have
shown that salinity contributes significantly to the stratification changes
at high latitudes (&gt; 40<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>) in both hemispheres and in the
North Pacific as a consequence of increases in precipitation (Bindoff et al.,
2007). From our comparisons, temperature dominates the stratification changes
in the tropical and subtropical regions (Fig. S2). Salinity dominates the
stratification changes in the much of the Arctic Ocean and in the
high-latitude North Atlantic. While stratification is a function of SSS and
SST to a good approximation (Cabré et al., 2015), stratification change
at high latitudes is also dependent on temperature and salinity at depth as
vertical mixing and exchange are stronger.</p>
      <p>In some regions, the spatial distributions and the driving process differs
substantially across models. Generally, the models agree well in the tropics
and in the subtropical gyres that surface warming drives the increase of stratification.
In the high-latitude North Atlantic, the subpolar Pacific and the western
Pacific Ocean, there is weaker agreement across the models. In the
subtropical gyre of the South Pacific, stratification changes in the IPSL and
CESM1(BGC) models have a stronger influence from temperature change, while
the other models exhibit more complicated spatial patterns. In the North
Atlantic, salinity contributes more in the IPSL and HadGEM2 models than in
the other models. The southeastern Pacific is more dominated by salinity in
the two GFDL models. In the Southern Ocean, the models show relatively large
contributions from both salinity and temperature but with complicated spatial
patterns that differ considerably across models. Projections for the regions
where the models do not agree even on the driving factor should be viewed
with more caution. Climate change and biogeochemical impacts in these regions
tend to be projected with less consistency across models (Bopp et al., 2013;
Cabré et al., 2015).</p>
      <p>Stratification increases globally in all the models with climate change
(Figs. 1 and 2). Nearly all the models predict large increases in
stratification in the western tropical Pacific, the tropical Indian Ocean,
the Arctic Ocean and in the high-latitude North Atlantic (particularly in
the Labrador Sea). The Southern Ocean has weaker increases in stratification,
partly because the surface layer mixing and upwelling are intensified due to
the poleward shift of strengthened westerly jets (Swart and Fyfe, 2012). Our
stratification index may underestimate the changes in the high-latitude North
Atlantic, as the relatively deep mixing means that temperature and salinity
at 200 m depth are changing much more rapidly than in other regions.
Reductions in the deep winter mixing and NADW formation in this region are a
common pattern seen in strong warming climate simulations (e.g., Cheng et
al., 2013; Schwinger et al., 2014). Less drastic increases in stratification
are seen over much of the rest of the oceans, with only a few small regions
showing decreases in some models. An exception is the HadGEM2-ES model, which
has large stratification reductions in the Arctic (Fig. 2).</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F2" specific-use="star"><caption><p>The spatial pattern is shown for changes in stratification between
the 1990s and the 2090s.</p></caption>
          <?xmltex \igopts{width=369.885827pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f02.pdf"/>

        </fig>

</sec>
<sec id="Ch1.S3.SS2">
  <title>Surface nutrient trends with climate change</title>
      <p>One of the key factors determining global NPP is nutrient availability in the
euphotic zone. Time series of global mean nutrient (0–100 m) concentrations
for nitrate (NO<inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mn mathvariant="normal">3</mml:mn></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>, phosphate (PO<inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>, silicic acid (SiO<inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> and
dissolved iron (dFe) are presented in Fig. 3. The magnitude of surface
nutrient concentrations differs substantially across the models (varying by a
factor of <inline-formula><mml:math display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 1.5–2, and by a factor of 5 for dissolved iron). The IPSL
models have relatively low surface nutrient concentrations. Compared to the
WOA09, two models overestimate phosphate (CESM1(BGC) and GFDL-ESM2G) and five
models overestimate nitrate. All of the models overestimate the silicic acid
observations, with the exception of CESM1(BGC). The CESM1(BGC) model
overestimates surface phosphate concentrations initially, due to excessive
nitrogen limitation, but then shows the strongest surface phosphate declines
over the 21st century (Fig. 4).</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F3"><caption><p>Time series of nitrate (NO<inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mn mathvariant="normal">3</mml:mn></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>, phosphate (PO<inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>, silicate
(SiO<inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mn mathvariant="normal">4</mml:mn></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> and dissolved iron (dFe) concentrations (0–100 m) are shown for
1850–2100. Red square indicates WOA2009 global mean values.</p></caption>
          <?xmltex \igopts{width=241.848425pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f03.pdf"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F4"><caption><p>Time series are displayed of mean changes (in percent) relative to
the 1990s for <bold>(a)</bold> NO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">3</mml:mn></mml:msub></mml:math></inline-formula>, <bold>(b)</bold> PO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">4</mml:mn></mml:msub></mml:math></inline-formula>,
<bold>(c)</bold> SiO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">4</mml:mn></mml:msub></mml:math></inline-formula> and <bold>(d)</bold> dFe (0–100 m) during 1850–2100.</p></caption>
          <?xmltex \igopts{width=241.848425pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f04.pdf"/>

        </fig>

      <p>Over the entire period from 1850–2100, the models all display decreasing
trends for surface nitrate, phosphate and silicic acid. Interestingly,
surface iron concentrations increase modestly in all but one of the models
by 4–10 %. Changes in iron concentrations impact marine productivity,
nitrogen fixation rates and oceanic net CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> uptake. In the CMIP5
simulations, iron inputs to the oceans from deposition and rivers are held
constant over time, so the increasing surface iron concentrations may reflect
increasing macronutrient limitation of phytoplankton growth, leading to
reduced biological uptake of iron. The reductions in the sinking export flux
also reduce the particle scavenging loss term for dissolved iron. In the
CESM1(BGC) model, increased production in the high-nutrient, low-chlorophyll
(HNLC) regions offset about 25 % of the reduction observed in the
macronutrient-limited areas with climate change, and changing circulation
patterns also altered the lateral transport of iron within the oceans (Moore
et al., 2013; Misumi et al., 2014).</p>
      <p>The relative changes in nutrient concentrations (0–100 m) (normalized to
1990s means) are presented in Fig. 4. The relative changes in the historical
run show a consistent pattern across the models for nitrate, phosphate and
dissolved iron (except for HadGEM2-ES). In the RCP8.5 projection, the models
show diverging estimates of the magnitude of the relative changes. For
nitrate, the reductions range between 3 and 14 %, whereas for phosphate
the reductions range between 3 and 20 %. Silicic acid and iron trends are
even more variable than for nitrate and phosphate. For silicic acid, three models
exhibit slight increases, while the others exhibit decreases ranging from
5–17 %. The variability in relative change in silicic acid concentration
in the RCP8.5 is likely associated with changes in plankton community and
variable diatom production (Bopp et al., 2005). All of the models include
some representation of diatoms (Table 2) but the match to observed silicic
acid concentrations for the current era is generally poor (Fig. 3).</p>
      <p>The spatial distributions of mean nitrate concentration for 0–100 m in the
1990s are shown in Fig. S3. The CMIP5 models reproduce key observed features
of the basin-scale distributions of surface nitrate. For example, all of the
models exhibit elevated nitrate concentrations in the eastern equatorial
Pacific, Southern Ocean, subarctic North Atlantic and subarctic Pacific. In
the subtropical gyres of the Atlantic and Pacific basins, mean nitrate
concentrations are low. These general patterns are consistent with the WOA09
observations. However, there are clear disagreements in some regions. For
example, the details of the high-nitrate surface water distributions vary
considerably in the eastern equatorial Pacific. The HNLC condition extends
too far north and south of the equator in some models, and too far to the
west in others (Fig. S3). The models also differ in the intensity and extent
of high nitrate concentration waters in the subarctic North Pacific, where
six of nine models show lower nitrate concentrations than the WOA09 data
(MPI-ESM-LR, MPI-ESM-MR and HadGEM2-ES are closest to the observations).
There are also differences in the Arabian Sea and Bay of Bengal, where most
models underestimate nitrate concentrations except the GFDL-ESM2M and
MPI-ESM-LR models.</p>
      <p>Intermodel spread in NPP during the 1990s is pronounced, with NPP as low as
29 PgC yr<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula> (IPSL-CM5A-LR and IPSL-CM5A-MR), while NPP in one model
exceeds 75 PgC yr<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula> (GFDL-ESM2M) (Table 3, Fig. 5). In addition, the
spatial pattern of NPP is not well represented by the multimodel mean (Bopp
et al., 2013). Satellite-based estimation of NPP is approximately
50 PgC yr<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula> (Behrenfeld et al., 2006; Carr et al., 2006). The
MPI-ES-MR and CESM1(BGC) models had NPP of 49.8 and 54.2 PgC yr<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>,
closer to the satellite-based estimates and the observationally constrained,
model estimate of 56 PgC yr<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula> by Buitenhuis et al. (2013). The
magnitude of EP also varies substantially across models in the 1990s, ranging
from 4.4 to 7.2 PgC yr<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula> (Table 3).</p>

