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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-15-6559-2018</article-id><title-group><article-title>Ecosystem carbon transit versus turnover times in response to climate
warming and rising atmospheric <inline-formula><mml:math id="M1" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> concentration</article-title><alt-title>Carbon transit versus turnover times</alt-title>
      </title-group><?xmltex \runningtitle{Carbon transit versus turnover times}?><?xmltex \runningauthor{X. Lu et al.}?>
      <contrib-group>
        <contrib contrib-type="author" corresp="yes" rid="aff1 aff2 aff3">
          <name><surname>Lu</surname><given-names>Xingjie</given-names></name>
          <email>xngj.lu@gmail.com</email>
        <ext-link>https://orcid.org/0000-0003-3732-1978</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff3">
          <name><surname>Wang</surname><given-names>Ying-Ping</given-names></name>
          
        <ext-link>https://orcid.org/0000-0002-4614-6203</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff2 aff4">
          <name><surname>Luo</surname><given-names>Yiqi</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff2">
          <name><surname>Jiang</surname><given-names>Lifen</given-names></name>
          
        </contrib>
        <aff id="aff1"><label>1</label><institution>School of Atmospheric Sciences, Sun Yat-sen University, Guangzhou
510275, China</institution>
        </aff>
        <aff id="aff2"><label>2</label><institution>Center for Ecosystem Science and Society, Department
of Biological Sciences, <?xmltex \hack{\newline}?>Northern Arizona University, Flagstaff
86011, USA</institution>
        </aff>
        <aff id="aff3"><label>3</label><institution>CSIRO Oceans and Atmosphere, Aspendale 3195, Australia</institution>
        </aff>
        <aff id="aff4"><label>4</label><institution>Department for Earth System Science, Tsinghua University, Beijing
100084, China</institution>
        </aff>
      </contrib-group>
      <author-notes><corresp id="corr1">Xingjie Lu (xngj.lu@gmail.com)</corresp></author-notes><pub-date><day>7</day><month>November</month><year>2018</year></pub-date>
      
      <volume>15</volume>
      <issue>21</issue>
      <fpage>6559</fpage><lpage>6572</lpage>
      <history>
        <date date-type="received"><day>7</day><month>April</month><year>2018</year></date>
           <date date-type="rev-request"><day>2</day><month>May</month><year>2018</year></date>
           <date date-type="rev-recd"><day>3</day><month>October</month><year>2018</year></date>
           <date date-type="accepted"><day>4</day><month>October</month><year>2018</year></date>
      </history>
      <permissions>
        
        
      <license license-type="open-access"><license-p>This work is licensed under the Creative Commons Attribution 4.0 International License. To view a copy of this licence, visit <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">https://creativecommons.org/licenses/by/4.0/</ext-link></license-p></license></permissions><self-uri xlink:href="https://bg.copernicus.org/articles/15/6559/2018/bg-15-6559-2018.html">This article is available from https://bg.copernicus.org/articles/15/6559/2018/bg-15-6559-2018.html</self-uri><self-uri xlink:href="https://bg.copernicus.org/articles/15/6559/2018/bg-15-6559-2018.pdf">The full text article is available as a PDF file from https://bg.copernicus.org/articles/15/6559/2018/bg-15-6559-2018.pdf</self-uri>
      <abstract>
    <p id="d1e141">Ecosystem carbon (C) transit time is a critical diagnostic parameter to
characterize land C sequestration. This parameter has different variants in
the literature, including a commonly used turnover time. However, we know
little about how different transit time and turnover time are in representing
carbon cycling through multiple compartments under a non-steady state. In this
study, we estimate both C turnover time as defined by the conventional
stock over flux and mean C transit time as defined by the mean age of C mass
leaving the system. We incorporate them into the Community
Atmosphere Biosphere Land Exchange (CABLE) model to estimate C turnover time
and transit time in response to climate warming and rising
atmospheric [<inline-formula><mml:math id="M2" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>]. Modelling analysis shows that both C turnover
time and transit time increase with climate warming but decrease with rising
atmospheric [<inline-formula><mml:math id="M3" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>]. Warming increases C turnover time by 2.4 years
and transit time by 11.8 years in 2100 relative to that at steady state in
1901. During the same period, rising atmospheric [<inline-formula><mml:math id="M4" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] decreases C
turnover time by 3.8 years and transit time by 5.5 years. Our analysis shows
that 65 % of the increase in global mean C transit time with climate
warming results from the depletion of fast-turnover C pool. The remaining
35 % increase results from accompanied changes in compartment C age
structures. Similarly, the decrease in mean C transit time with rising
atmospheric [<inline-formula><mml:math id="M5" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] results approximately equally from replenishment
of C into fast-turnover C pool and subsequent decrease in compartment C age
structure. Greatly different from the transit time, the turnover time, which
does not account for changes in either C age structure or composition of
respired C, underestimated impacts of warming and rising atmospheric
[<inline-formula><mml:math id="M6" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] on C diagnostic time and potentially led to deviations in
estimating land C sequestration in multi-compartmental ecosystems.</p>
  </abstract>
    </article-meta>
  </front>
<body>
      

<sec id="Ch1.S1" sec-type="intro">
  <title>Introduction</title>
      <p id="d1e206">The terrestrial ecosystem plays an important role in mitigation of climate change
through sequestering carbon (C) from the atmosphere. Terrestrial C storage is
co-determined by C input and C transit time, which is defined as the mean age
of C mass leaving the system (Luo et al., 2001; Taylor and Lloyd, 1992; Nir
and Lewis, 1975; Sierra et al., 2016; Manzoni et al., 2009; Eriksson, 1971;
Bolin and Rodhe, 1973). As transit time cannot be easily estimated from
observation, its variant, C turnover time, has been commonly used in the
literature (Sierra et al., 2016). A recent model inter-comparison study
indicated that a major cause of uncertainty in predicting future terrestrial
C sequestration is the variation in C turnover time among the models (Friend
et al., 2014). Up to 40 % of soil C sequestration potential can be
overestimated due to underestimation of C turnover time in current CMIP5
models (He et al., 2016). These examples highlight the importance of C
turnover time in understanding C cycle uncertainties. However, the C turnover
time has been mostly estimated with a conventional stock-over-flux method
(Carvalhais et al., 2014; Chen et al., 2013; Yan et al., 2017), which was
probably first introduced by Olson (1963) and based on<?pagebreak page6560?> a steady-state
assumption. In response to climate change, terrestrial ecosystem C dynamics
move away from steady states to dynamic disequilibrium (Luo and Weng, 2011).
Estimation of C turnover time likely deviates from C transit time in response
to climate change (Sierra et al., 2016). It is not clear how much the
estimate of C turnover time deviates from mean C transit time and what causes
their deviation under climate change.</p>
      <p id="d1e209">The C transit time as the mean age of C mass leaving the system can be
estimated only from age structure of C atoms in a multi-compartment
ecosystem. In contrast, the C turnover time is estimated without any
information of age structure of C atoms among compartments. Thus, C turnover
time is equivalent to mean C transit time only in the autonomous (i.e.
time-invariant) system at steady state (Sierra et al., 2016) with two
conditions to be satisfied. The first condition is that C fluxes and turnover
rates of individual pools do not change with time (i.e. time invariant or
autonomous). The second is that C influx to each pool equals the C efflux from
the pool (i.e. at steady state). However, the autonomous and steady-state
conditions are usually too strict to completely meet for real-world
ecosystems. For example, ecosystem C input via photosynthesis has diurnal
variation, a seasonal cycle, and inter-annual variability. C turnover time also
exhibits strong seasonal variation (Luo et al., 2017). With seasonal cycles
and inter-annual variability in both C input and turnover time, the ecosystem C
cycle is rarely at steady state and is rather mostly at dynamic disequilibrium
(Luo and Weng, 2011). Therefore, C turnover time may not equal C transit time
in the real world, especially when land C cycle is under transient dynamics
in response to climate change.</p>
      <p id="d1e212">The estimates of C transit time require information of C age structure in
ecosystems so that the mean age of the C atoms at a time when they leave the
system can be calculated (Manzoni et al., 2009). In a multi-compartmental
ecosystem, the C age within each compartment is represented by a single
compartment's mean C age and different compartments have different mean C
ages (Rasmussen et al., 2016). Thus, the C transit time is the weighed mean
of ages of C atoms leaving different compartments according to the fraction
of C loss from each pool to the total C loss. In response to rising
atmospheric [<inline-formula><mml:math id="M7" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>], increased C input of a young age into an
ecosystem is usually allocated more to fast- than slow-turnover pools,
leading to changes in the C age structure of the ecosystem. The fast-turnover
pools usually contribute more to respiratory loss than the slow pools. Thus,
it is expected that rising atmospheric [<inline-formula><mml:math id="M8" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] decreases C transit
time due to both changes in the C age structure and fractions of different
pools to total C loss from the ecosystem. Although it may change in response
to rising atmospheric [<inline-formula><mml:math id="M9" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] due to changes in both C fluxes and
pools, C turnover time does not account for changes in the C age structure
and the pool fractions of respiratory C loss over total ecosystem respiration. It is necessary to understand the theoretical deviation between C
transit time and C turnover time under a non-steady state.</p>
      <p id="d1e248">In this study, we aim to answer following questions:
<list list-type="order"><list-item>
      <p id="d1e253">How do both C turnover time and C transit time change in response to
climate warming and rising atmospheric [<inline-formula><mml:math id="M10" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>]?</p></list-item><list-item>
      <p id="d1e268">How much does the C
turnover time deviate from C transit time under future climate change?</p></list-item><list-item>
      <p id="d1e272">What mechanisms cause deviation between the two methods?</p></list-item><list-item>
      <p id="d1e276">Which
regions show the greatest deviations under different climate change
scenarios?</p></list-item></list>
To answer those questions, we incorporated a new algorithm into the Community
Atmosphere Biosphere Land surface Exchange (CABLE) model (Wang et al., 2010,
2011) to calculate both C turnover time and transit time. We ran the modified
CABLE model under three climate change scenarios, climate warming only, rising
atmospheric [<inline-formula><mml:math id="M11" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] only, and both climate warming and rising
atmospheric [<inline-formula><mml:math id="M12" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>], to compare changes in C transit time with those in
C turnover time.</p>
</sec>
<sec id="Ch1.S2">
  <title>Materials and methods</title>
<sec id="Ch1.S2.SS1">
  <title>The CABLE model</title>
      <p id="d1e313">CABLE is a global land surface model as described by Kowalczyk et al. (2006)
and incorporates global carbon, nitrogen, and phosphorus cycles (Wang et al.,
2010, 2011). This study does not activate phosphorus cycle in the model
largely because phosphorus has minor impacts on C cycle (Zhang et al., 2011).
Leaf photosynthesis, stomatal conductance, and heat and water transfer in
CABLE are calculated using the two-leaf approach (Wang and Leuning, 1998).</p>