<?xmltex \floatpos{p}?><table-wrap id="Ch1.T3" orientation="landscape"><caption><p>Global average of sea surface temperature (SST), sea surface
salinity (SSS), nitrate (<inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="normal">NO</mml:mi><mml:mn mathvariant="normal">3</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>), phosphate (<inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="normal">PO</mml:mi><mml:mn mathvariant="normal">4</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>), NPP,
EP, pe-ratio, stratification index (SI) defined as
density difference between 200 m and the surface
and  NPP by diatom (%) for the 1990s and 2090s. Observed estimates for the 1990s are obtained from
WOA09 data for SST, SSS, nitrate and phosphate, from Carr et al. (2006) for NPP.</p></caption><oasis:table frame="topbot"><?xmltex \begin{scaleboxenv}{.98}[.98]?><oasis:tgroup cols="19">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="right"/>
     <oasis:colspec colnum="3" colname="col3" align="right" colsep="1"/>
     <oasis:colspec colnum="4" colname="col4" align="right"/>
     <oasis:colspec colnum="5" colname="col5" align="right" colsep="1"/>
     <oasis:colspec colnum="6" colname="col6" align="right"/>
     <oasis:colspec colnum="7" colname="col7" align="right" colsep="1"/>
     <oasis:colspec colnum="8" colname="col8" align="right"/>
     <oasis:colspec colnum="9" colname="col9" align="right" colsep="1"/>
     <oasis:colspec colnum="10" colname="col10" align="right"/>
     <oasis:colspec colnum="11" colname="col11" align="right" colsep="1"/>
     <oasis:colspec colnum="12" colname="col12" align="right"/>
     <oasis:colspec colnum="13" colname="col13" align="right" colsep="1"/>
     <oasis:colspec colnum="14" colname="col14" align="right"/>
     <oasis:colspec colnum="15" colname="col15" align="right" colsep="1"/>
     <oasis:colspec colnum="16" colname="col16" align="right"/>
     <oasis:colspec colnum="17" colname="col17" align="right" colsep="1"/>
     <oasis:colspec colnum="18" colname="col18" align="right"/>
     <oasis:colspec colnum="19" colname="col19" align="right"/>
     <oasis:thead>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry namest="col2" nameend="col3" align="center" colsep="1">SST </oasis:entry>  
         <oasis:entry namest="col4" nameend="col5" align="center" colsep="1">SSS </oasis:entry>  
         <oasis:entry namest="col6" nameend="col7" align="center" colsep="1">NO3(0–100 m) </oasis:entry>  
         <oasis:entry namest="col8" nameend="col9" align="center" colsep="1">PO4(0–100 m) </oasis:entry>  
         <oasis:entry namest="col10" nameend="col11" align="center" colsep="1">NPP </oasis:entry>  
         <oasis:entry namest="col12" nameend="col13" align="center" colsep="1">EP </oasis:entry>  
         <oasis:entry namest="col14" nameend="col15" align="center" colsep="1">pe-ratio </oasis:entry>  
         <oasis:entry namest="col16" nameend="col17" align="center" colsep="1">SI </oasis:entry>  
         <oasis:entry namest="col18" nameend="col19" align="center">%Diat </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry namest="col2" nameend="col3" align="center" colsep="1"><inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup><mml:mi mathvariant="normal">C</mml:mi></mml:mrow></mml:math></inline-formula></oasis:entry>  
         <oasis:entry namest="col4" nameend="col5" align="center" colsep="1">psu </oasis:entry>  
         <oasis:entry namest="col6" nameend="col7" align="center" colsep="1">mmol m<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry namest="col8" nameend="col9" align="center" colsep="1">mmol m<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry namest="col10" nameend="col11" align="center" colsep="1">PgC yr<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry namest="col12" nameend="col13" align="center" colsep="1">PgC yr<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry namest="col14" nameend="col15" align="center" colsep="1"><inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">%</mml:mi></mml:math></inline-formula></oasis:entry>  
         <oasis:entry namest="col16" nameend="col17" align="center" colsep="1">kg m<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula></oasis:entry>  
         <oasis:entry namest="col18" nameend="col19" align="center"><inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">%</mml:mi></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">1990s</oasis:entry>  
         <oasis:entry colname="col3">2090s</oasis:entry>  
         <oasis:entry colname="col4">1990s</oasis:entry>  
         <oasis:entry colname="col5">2090s</oasis:entry>  
         <oasis:entry colname="col6">1990s</oasis:entry>  
         <oasis:entry colname="col7">2090s</oasis:entry>  
         <oasis:entry colname="col8">1990s</oasis:entry>  
         <oasis:entry colname="col9">2090s</oasis:entry>  
         <oasis:entry colname="col10">1990s</oasis:entry>  
         <oasis:entry colname="col11">2090s</oasis:entry>  
         <oasis:entry colname="col12">1990s</oasis:entry>  
         <oasis:entry colname="col13">2090s</oasis:entry>  
         <oasis:entry colname="col14">1990s</oasis:entry>  
         <oasis:entry colname="col15">2090s</oasis:entry>  
         <oasis:entry colname="col16">1990s</oasis:entry>  
         <oasis:entry colname="col17">2090s</oasis:entry>  
         <oasis:entry colname="col18">1990s</oasis:entry>  
         <oasis:entry colname="col19">2090s</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>  
         <oasis:entry colname="col1">Observations</oasis:entry>  
         <oasis:entry colname="col2">18.3</oasis:entry>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4">34.57</oasis:entry>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6">6.73</oasis:entry>  
         <oasis:entry colname="col7"/>  
         <oasis:entry colname="col8">0.63</oasis:entry>  
         <oasis:entry colname="col9"/>  
         <oasis:entry colname="col10">50.0</oasis:entry>  
         <oasis:entry colname="col11"/>  
         <oasis:entry colname="col12"/>  
         <oasis:entry colname="col13"/>  
         <oasis:entry colname="col14"/>  
         <oasis:entry colname="col15"/>  
         <oasis:entry colname="col16">1.81</oasis:entry>  
         <oasis:entry colname="col17"/>  
         <oasis:entry colname="col18"/>  
         <oasis:entry colname="col19"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">GFDL-ESM2G</oasis:entry>  
         <oasis:entry colname="col2">18.5</oasis:entry>  
         <oasis:entry colname="col3">20.4</oasis:entry>  
         <oasis:entry colname="col4">34.06</oasis:entry>  
         <oasis:entry colname="col5">33.98</oasis:entry>  
         <oasis:entry colname="col6">6.65</oasis:entry>  
         <oasis:entry colname="col7">6.10</oasis:entry>  
         <oasis:entry colname="col8">0.66</oasis:entry>  
         <oasis:entry colname="col9">0.58</oasis:entry>  
         <oasis:entry colname="col10">57.8</oasis:entry>  
         <oasis:entry colname="col11">57.5</oasis:entry>  
         <oasis:entry colname="col12">4.40</oasis:entry>  
         <oasis:entry colname="col13">4.10</oasis:entry>  
         <oasis:entry colname="col14">7.60</oasis:entry>  
         <oasis:entry colname="col15">7.02</oasis:entry>  
         <oasis:entry colname="col16">2.35</oasis:entry>  
         <oasis:entry colname="col17">2.75</oasis:entry>  
         <oasis:entry colname="col18">10.7</oasis:entry>  
         <oasis:entry colname="col19">9.7</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">GFDL-ESM2M</oasis:entry>  
         <oasis:entry colname="col2">18.8</oasis:entry>  
         <oasis:entry colname="col3">20.6</oasis:entry>  
         <oasis:entry colname="col4">34.32</oasis:entry>  
         <oasis:entry colname="col5">34.24</oasis:entry>  
         <oasis:entry colname="col6">8.67</oasis:entry>  
         <oasis:entry colname="col7">8.22</oasis:entry>  
         <oasis:entry colname="col8">0.58</oasis:entry>  
         <oasis:entry colname="col9">0.55</oasis:entry>  
         <oasis:entry colname="col10">77.6</oasis:entry>  
         <oasis:entry colname="col11">78.1</oasis:entry>  
         <oasis:entry colname="col12">6.54</oasis:entry>  
         <oasis:entry colname="col13">6.06</oasis:entry>  
         <oasis:entry colname="col14">8.44</oasis:entry>  
         <oasis:entry colname="col15">7.77</oasis:entry>  
         <oasis:entry colname="col16">1.95</oasis:entry>  
         <oasis:entry colname="col17">2.31</oasis:entry>  
         <oasis:entry colname="col18">9.4</oasis:entry>  
         <oasis:entry colname="col19">8.8</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">MPI-ESM-LR</oasis:entry>  
         <oasis:entry colname="col2">18.3</oasis:entry>  
         <oasis:entry colname="col3">20.7</oasis:entry>  
         <oasis:entry colname="col4">34.38</oasis:entry>  