<?xmltex \floatpos{t}?><table-wrap id="Ch1.T1" specific-use="star"><caption><p id="d1e319">Summary of scenarios and forcing data.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="4">
     <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:thead>
       <oasis:row>
         <oasis:entry colname="col1">Scenario name</oasis:entry>
         <oasis:entry colname="col2">Simulation</oasis:entry>
         <oasis:entry colname="col3">Climate forcing</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M20" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> data</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">abbreviation</oasis:entry>
         <oasis:entry colname="col3"/>
         <oasis:entry colname="col4"/>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>
         <oasis:entry colname="col1">Climate warming scenario</oasis:entry>
         <oasis:entry colname="col2">S1</oasis:entry>
         <oasis:entry colname="col3">Climate warming<inline-formula><mml:math id="M21" display="inline"><mml:msup><mml:mi/><mml:mi mathvariant="normal">a</mml:mi></mml:msup></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col4">Pre-industrial<inline-formula><mml:math id="M22" display="inline"><mml:msup><mml:mi/><mml:mi mathvariant="normal">d</mml:mi></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><inline-formula><mml:math id="M23" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> direct effect scenario</oasis:entry>
         <oasis:entry colname="col2">S2</oasis:entry>
         <oasis:entry colname="col3">Pre-industrial<inline-formula><mml:math id="M24" display="inline"><mml:msup><mml:mi/><mml:mi mathvariant="normal">b</mml:mi></mml:msup></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M25" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> increase<inline-formula><mml:math id="M26" display="inline"><mml:msup><mml:mi/><mml:mi mathvariant="normal">c</mml:mi></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Full effect scenario</oasis:entry>
         <oasis:entry colname="col2">S3</oasis:entry>
         <oasis:entry colname="col3">Climate warming<inline-formula><mml:math id="M27" display="inline"><mml:msup><mml:mi/><mml:mi mathvariant="normal">a</mml:mi></mml:msup></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M28" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> increase<inline-formula><mml:math id="M29" display="inline"><mml:msup><mml:mi/><mml:mi mathvariant="normal">c</mml:mi></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table><table-wrap-foot><p id="d1e322"><inline-formula><mml:math id="M13" display="inline"><mml:msup><mml:mi/><mml:mi mathvariant="normal">a</mml:mi></mml:msup></mml:math></inline-formula> Climate warming forcing data from 1901 to 2005 use
the
CRUNCEP dataset. The forcing data from 2006 to 2100 use CESM output under
Representative Concentration Pathways with radiative forcing increased by
8.5 W m<inline-formula><mml:math id="M14" 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> (RCP8.5).<?xmltex \hack{\newline}?>
<inline-formula><mml:math id="M15" display="inline"><mml:msup><mml:mi/><mml:mi mathvariant="normal">b</mml:mi></mml:msup></mml:math></inline-formula> The pre-industrial
climate forcing repeatedly uses 1-year climatology data averaged over 1901
to 1910 from the CRUNCEP dataset.<?xmltex \hack{\newline}?>
<inline-formula><mml:math id="M16" display="inline"><mml:msup><mml:mi/><mml:mi mathvariant="normal">c</mml:mi></mml:msup></mml:math></inline-formula> <inline-formula><mml:math id="M17" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>
concentration data are from the 200-year CMIP5 dataset under historical and
future scenarios (RCP8.5).<?xmltex \hack{\newline}?>
<inline-formula><mml:math id="M18" display="inline"><mml:msup><mml:mi/><mml:mi mathvariant="normal">d</mml:mi></mml:msup></mml:math></inline-formula> The pre-industrial
<inline-formula><mml:math id="M19" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> concentration is from the CMIP5 dataset for the year 1901.</p></table-wrap-foot></table-wrap>

      <p id="d1e581">Gross primary production (GPP) is calculated for both C<inline-formula><mml:math id="M30" display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">3</mml:mn></mml:msub></mml:math></inline-formula> and C<inline-formula><mml:math id="M31" display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">4</mml:mn></mml:msub></mml:math></inline-formula> plants
(Farquhar et al., 1980; Kowalczyk et al., 2006). The Farquhar model is a
biochemical model and modified in CABLE to calculate <inline-formula><mml:math id="M32" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>
assimilation rate at canopy level as a minimum of three potential limitation
processes of photosynthesis: light, enzyme, and C sink. Generally, all
three of these photosynthetic limitations are positively related to maximal
carboxylation rate (<inline-formula><mml:math id="M33" display="inline"><mml:mrow><mml:msub><mml:mi>V</mml:mi><mml:mi mathvariant="normal">cmax</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) or maximal potential electron
transport rate (<inline-formula><mml:math id="M34" display="inline"><mml:mrow><mml:msub><mml:mi>J</mml:mi><mml:mi mathvariant="normal">max</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) and intercellular <inline-formula><mml:math id="M35" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>
concentration (<inline-formula><mml:math id="M36" display="inline"><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>). Both <inline-formula><mml:math id="M37" display="inline"><mml:mrow><mml:msub><mml:mi>V</mml:mi><mml:mi mathvariant="normal">cmax</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula><mml:math id="M38" display="inline"><mml:mrow><mml:msub><mml:mi>J</mml:mi><mml:mi mathvariant="normal">max</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> are
temperature dependent (Leuning, 2002) and are maximized at around
30 <inline-formula><mml:math id="M39" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C. Thus, in response to warming, the model usually predicts a
positive response in GPP in cold and temperate regions but a negative
response in GPP in hot regions. <inline-formula><mml:math id="M40" display="inline"><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> depends on the stomata conductance
and atmospheric [<inline-formula><mml:math id="M41" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>]. GPP in CABLE positively responds to rising
atmospheric [<inline-formula><mml:math id="M42" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>]. CABLE photosynthesis is also controlled by soil
moisture.</p>
      <?pagebreak page6561?><p id="d1e723">Autotrophic respiration (<inline-formula><mml:math id="M43" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mi mathvariant="normal">a</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) in CABLE is also temperature
dependent and follows the modified Arrhenius formula (Ryan, 1991; Sitch et
al., 2003). At the canopy scale, <inline-formula><mml:math id="M44" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mi mathvariant="normal">a</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> is proportional to vegetation
nitrogen content and a temperature-related coefficient. <inline-formula><mml:math id="M45" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mi mathvariant="normal">a</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> will
positively respond to warming climate. Heterotrophic respiration
(<inline-formula><mml:math id="M46" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mi mathvariant="normal">H</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) is proportional to litter and soil decomposition rate and C
pool sizes. The decomposition rates in the model are controlled by soil