         <oasis:entry colname="col5">34.23</oasis:entry>  
         <oasis:entry colname="col6">7.20</oasis:entry>  
         <oasis:entry colname="col7">6.61</oasis:entry>  
         <oasis:entry colname="col8">0.57</oasis:entry>  
         <oasis:entry colname="col9">0.50</oasis:entry>  
         <oasis:entry colname="col10">45.7</oasis:entry>  
         <oasis:entry colname="col11">41.6</oasis:entry>  
         <oasis:entry colname="col12">7.23</oasis:entry>  
         <oasis:entry colname="col13">6.05</oasis:entry>  
         <oasis:entry colname="col14">15.84</oasis:entry>  
         <oasis:entry colname="col15">14.56</oasis:entry>  
         <oasis:entry colname="col16">1.88</oasis:entry>  
         <oasis:entry colname="col17">2.41</oasis:entry>  
         <oasis:entry colname="col18">78.7</oasis:entry>  
         <oasis:entry colname="col19">80.1</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">MPI-ESM-MR</oasis:entry>  
         <oasis:entry colname="col2">18.4</oasis:entry>  
         <oasis:entry colname="col3">20.9</oasis:entry>  
         <oasis:entry colname="col4">34.41</oasis:entry>  
         <oasis:entry colname="col5">34.25</oasis:entry>  
         <oasis:entry colname="col6">6.96</oasis:entry>  
         <oasis:entry colname="col7">6.45</oasis:entry>  
         <oasis:entry colname="col8">0.53</oasis:entry>  
         <oasis:entry colname="col9">0.47</oasis:entry>  
         <oasis:entry colname="col10">47.9</oasis:entry>  
         <oasis:entry colname="col11">43.0</oasis:entry>  
         <oasis:entry colname="col12">6.56</oasis:entry>  
         <oasis:entry colname="col13">5.67</oasis:entry>  
         <oasis:entry colname="col14">13.70</oasis:entry>  
         <oasis:entry colname="col15">13.20</oasis:entry>  
         <oasis:entry colname="col16">1.97</oasis:entry>  
         <oasis:entry colname="col17">2.50</oasis:entry>  
         <oasis:entry colname="col18">91.1</oasis:entry>  
         <oasis:entry colname="col19">92.2</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">IPSL-CM5A-LR</oasis:entry>  
         <oasis:entry colname="col2">17.7</oasis:entry>  
         <oasis:entry colname="col3">21.0</oasis:entry>  
         <oasis:entry colname="col4">34.52</oasis:entry>  
         <oasis:entry colname="col5">34.43</oasis:entry>  
         <oasis:entry colname="col6">5.62</oasis:entry>  
         <oasis:entry colname="col7">4.81</oasis:entry>  
         <oasis:entry colname="col8">0.43</oasis:entry>  
         <oasis:entry colname="col9">0.36</oasis:entry>  
         <oasis:entry colname="col10">28.9</oasis:entry>  
         <oasis:entry colname="col11">27.0</oasis:entry>  
         <oasis:entry colname="col12">5.96</oasis:entry>  
         <oasis:entry colname="col13">4.87</oasis:entry>  
         <oasis:entry colname="col14">20.61</oasis:entry>  
         <oasis:entry colname="col15">18.05</oasis:entry>  
         <oasis:entry colname="col16">2.05</oasis:entry>  
         <oasis:entry colname="col17">2.63</oasis:entry>  
         <oasis:entry colname="col18">23.1</oasis:entry>  
         <oasis:entry colname="col19">20.3</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">IPSL-CM5A-MR</oasis:entry>  
         <oasis:entry colname="col2">18.2</oasis:entry>  
         <oasis:entry colname="col3">21.5</oasis:entry>  
         <oasis:entry colname="col4">34.42</oasis:entry>  
         <oasis:entry colname="col5">34.33</oasis:entry>  
         <oasis:entry colname="col6">5.82</oasis:entry>  
         <oasis:entry colname="col7">4.99</oasis:entry>  
         <oasis:entry colname="col8">0.45</oasis:entry>  
         <oasis:entry colname="col9">0.38</oasis:entry>  
         <oasis:entry colname="col10">31.8</oasis:entry>  
         <oasis:entry colname="col11">29.3</oasis:entry>  
         <oasis:entry colname="col12">6.33</oasis:entry>  
         <oasis:entry colname="col13">5.28</oasis:entry>  
         <oasis:entry colname="col14">19.94</oasis:entry>  
         <oasis:entry colname="col15">17.99</oasis:entry>  
         <oasis:entry colname="col16">2.12</oasis:entry>  
         <oasis:entry colname="col17">2.75</oasis:entry>  
         <oasis:entry colname="col18">22.0</oasis:entry>  
         <oasis:entry colname="col19">19.7</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">HadGEM2-ES</oasis:entry>  
         <oasis:entry colname="col2">18.3</oasis:entry>  
         <oasis:entry colname="col3">21.5</oasis:entry>  
         <oasis:entry colname="col4">34.06</oasis:entry>  
         <oasis:entry colname="col5">33.83</oasis:entry>  
         <oasis:entry colname="col6">6.56</oasis:entry>  
         <oasis:entry colname="col7">5.82</oasis:entry>  
         <oasis:entry colname="col8">0.44</oasis:entry>  
         <oasis:entry colname="col9">0.36</oasis:entry>  
         <oasis:entry colname="col10">34.5</oasis:entry>  
         <oasis:entry colname="col11">29.7</oasis:entry>  
         <oasis:entry colname="col12">4.77</oasis:entry>  
         <oasis:entry colname="col13">4.10</oasis:entry>  
         <oasis:entry colname="col14">13.82</oasis:entry>  
         <oasis:entry colname="col15">13.79</oasis:entry>  
         <oasis:entry colname="col16">2.45</oasis:entry>  
         <oasis:entry colname="col17">3.18</oasis:entry>  
         <oasis:entry colname="col18">58.8</oasis:entry>  
         <oasis:entry colname="col19">58.3</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">CESM1(BGC)</oasis:entry>  
         <oasis:entry colname="col2">19.0</oasis:entry>  
         <oasis:entry colname="col3">21.4</oasis:entry>  
         <oasis:entry colname="col4">34.23</oasis:entry>  
         <oasis:entry colname="col5">34.18</oasis:entry>  
         <oasis:entry colname="col6">7.60</oasis:entry>  
         <oasis:entry colname="col7">6.56</oasis:entry>  
         <oasis:entry colname="col8">0.71</oasis:entry>  
         <oasis:entry colname="col9">0.54</oasis:entry>  
         <oasis:entry colname="col10">54.2</oasis:entry>  
         <oasis:entry colname="col11">52.1</oasis:entry>  
         <oasis:entry colname="col12">6.97</oasis:entry>  
         <oasis:entry colname="col13">6.26</oasis:entry>  
         <oasis:entry colname="col14">12.86</oasis:entry>  
         <oasis:entry colname="col15">12.03</oasis:entry>  
         <oasis:entry colname="col16">2.25</oasis:entry>  
         <oasis:entry colname="col17">2.63</oasis:entry>  
         <oasis:entry colname="col18">35.7</oasis:entry>  
         <oasis:entry colname="col19">33.2</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">NorESM1-ME</oasis:entry>  
         <oasis:entry colname="col2">18.1</oasis:entry>  
         <oasis:entry colname="col3">20.2</oasis:entry>  
         <oasis:entry colname="col4">34.34</oasis:entry>  
         <oasis:entry colname="col5">34.26</oasis:entry>  
         <oasis:entry colname="col6">7.01</oasis:entry>  
         <oasis:entry colname="col7">6.18</oasis:entry>  
         <oasis:entry colname="col8">0.60</oasis:entry>  
         <oasis:entry colname="col9">0.51</oasis:entry>  
         <oasis:entry colname="col10">38.6</oasis:entry>  
         <oasis:entry colname="col11">35.3</oasis:entry>  
         <oasis:entry colname="col12">6.81</oasis:entry>  
         <oasis:entry colname="col13">6.18</oasis:entry>  
         <oasis:entry colname="col14">17.64</oasis:entry>  
         <oasis:entry colname="col15">17.52</oasis:entry>  
         <oasis:entry colname="col16">1.74</oasis:entry>  
         <oasis:entry colname="col17">2.01</oasis:entry>  
         <oasis:entry colname="col18"/>  
         <oasis:entry colname="col19"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Model mean</oasis:entry>  
         <oasis:entry colname="col2">18.4</oasis:entry>  
         <oasis:entry colname="col3">20.9</oasis:entry>  
         <oasis:entry colname="col4">34.30</oasis:entry>  
         <oasis:entry colname="col5">34.19</oasis:entry>  
         <oasis:entry colname="col6">6.90</oasis:entry>  
         <oasis:entry colname="col7">6.19</oasis:entry>  
         <oasis:entry colname="col8">0.55</oasis:entry>  
         <oasis:entry colname="col9">0.47</oasis:entry>  
         <oasis:entry colname="col10">46.3</oasis:entry>  
         <oasis:entry colname="col11">43.7</oasis:entry>  
         <oasis:entry colname="col12">6.17</oasis:entry>  
         <oasis:entry colname="col13">5.39</oasis:entry>  
         <oasis:entry colname="col14">14.49</oasis:entry>  
         <oasis:entry colname="col15">13.55</oasis:entry>  
         <oasis:entry colname="col16">2.08</oasis:entry>  
         <oasis:entry colname="col17">2.57</oasis:entry>  
         <oasis:entry colname="col18">41.2</oasis:entry>  
         <oasis:entry colname="col19">40.3</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup><?xmltex \end{scaleboxenv}?></oasis:table></table-wrap>