temperature and water. The temperature response is based on a <inline-formula><mml:math id="M47" display="inline"><mml:mrow><mml:msub><mml:mi>Q</mml:mi><mml:mn mathvariant="normal">10</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>
equation. Decomposition rates will positively respond to warming. The water
response function is from the daily time step ecosystem model (DAYCENT)
(Kelly et al., 2000) and the decomposition rate positively responds to wetter
soil conditions.</p>
      <p id="d1e782">CABLE has three vegetation compartments (leaf, wood, and root), three
litter compartments (metabolic litter, structure litter, and coarse wood
debris), and three soil compartments (fast soil pool, slow soil pool, and
passive soil pool) (Wang et al., 2010).</p>
</sec>
<sec id="Ch1.S2.SS2">
  <title>Simulation design</title>
      <p id="d1e791">We use the meteorological datasets from the National Centers for Environmental
Prediction and Climatic Research Unit (CRUNCEP) to drive our model. The
meteorological inputs from 1901 to 2100 include temperature, specific
humidity, air pressure, downward solar radiation, downward long-wave
radiation, rainfall, snowfall, and wind speed. The meteorological variables
of CRUNCEP data from 1901 to 2005 are interpolated from 6-hourly into
hourly (Qian et al., 2006) and re-gridded from
0.5<inline-formula><mml:math id="M48" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> <inline-formula><mml:math id="M49" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 0.5<inline-formula><mml:math id="M50" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> to
1.875<inline-formula><mml:math id="M51" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> <inline-formula><mml:math id="M52" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 2.5<inline-formula><mml:math id="M53" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> spatial resolution. From 2006 to 2100,
the hourly meteorological variables are generated from the Community Earth System
Model version 1.0 (CESM) (Li et al., 2016; Hurrell et al., 2013) for
Representative Concentration Pathway (RCP) 8.5.</p>
      <p id="d1e845">C storage for all three scenarios (climate warming, rising atmospheric
[<inline-formula><mml:math id="M54" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>], and both together) is initialized at pre-industrial steady
states, which is achieved by a spin-up approach. The spin-up method cycles
10-year CRUNCEP data (1901–1910) to drive CABLE, with [<inline-formula><mml:math id="M55" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] constant at the 1901 level. A semi-analytic solution is used to accelerate
spin-up simulation (Xia et al., 2012).</p>
      <p id="d1e870">The description of three scenarios in this study is summarized in Table 1.
Simulation one (S1) fixes the atmospheric [<inline-formula><mml:math id="M56" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] but uses changing
climate forcing. Simulation two (S2) fixes climate forcing but increases
atmospheric [<inline-formula><mml:math id="M57" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>]. Simulation three (S3) uses both changing climate
forcing and increasing atmospheric [<inline-formula><mml:math id="M58" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>].</p>
</sec>
<sec id="Ch1.S2.SS3">
  <title>Calculation of mean ecosystem C age</title>
      <p id="d1e912">Mean C age is defined as the mean time elapsed since C atoms (current in the
system) entered the system, which is important for understanding C transit
time described below. Following Rasmussen et al. (2016), mean C age
(<inline-formula><mml:math id="M59" display="inline"><mml:mover accent="true"><mml:mi>a</mml:mi><mml:mo mathvariant="normal">‾</mml:mo></mml:mover></mml:math></inline-formula>) can be formulated

                <disp-formula id="Ch1.E1" content-type="numbered"><mml:math id="M60" display="block"><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:mover accent="true"><mml:mi>a</mml:mi><mml:mo mathvariant="normal">‾</mml:mo></mml:mover><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced close=")" open="("><mml:mi>t</mml:mi></mml:mfenced><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow><mml:mrow><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:mfrac></mml:mstyle><mml:mo>.</mml:mo></mml:mrow></mml:math></disp-formula>

          In Eq. (1), <inline-formula><mml:math id="M61" display="inline"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> represents the mean age of C in the <inline-formula><mml:math id="M62" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th compartment, <inline-formula><mml:math id="M63" display="inline"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>
represents C pool size of the <inline-formula><mml:math id="M64" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th compartment, and <inline-formula><mml:math id="M65" display="inline"><mml:mi>d</mml:mi></mml:math></inline-formula> is the total number
of C compartments.</p>
      <?pagebreak page6562?><p id="d1e1049">Mixing fresh C input into old ecosystem C may reduce the mean ecosystem C
age. Meanwhile, C remaining in the system will age with time. As shown by
Rasmussen et al. (2016), dynamics of mean compartment C age can be described
by the following differential equation:

                <disp-formula specific-use="align" content-type="numbered"><mml:math id="M66" display="block"><mml:mtable displaystyle="true"><mml:mlabeledtr id="Ch1.E2"><mml:mtd/><mml:mtd><mml:mrow><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:mi mathvariant="normal">d</mml:mi><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow><mml:mrow><mml:mi mathvariant="normal">d</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:mfrac></mml:mstyle></mml:mrow></mml:mtd><mml:mtd><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:mtd></mml:mlabeledtr><mml:mtr><mml:mtd><mml:mstyle class="stylechange" displaystyle="true"/></mml:mtd><mml:mtd><mml:mrow><mml:mstyle class="stylechange" displaystyle="true"/><mml:mo>+</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:msubsup><mml:mo>∑</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:msubsup><mml:mfenced close=")" open="("><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:mfenced close=")" open="("><mml:mi>t</mml:mi></mml:mfenced><mml:mo>-</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced open="(" close=")"><mml:mi>t</mml:mi></mml:mfenced></mml:mrow></mml:mfenced><mml:msub><mml:mi>b</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:msub><mml:mi>k</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mo>-</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced open="(" close=")"><mml:mi>t</mml:mi></mml:mfenced><mml:msub><mml:mi>s</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:mfrac></mml:mstyle><mml:mo>.</mml:mo></mml:mrow></mml:mtd></mml:mtr></mml:mtable></mml:math></disp-formula>