</sec>
<sec id="Ch1.S3.SS3">
  <title>Climate change impacts on NPP and EP</title>
      <p>All of the models exhibit decreasing trends in global NPP and EP with climate
change, as shown in previous studies (Bopp et al., 2013; Dutkiewicz et al.,
2013) and most models show more rapid decreases during the middle to latter
part of the 21st century (Figs. 5–6, Table 3). All nine models project
decreases in export production by the 2090s, exceeding 5 % relative to
levels in the 1990s, whereas the response for NPP is divided into two groups
after 2020. The CESM1(BGC) and GFDL models experience smaller changes in NPP
(&lt; 5 % relative to 1990s) while other models have larger
decreases (8–16 %). The largest relative change for NPP is about
<inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>16 % (MPI-ESM-LR). The EP decreases range from 7 % for GFDL-ESM2G to
28 % for IPSL-CM5A-LR. Cabré et al. (2015) found reductions in NPP
and EP for all biomes, except at the highest latitudes. The reductions in
global NPP and EP covary with the increases in stratification (Fig. 6). By
the 2090s, stratification increases by about 16 % in GFDL-ESM2M and up to
33 % in HadGEM1-ES. The rate of stratification increase is slower in the
two GFDL models and CESM1(BGC), which also agrees with the slower rates of
relative NPP and EP change.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F5"><caption><p>Time series of global mean net primary production, export
production and the particle export ratio over 1850–2100 are shown for each
model.</p></caption>
          <?xmltex \igopts{width=241.848425pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f05.pdf"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F6"><caption><p>Time series are displayed of the percent changes in net primary
production, export production and the particle export ratio and
stratification over the period 1850–2100 (each relative to their 1990s
means).</p></caption>
          <?xmltex \igopts{width=241.848425pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f06.pdf"/>