            In Eq. (2), <inline-formula><mml:math id="M67" display="inline"><mml:mrow><mml:msub><mml:mi>s</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>(<inline-formula><mml:math id="M68" display="inline"><mml:mi>t</mml:mi></mml:math></inline-formula>) is the direct C input rate from net primary
production (NPP) to the <inline-formula><mml:math id="M69" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th compartment in g C m<inline-formula><mml:math id="M70" 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> yr<inline-formula><mml:math id="M71" 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>, <inline-formula><mml:math id="M72" display="inline"><mml:mrow><mml:msub><mml:mi>b</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>
is the proportion of decomposed carbon from the <inline-formula><mml:math id="M73" display="inline"><mml:mi>j</mml:mi></mml:math></inline-formula>th compartment to be
transferred to the <inline-formula><mml:math id="M74" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th compartment. <inline-formula><mml:math id="M75" display="inline"><mml:mrow><mml:msub><mml:mi>k</mml:mi><mml:mi>j</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> is the decomposition rate of
the <inline-formula><mml:math id="M76" display="inline"><mml:mi>j</mml:mi></mml:math></inline-formula>th compartment; the unit is per year. Thus, change in
compartment C age depends on C aging, network C transfers among pools with
different ages, and C input. Note that this equation works only for linear
models.</p>
      <p id="d1e1295">With a time step <inline-formula><mml:math id="M77" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:math></inline-formula>, the C transferred from the <inline-formula><mml:math id="M78" display="inline"><mml:mi>j</mml:mi></mml:math></inline-formula>th compartment
to the <inline-formula><mml:math id="M79" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th compartment (<inline-formula><mml:math id="M80" display="inline"><mml:mrow><mml:msub><mml:mi>F</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>) equals
<inline-formula><mml:math id="M81" display="inline"><mml:mrow><mml:msub><mml:mi>b</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:msub><mml:mi>k</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mi mathvariant="normal">Δ</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:math></inline-formula> and C input (<inline-formula><mml:math id="M82" display="inline"><mml:mrow><mml:msub><mml:mi>S</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) equals
<inline-formula><mml:math id="M83" display="inline"><mml:mrow><mml:msub><mml:mi>S</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>(t) <inline-formula><mml:math id="M84" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> s<inline-formula><mml:math id="M85" display="inline"><mml:msub><mml:mi/><mml:mi>i</mml:mi></mml:msub></mml:math></inline-formula>(t)<inline-formula><mml:math id="M86" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:math></inline-formula>, Eq. (2) can be rewritten in a
finite-element form to represent C age dynamics:

                <disp-formula id="Ch1.E3" content-type="numbered"><mml:math id="M87" display="block"><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mi mathvariant="normal">Δ</mml:mi><mml:mi>t</mml:mi><mml:mo>+</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:msubsup><mml:mo>∑</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:msubsup><mml:mfenced open="(" close=")"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:mfenced open="(" close=")"><mml:mi>t</mml:mi></mml:mfenced><mml:mo>-</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced open="(" close=")"><mml:mi>t</mml:mi></mml:mfenced></mml:mrow></mml:mfenced><mml:msub><mml:mi>F</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:mo>-</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced close=")" open="("><mml:mi>t</mml:mi></mml:mfenced><mml:msub><mml:mi>S</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:mfrac></mml:mstyle><mml:mo>.</mml:mo></mml:mrow></mml:math></disp-formula>