        </fig>

      <p>The variability across models in NPP is substantially larger than that seen
in EP (Table 3). The normalized standard deviation was <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula>27 % for
NPP, but only <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula>12 % for EP in the 1990s. The large spread in
simulated NPP and its response to climate change was also noted by
(Laufkötter et al., 2015). Seven of the nine models have an EP between
6.0 and 7.2 PgC yr<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula> in the 1990s, with the remaining two (HadGEM2-ES
and GFDL-ESM2G) having considerably lower EP (&lt; 5 PgC yr<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>. EP
is tightly coupled to new nutrient inputs to the euphotic zone in these
models. NPP is less tightly coupled as the fraction of regenerated production
varies across the models, and can vary spatially and temporally within some
models. Thus, the large spread in NPP is not just a function of the different
physical models and their transport of nutrients to the euphotic zone, but
rather it is also strongly impacted by the phytoplankton community structure
and export efficiency inherent in the models, and the resulting varying
levels of regenerated production.</p>
      <p>The sinking carbon flux out of the euphotic zone to net primary production
ratio (particle export ratio or pe-ratio) is a measure of the export
efficiency and also reflects the variable contribution of regenerated
production to total NPP (Dugdale and Goering, 1967; Eppley and Peterson,
1979; Dunne et al., 2007). High pe-ratio values are typically associated with
productive ecosystems dominated by larger phytoplankton (often diatoms;
Buesseler, 1998; Boyd and Newton, 1999), while low pe-ratios are associated
with oligotrophic food webs with most carbon flow through the microbial loop
(Pomeroy, 1974; Azam et al., 1983). The CMIP5 models that include both large
and small phytoplankton, assume a higher export efficiency for the large
phytoplankton (Aumont and Bopp, 2006; Seferian et al., 2013; Tjiputra et al.,
2013). The fraction of grazed material routed to sinking export is higher,
often by a factor of 3 or more, than the fraction routed to sinking export
for the small phytoplankton. Diatoms are also likely to dominate
phytoplankton blooms in these models. This can drive additional, very
efficient, export through aggregation, further enhancing the differences in
export efficiency between large and small phytoplankton. Relative to the
1990s, six of the nine models show decreasing trends in the pe-ratio (up to a
10 % reduction) (Figs. 5–6, Table 3; see also Cabré et al., 2015).
Diatoms account for a smaller percentage of NPP in the 2090s than in the
1990s in all the models, except for the MPI model, where nearly all of the
production is by diatoms and the smallest phytoplankton are not explicitly
represented (Table 3).</p><?xmltex \hack{\newpage}?>
</sec>
<sec id="Ch1.S3.SS4">
  <title>Increasing stratification and declining nutrients, NPP and EP</title>
      <p>We quantify the relations between stratification and key biogeochemical
variables with annual model output over the entire time period of 1850–2100.
This approach captures physical–biological interactions over interannual to
centennial timescales, and is thus more robust than the earlier work
comparing two end points from the beginning and end of the 21st century
(Bopp et al., 2013; Cabré et al., 2015). Changes in global NPP relative
to the 1990s are plotted against the relative change in stratification in
Fig. 7a. Across all the ESMs, a relatively good relationship exists with a
correlation <inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo></mml:mrow></mml:math></inline-formula> 0.72. Larger relative increases in stratification
correspond to larger relative declines in NPP. In addition, the
globally fitted line with a slope of 0.38 separates the models into two
groups. In one group (GFDL, IPSL and CESM1(BGC)), the NPP reductions are more
modest as stratification increases; the other group is composed of the two
MPI models, HadGEM1-ES and the NorESM model, which show more intense and
linear reductions in NPP with increasing stratification. The reduction of NPP
can be partly explained by nutrient changes responding to stratification
increases. Across the models, surface nitrate and phosphate concentrations
clearly decline as the stratification is enhanced (Fig. 7c and d, with
<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> of 0.80 and 0.82, respectively). Note that all of these trends are
robust across the full time series. Compared to the 1990s, the preindustrial
stratification is weaker, surface nutrient concentrations are higher, and NPP
and EP are elevated (Figs. 3–7).</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F7"><caption><p>Relationships are shown between the relative percent change in
surface stratification with climate and the relative change in several
biogeochemical variables including net primary production (NPP) <bold>(a)</bold>,
silicate <bold>(b)</bold>, nitrate <bold>(c)</bold>, phosphate <bold>(d)</bold>, export
production (EP) <bold>(e)</bold>, the fraction of NPP by diatoms
<bold>(f)</bold>. EP is plotted against the
change in the fraction of NPP by diatoms <bold>(g)</bold> and against the change
in NPP <bold>(h)</bold>. All changes are relative to the 1990s and plotted over
1850–2100. These time series are derived from global annual mean data.</p></caption>
          <?xmltex \igopts{width=241.848425pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f07.png"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F8" specific-use="star"><caption><p>The fraction of total NPP by diatoms for the 1990s is shown for each
model (data for NorESM not available).</p></caption>
          <?xmltex \igopts{width=369.885827pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f08.pdf"/>

        </fig>

      <p>This indicates significant climate change impacts on ocean biogeochemistry,
prior to the 1990s and the modern era when most ocean observations have been
made. The response of surface silicic acid to increasing stratification is
more variable. The projected changes are more divided, as three models
(MPI-ESM-LR, MPI-ESM-MR and HadGEM1-ES) show slight increases and the others
show reductions in surface silicic acid concentrations (Fig. 7b).</p>
      <p>EP is even more closely related to the stratification changes
(<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo></mml:mrow></mml:math></inline-formula> 0.89) than NPP (<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo></mml:mrow></mml:math></inline-formula> 0.72; Fig. 7e). The EP change is also
closely related to the NPP change (<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo></mml:mrow></mml:math></inline-formula> 0.85). EP decreases by up to
20 % (Fig. 7e), whereas NPP decreases by 10–18 %. The models display
two patterns in terms of the response of NPP and EP to climate change. The
first group includes five models (the two IPSL models, CESM1(BGC) and the two
GFDL models) where the relative declines in NPP are smaller than the relative
declines in EP by a factor of 2 or more (Fig. 6 and Table 3). In this group,
the EP drops by about 10 % and the NPP decreases by 5 %. In the
remaining models, the relative declines in EP and NPP are larger and more
similar in magnitude. For example, both EP and NPP decrease by about 14 %
in the HadGEM2-ES model. The differential declines in NPP and EP in the first
group of models documents declining export efficiency for the ocean
biological pump, driven by phytoplankton community shifts and a decreased
contribution to NPP by large phytoplankton (diatoms) (see below and
Figs. 6–10; also Cabré et al., 2015).</p>
      <p>Reduced nutrient availability is a major contributor to the declines in NPP
and EP. However, the relationship varies from one model to another because
growth and export are complicated functions of macronutrient limitation,
temperature, irradiance and iron limitation, as well as the routing of
organic matter within the ecosystem that drives export efficiency. Higher
metabolic rates with warming can be compensated to a large degree by changes
in the supply of nutrients in terms of globally integrated productivity
(Dutkiewicz et al., 2013). The NPP response is also strongly impacted by
phytoplankton community structure, which modifies export efficiency, and the
corresponding magnitude of regenerated primary production. For the IPSL,
CESM1(BGC) and GFDL models that show larger declines in EP than in NPP, this
pattern is driven by a decreasing contribution to total NPP by large
phytoplankton (Table 3, Figs. 8–9). Most of the primary production in these
models is by smaller phytoplankton. The GFDL models express this pattern most
strongly, with minimal declines in NPP, despite declines in EP approaching
10 % (Fig. 6 and Table 3). The other models tend to have production that
is dominated by diatoms, and do not capture the community shifts towards
increasing small phytoplankton dominance (and reduced export efficiency)
under increasing nutrient stress. The declines in NPP with increasing
stratification are more linear and more similar in magnitude to the declines
in EP (Fig. 7a, b and h). Thus, there are also very strong correlations
between the climate-driven changes in the fractional contribution of diatoms
to NPP and both the changes in stratification and the changes in EP (Fig. 7f
and g, correlations of <inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo></mml:mrow></mml:math></inline-formula> 0.85 and <inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo></mml:mrow></mml:math></inline-formula> 0.95, both much higher
than the correlation between changing stratification and NPP,
<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo></mml:mrow></mml:math></inline-formula> 0.71). Cabré et al. (2015) found similar patterns relating
community composition, NPP and EP comparing the period of 1980–1999 with
2080–2099, across low to midlatitude biomes.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F9" specific-use="star"><caption><p>The percent change in NPP by diatoms between the 2090s and the
1990s.</p></caption>
          <?xmltex \igopts{width=369.885827pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f09.pdf"/>

        </fig>

      <p>Some of these patterns are illustrated in Fig. 8, which shows the
contribution of diatoms (large phytoplankton) to NPP for the 1990s. Most of
the models show elevated diatom production at high latitudes and low diatom
contributions in the subtropical gyres. However, there are large
discrepancies in the magnitude of the diatom contribution, ranging from about
30 % to more than 90 % in the Arctic Ocean, for example. At the
global scale, diatoms account for only 9.4 % of NPP in the GFDL-ESM2M
model and reach a maximum of 91 % in the MPI-ESM-MR model (Table 3). The
large variability across the models reflects, in part, the lack of
observational data sets to constrain phytoplankton community composition at
the time these models were being developed. The new globally gridded ocean
atlas of plankton functional types, Marine Ecosystem Data (MAREDAT)
(Buitenhuis et al., 2013) has started to fill this gap, and should lead to
improved representations of plankton community structure in the future as the
data set becomes increasingly populated and is entrained into model
development and validation. Remote sensing estimates of phytoplankton
community composition and size class structures are also providing useful
constraints for global-scale modeling efforts (e.g., Alvain et al., 2005;
Hirata et al., 2008; Kostadinov et al., 2009; Siegel et al., 2014).</p>
      <p>The spatial patterns of the shifts in phytoplankton community composition
with climate change are illustrated in Figure 9; which shows the change in
the percentage of NPP by diatoms (2090s–1990s). There are some robust
trends across the models. One of the areas with the biggest declines in
diatom production is the high-latitude North Atlantic. This region typically
has some of the biggest stratification increases with climate change, greatly
reducing the deep winter mixing that entrains nutrients to the surface (Moore
et al., 2013; Randerson et al., 2015). Nearly all the models also show large
declines in diatom contributions to production in the Arctic Ocean. The CMIP5
models show consistent trends of increasing stratification, declining surface
nutrient concentrations and a longer growing season with climate change in
the Arctic (Vancoppenolle et al., 2013). Increasing surface temperatures and
substantial declines in sea ice cover allow for a longer growing season with
climate change. Thus, nutrients in surface waters are more completely used up
by summer's end, leading to community shifts with decreased diatom production
and an increased fraction of production by smaller phytoplankton. In the
CESM1(BGC) model, this community shift allows for a small increase in central
Arctic NPP, even as export production and surface nutrient concentrations
decline, due to the increased fraction of NPP from small phytoplankton and
the resulting increases in regenerated production (Moore et al., 2013).</p>
      <p>All of the models show some increase in the fraction of NPP by diatoms in the
Southern Ocean (Fig. 9). The increase is particularly strong in the
CESM1(BGC), IPSL and GFDL models. Most of the models also show some
increased diatom production in the tropical Pacific. Bopp et al. (2005)
reported decreasing diatom production in the Arctic and high-latitude North
Atlantic, with some increases in the Southern Ocean under a strong warming
climate scenario. Steinacher et al. (2010) also found declining productivity
in the North Atlantic and shifts in the export ratio due to phytoplankton
community shifts with decreasing diatom production. An earlier version of the
CESM used in that study (CCSM3) showed only small shifts in export ratios
with climate change, as the range in export ratios and the differences in
export efficiencies between large and small phytoplankton were smaller than
in CESM1(BGC) (Steinacher et al., 2010; Moore et al., 2013). Three models in
this study (HadGEM2-ES and the two MPI models) show increases in diatom
production in the low latitudes (Fig. 9). However, the diatoms dominate
production nearly everywhere in these three models (Fig. 8).</p>
      <p>There are also large intermodel differences in the spatial patterns of the
pe-ratio (Fig. 10). Some of the models (GFDL, IPSL, CESM1(BGC)) show a
close correlation between the pe-ratio and diatom production (compare Figs. 8 and 10),
due to the enhanced export efficiency for diatoms (large
phytoplankton) built into the models. Thus, there is a very high correlation
between the changing contribution of diatoms to NPP and the changes in EP
(Fig. 7g, Table 3). The MPI model includes one phytoplankton group
and has an essentially constant pe-ratio of 0.15, explaining the linearity of
the changes in NPP and EP with warming (Figs. 8 and 10). Production in the
HadGEM1-ES model is dominated nearly everywhere by the diatoms (Fig. 8).
Therefore, the MPI and HadGEM models cannot capture a shift towards
increasing small phytoplankton dominance under declining surface nutrient
concentrations. This leads to export production being closely correlated with
diatom production in these models as most production is by diatoms, as well
as in the other models where diatoms are assumed to export more efficiently
but account for a smaller fraction of total NPP (Table 3).</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F10" specific-use="star"><caption><p>The mean pe-ratio for the 1990s is shown for
each model.</p></caption>
          <?xmltex \igopts{width=369.885827pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f10.pdf"/>

        </fig>

      <p>There is also a strong correlation between the declines in the fraction of
NPP by diatoms and declines in the pe-ratio (compare Figs. 7, 9 and 11).
The largest declines in the pe-ratio are seen in the Arctic and the
high-latitude North Atlantic, regions where diatom production also decreases
the most. The GFDL, IPSL and CESM1(BGC) models show some reductions in
pe-ratio in the subarctic North Pacific, but the spatial patterns are
inconsistent (Fig. 11). The models display considerable variability in the
degree of stratification increase and in the dominant factor driving these
changes in the subarctic North Pacific (Figs. S2 and 2).</p>
      <p>The correlation for the relationship between the changing percentage of NPP
by diatoms vs. the changes in EP across all the models has an <inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> of
0.96 and a slope close to 1 (0.94, Fig. 7g), indicating that phytoplankton
community structure plays a dominant role in determining the responses of
NPP, EP and the pe-ratio to climate change. The biggest declines in the
fraction of production by diatoms and pe-ratios are in precisely the areas
where some of the largest increases in upper ocean stratification are seen,
along with declining surface nutrient concentrations, as in the Arctic Ocean
and in the high-latitude North Atlantic (Figs. 6–8; see also Steinacher et
al., 2010; Moore et al., 2013).</p>
</sec>
<sec id="Ch1.S3.SS5">
  <title>Projected changes in NPP, EP and stratification biases</title>
      <p>At the global scale, the CMIP5 models show considerable stratification biases
for the 1990s when compared to the WOA09 data (Fig. 1, Table 3). Only the
GFDL-ESM2M model is within 10 % of the observed value. From the density
profiles as well (Fig. S1), it is apparent that most of the models have
stronger stratification in the 1990s than seen in the observations. Liu et
al. (2014) argue that climate bias is important when projecting the impact of
climate change on land surface processes and Hoffman et al. (2014) document
this for atmospheric CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> mole fractions. Here, we examine how
stratification biases in the 1990s may affect model projections of NPP and EP
in the 2090s.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F11" specific-use="star"><caption><p>The percent change in the pe-ratio between
the 2090s and the 1990s).</p></caption>
          <?xmltex \igopts{width=369.885827pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f11.pdf"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F12"><caption><p>The stratification bias for the 1990s is plotted for each model
vs. the relative changes in NPP <bold>(a)</bold>, EP <bold>(b)</bold> and stratification <bold>(c)</bold> with
climate change (2090s–1990s). All three linear regressions are
statistically significant at a level &gt; 95 %.</p></caption>
          <?xmltex \igopts{width=227.622047pt}?><graphic xlink:href="https://bg.copernicus.org/articles/13/5151/2016/bg-13-5151-2016-f12.png"/>

        </fig>

      <p>Models with stronger bias in the 1990s for surface stratification tend to
predict larger climate-induced declines in both NPP and EP (Fig. 12,
<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo></mml:mrow></mml:math></inline-formula> 0.47 and <inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo></mml:mrow></mml:math></inline-formula> 0.54, respectively). The slopes are plotted when the
correlation is significant at &gt; 95 % level. Five of the models
have positive biases in stratification for the current era that exceed
20 %. These models also show the largest relative increases in
stratification with climate change of 26–30 % (Fig. 12, Table 3). The
remaining four models (GFDL models, CESM1(BGC) and NorESM1-ME) do a better
job of simulating observed stratification for the current era and predict
relative increases in stratification over the 21st century that are
roughly half as large, ranging <inline-formula><mml:math display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 15–18 %. This suggests that
the more biased models (for the 1990s) may be overestimating the projected
reductions in NPP and EP for the end of the century.</p>
</sec>
</sec>
<sec id="Ch1.S4" sec-type="conclusions">
  <title>Discussion and conclusions</title>
      <p>The ESMs analyzed here have different resolutions and incorporate marine
biogeochemical-ecosystem models with different mechanisms and degrees of
complexity. We find this set of models has consistent trends of increasing
stratification and decreasing NPP and EP. However, a large model spread is
apparent for the 1990s, particularly for NPP, and in the relative changes to
NPP and EP over the 21st century due to climate change. NPP is reduced
by 2–18 % in the 2090s and EP is reduced by 7–20 %. Mean
stratification increased by 16 % (GFDL-ESM2M) up to 33 % (HadGEM1-ES)
from the 1990s to the 2090s. Under strong warming scenarios like RCP8.5,
ocean stratification will continue to rapidly increase after the year 2100 in
all of these models (Randerson et al., 2015).</p>
      <p>The strongly linear relationship between stratification increases and EP
decreases seen within each model and across all the models (Figs. 7 and 12)
indicates a strong bottom-up control on EP, through declining upward nutrient
flux to the euphotic zone. Declining surface nutrient concentrations are seen
in all the models with climate change under the RCP8.5 scenario (Figs. 5–6).
Nitrate is reduced by 3–14 % and phosphate is reduced by 3–20 %.
Changes in surface silicic acid and iron concentrations are more
variable across the models. For silicic acid, there are three models showing
slight increases, while the others exhibit decreases of 5–17 %. With
respect to iron, eight models indicate an increase of 4–10 % relative to the
1990s, with the exception being the NorESM-ME model, which is reduced by
3 %. Changes in the temperature and light fields also have impacts on EP
in some regions, but increasing stratification and nutrient stress and the
resulting impacts on phytoplankton community composition and EP is the
dominant process at the global scale. On a global scale, over the full
1850–2100 time period, the changes in NPP and EP are more highly correlated
with the changes in stratification than with the changes in SST
(<inline-formula><mml:math display="inline"><mml:mrow><mml:msup><mml:mi>r</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo></mml:mrow></mml:math></inline-formula> 0.72 for stratification–NPP and
0.66 for SST–NPP, Fig. 7). This is because that the stratification metric
captures both the temperature-driven changes that dominate at low to
midlatitudes and the salinity-driven changes at higher latitudes. The
temperature-driven increases in growth rates are partially offset by reduced
nutrient supply in many regions as stratification increases (Bopp et al.,
2005; Cabré et al., 2015).</p>
      <p>Simulated NPP and its response to climate change are both more variable
across the models than EP, and are less strongly correlated with changes in
stratification (Fig. 7). This is driven by model differences in the export
efficiency of the biological pump and its relation to phytoplankton community
structure. The models that allow for shifts in phytoplankton community
structure show strongly nonlinear NPP response to climate change. NPP
declines less rapidly than EP with increasing nutrient stress, as the
percentage of NPP by large cells declines and export efficiency decreases
(and the regenerated production fraction increases). Models without this
dynamic community composition and export efficiency show a much more linear
NPP response to climate change (Fig. 7). Thus, projections of the response
of NPP to climate change in the CMIP5 models are critically dependent on the
simulated phytoplankton community structure, the efficiency of the biological
pump and the resulting (highly variable) levels of regenerated production.</p>
      <p>Spatial patterns of diatom productivity are influenced by changes in surface
nutrients and the resulting shifts in plankton community composition. The
response of the %NPP by diatoms depends on several factors, including
whether they were a small or large component of the community initially.
Therefore, the spatial patterns of changes in stratification and %NPP by
diatoms can differ (Figs. 2 and 9). The largest decreases are seen in areas
with high diatom production initially and large increases in stratification,
particularly in the Northern Hemisphere, leading to a north–south
hemispheric asymmetry (Marinov et al., 2013; Cabré et al., 2015). In the
Southern Ocean, the winds that drive upwelling, strengthen and shift poleward
with climate change, influencing iron supply and productivity patterns (Moore
et al., 2013; Misumi et al., 2014; Leung et al., 2015).</p>
      <p>The large spread in the simulated NPP rates for the 1990s and the variability
seen across models in the response of NPP to climate change introduce
challenges for climate impact and risk assessment, as NPP is a key product of
both terrestrial and marine ecosystem models, and changes to NPP are perhaps
the most cited result from this class of models. We have demonstrated that
the wide spread seen in simulated NPP is not due to the different physical
circulation models and the flux of nutrients they deliver to surface waters,
but rather to the efficiency of the biological pump (tied to community
structure in most models) and the resulting levels of regenerated primary
production. Changes in EP are an additional useful metric of climate impacts
on marine ecosystems. EP is more strongly tied to the climate feedback, as it
is mainly the fixed carbon sequestered to the deeper ocean by the biological
pump that will impact air–sea CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> exchange. In addition, in terms of
impacts up the food chain, EP may be a better metric than NPP. Friedland et
al. (2012) demonstrated that there is no correlation between fishery yield
and NPP at the global scale, but that there are strong correlations between
fishery yield and several other variables including chlorophyll
concentration, the pe-ratio and EP. These three proxies all correlate with
the fraction of primary production by large phytoplankton. In this context,
the results presented here suggest large future declines in fishery yield
across the high-latitude North Atlantic.</p>
      <p>Laufkötter et al. (2015) suggest a strong impact of temperature
modification of phytoplankton growth rates and other ecosystem processes
(including zooplankton growth and grazing rates) to infer a strong top-down
grazing influence on the NPP response to climate change, noting that
phytoplankton growth rates appear to increase at low latitudes in some
models, even as available nutrient concentrations decline. However, many of
the key fluxes and fields needed to support this hypothesis were not
available in the archived output from the CMIP5 models. This study relied on
estimated nutrient limitation factors and growth rates for only the surface
ocean in their analysis. Temperature warming is strongest at the surface
(Fig. 1S). Thus, the analysis may overestimate the temperature impacts for
the whole euphotic zone. Their conclusions were based on diatom-specific
nutrient limitation patterns, on the phytoplankton group with the largest
changes in limitation factors and on comparing total grazing with total NPP
for some models (Figs. 6–8, Laufkötter et al., 2015). These may not be
representative of the community nutrient limitation patterns and growth
response. At low latitudes the diatoms might show the biggest declines in
growth due to nutrient limitation, but they are only a small component of the
community in many of the models (Fig. 8). Under increasing nutrient stress,
phytoplankton community growth rates may increase simply due to a declining
contribution from diatoms, as the smaller phytoplankton will typically grow
faster at low nutrient concentrations. Grazing rates are also higher on the
small phytoplankton in models with multiple groups. Thus, comparing total
grazing rates to NPP cannot account for these influences of phytoplankton
community shifts. A community shift towards smaller phytoplankton will likely
increase mean community growth rate and total grazing, even with no change in
temperature.</p>
      <p>We agree that temperature effects may be important in the NPP climate change
response, and that the temperature influence on phytoplankton growth rates
and on the ecosystem processing of NPP that leads to export are highly
uncertain (Laufkötter et al., 2015).  Sherman et al. (2016) compiled in
situ estimates of phytoplankton community growth rates at the global scale
and found a relatively weak apparent temperature effect (apparent <inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi>Q</mml:mi><mml:mn>10</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>
<inline-formula><mml:math display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 1.5). These observational estimates of phytoplankton community growth
rates were compared with the CESM and GFDL CMIP5 simulations analyzed here.
ESMs used in climate change studies need to ensure that the emergent,
community temperature–growth relation matches this observed value (even
though higher explicit <inline-formula><mml:math display="inline"><mml:mrow><mml:msub><mml:mi>Q</mml:mi><mml:mn>10</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> values may be prescribed for individual plankton
functional types) to avoid biases in the response to temperature change
(Sherman et al., 2016).</p>
      <p>Some of the CMIP5 models have an assumed higher export efficiency for diatoms
relative to small phytoplankton, building on a long-standing paradigm,
strengthened by results from the detailed ecosystem studies of the Joint
Global Flux Study (JGOFS) program (Buesseler, 1998; Boyd and Newton, 1999).
In the current models, the spectrum of phytoplankton size structure is often
represented very simply with only the end members of one large and one small
phytoplankton group. Thus, the diatom group is a proxy for larger,
efficiently exporting, blooming phytoplankton functional types. DOM cycling,
heterotrophic bacteria, microzooplankton and the microbial loop are
typically treated in an idealized, implicit manner in the current models as
well.</p>
      <p>To accurately predict the response of NPP and EP to climate change, it may be
necessary to develop more robust ecosystem models with additional explicit
phytoplankton, heterotrophic microbial and zooplankton groups, including
their impacts on nutrient cycling, export efficiency and the downward
transport of organic matter. Models that include much greater diversity in
the phytoplankton show large community composition shifts with climate
change  (Dutkiewicz et al., 2013). Quantifying the links between NPP, EP and
community composition in observational data sets is a high priority. There are
only limited field observations of the pe-ratio, some of which rely on
nutrient drawdown and other indirect estimates of the sinking particle flux
(Dunne et al., 2007). Further progress to improve model performance
requires combined efforts from satellite, field and laboratory observations,
empirical and inverse modeling approaches, as well as process-based, forward
models.</p>
      <p>The large model spread in EP and NPP, and significant biases seen in key
nutrient fields for the 1990s suggest that the current ocean biogeochemical
models are far from perfect and their results must be interpreted with
caution. However, the relationships between stratification and EP, NPP and
nutrients do reveal some common mechanisms driving the climate change
response. The large intermodel differences for the current era in NPP, EP
and nutrient concentrations are partially associated with how these
biogeochemical models are initialized and spun up for these experiments. The
ocean biogeochemical models are often integrated in an offline mode for a
thousand years or more before coupling to other components of the ESM
(Séférian et al., 2016). The achieved preindustrial, near-steady
state of biogeochemical fields may deviate substantially from the observed
climatology, driven by biases in the physics and biogeochemistry. These
differences typically persist in the present-day simulations and future
projections. The advantage of the initialization and spin-up process is that
the biogeochemical fields are consistent with the simulated ocean
circulation, and will respond to climate-driven changes appropriately. The
strong intrinsic variability helps to reduce model drift and generate
reasonable longer-term variability. As a result, these long-term simulations
are suitable for analyzing climate trends, variability and sensitivities. RCP8.5
is a strong warming scenario and the relationship between stratification
changes and NPP/EP changes may be somewhat different under other RCP
scenarios. Although the relations between the degree of surface warming and
the ocean biogeochemical responses were largely linear across RCP4.5 and 8.5
for the CESM(BGC) (Moore et al., 2013), some potentially important marine
biogeochemical feedbacks on the climate
system were missing completely or not well represented in the CMIP5 models,
including important feedbacks through aerosol transport and deposition on
the marine iron cycle, feedbacks involving the oxygen minimum zones and the
marine nitrogen cycle, and the impacts on ocean biology by ongoing ocean
acidification. Each of these feedbacks could impact phytoplankton and
zooplankton community structures, NPP, EP and pe-ratios in the future.</p>
      <p>It is also important to consider the longer-term climate change responses of
both ocean physics and marine biogeochemistry. Moore et al. (2013) noted that
climate impacts on the oceans were still accelerating in the year 2100 under the
RCP8.5 scenario (but not under the more moderate RCP4.5 scenario).
Randerson et al. (2015) extended the CESM1(BGC) RCP8.5 scenario simulation
examined here to the year 2300. In these longer simulations, the climate
impacts on ocean physical fields and biogeochemistry led to even stronger
perturbations after 2100 than those presented here. In addition, the ocean
contribution to the climate-carbon feedback exceeded the land contribution
after the year 2100 (Randerson et al., 2015).</p>
</sec>
<sec id="Ch1.S5">
  <title>Data availability</title>
      <p>CMIP5 Data used in all figures can be obtained from <uri>ftp://ftp.ceda.ac.uk//badc/cmip5/data/cmip5/output1/</uri>.</p>
</sec>

      
      </body>
    <back><app-group>
        <supplementary-material position="anchor"><p><bold>The Supplement related to this article is available online at <inline-supplementary-material xlink:href="http://dx.doi.org/10.5194/bg-13-5151-2016-supplement" xlink:title="pdf">doi:10.5194/bg-13-5151-2016-supplement</inline-supplementary-material>.</bold></p></supplementary-material>
        </app-group><ack><title>Acknowledgements</title><p>We are grateful for support from the US Department of Energy Office of
Science and the National Science Foundation (NSF). This contribution was
supported by a grant to UCI as a part of the BGC Feedbacks Scientific Focus
Area within the Regional and Global Climate Modeling (RGCM) Program in the
Climate and Environmental Sciences Division (CESD) of the Biological and
Environmental Research (BER) Program in the US Department of Energy Office of
Science. We also received funding from the NSF project “Collaborative
Research: Improved Regional and Decadal Predictions of the Carbon Cycle”
(AGS-1048890). We would also like to thank all those in the CMIP5 project
efforts which made this work possible.<?xmltex \hack{\newline}?><?xmltex \hack{\newline}?> Edited
by: F. Chai <?xmltex \hack{\newline}?> Reviewed by: two anonymous referees</p></ack><ref-list>
    <title>References</title>

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(EP) regulated by increasing stratification and phytoplankton community
structure in the CMIP5 models</article-title-html>
<abstract-html><p class="p">We
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CMIP5 models show an increase in stratification in response to surface–ocean
warming and freshening, which is accompanied by decreases in surface
nutrients, NPP and EP.</p><p class="p">There is considerable variability across the models in the magnitudes of NPP,
EP, surface nutrient concentrations and their perturbations by climate
change. The negative response of NPP and EP to increasing stratification
reflects primarily a bottom-up control, as upward nutrient flux declines at
the global scale. Models with dynamic phytoplankton community structure show
larger declines in EP than in NPP. This pattern is driven by phytoplankton
community composition shifts, with reductions in productivity by large
phytoplankton as smaller phytoplankton (which export less efficiently) are
favored under the increasing nutrient stress. Thus, the projections of the
NPP response to climate change are critically dependent on the simulated
phytoplankton community structure, the efficiency of the biological pump and
the resulting levels of regenerated production, which vary widely across the
models. Community structure is represented simply in the CMIP5 models, and
should be expanded to better capture the spatial patterns and climate-driven
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