          In Eq. (3), the first term, <inline-formula><mml:math id="M88" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:math></inline-formula>, indicates natural C aging.
the second term,<inline-formula><mml:math id="M89" display="inline"><mml:mrow><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mstyle displaystyle="false"><mml:mfrac style="text"><mml:mrow><mml:msubsup><mml:mo>∑</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:msubsup><mml:mfenced open="(" close=")"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:mfenced close=")" open="("><mml:mi>t</mml:mi></mml:mfenced><mml:mo>-</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced open="(" close=")"><mml:mi>t</mml:mi></mml:mfenced></mml:mrow></mml:mfenced><mml:msub><mml:mi>F</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:mo>-</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced open="(" close=")"><mml:mi>t</mml:mi></mml:mfenced><mml:msub><mml:mi>S</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:mfrac></mml:mstyle></mml:mrow></mml:math></inline-formula>, represents the mean age change of the <inline-formula><mml:math id="M90" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th
compartment due to mixing with transferred C from other compartments or
external C input (i.e. NPP).</p>
      <p id="d1e1633">After the C cycle spin-up, we obtain the steady-state C ages in each
compartment by solving Eq. (2) with the Euler method. The changes of mean
compartment C age are less than 0.1 % between two successive cycles.</p>
</sec>
<sec id="Ch1.S2.SS4">
  <title>Ecosystem C transit time</title>
      <p id="d1e1642">C transit time is defined as the average time for a C atom to spend in the
ecosystem until its exit, or the time from entering the ecosystem to leaving
the ecosystem (or residence time; Luo et al., 2001). For a
multiple-compartment system, the mean C transit time,
<inline-formula><mml:math id="M91" display="inline"><mml:mrow><mml:msub><mml:mover accent="true"><mml:mi mathvariant="italic">τ</mml:mi><mml:mo mathvariant="normal">‾</mml:mo></mml:mover><mml:mi mathvariant="normal">ts</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>, can be calculated using the following equation
(Rasmussen et al., 2016):

                <disp-formula id="Ch1.E4" content-type="numbered"><mml:math id="M92" display="block"><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:msub><mml:mover accent="true"><mml:mi mathvariant="italic">τ</mml:mi><mml:mo mathvariant="normal">‾</mml:mo></mml:mover><mml:mi mathvariant="normal">ts</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced close=")" open="("><mml:mi>t</mml:mi></mml:mfenced><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mo>(</mml:mo><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>b</mml:mi><mml:mrow><mml:mi>j</mml:mi><mml:mi>i</mml:mi></mml:mrow></mml:msub><mml:mo>)</mml:mo><mml:msub><mml:mi>k</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow><mml:mrow><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mo>(</mml:mo><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>b</mml:mi><mml:mrow><mml:mi>j</mml:mi><mml:mi>i</mml:mi></mml:mrow></mml:msub><mml:mo>)</mml:mo><mml:msub><mml:mi>k</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:mfrac></mml:mstyle><mml:mo>.</mml:mo></mml:mrow></mml:math></disp-formula>

          When <inline-formula><mml:math id="M93" display="inline"><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mi>j</mml:mi></mml:mrow></mml:math></inline-formula>, <inline-formula><mml:math id="M94" display="inline"><mml:mrow><mml:msub><mml:mi>b</mml:mi><mml:mrow><mml:mi>j</mml:mi><mml:mi>i</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> equals <inline-formula><mml:math id="M95" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>1, indicating one unit of C exited from the
<inline-formula><mml:math id="M96" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th compartment. When <inline-formula><mml:math id="M97" display="inline"><mml:mrow><mml:mi>i</mml:mi><mml:mo>≠</mml:mo><mml:mi>j</mml:mi></mml:mrow></mml:math></inline-formula>, <inline-formula><mml:math id="M98" display="inline"><mml:mrow><mml:msub><mml:mi>b</mml:mi><mml:mrow><mml:mi>j</mml:mi><mml:mi>i</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> represents the proportion of
exited C of the <inline-formula><mml:math id="M99" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th compartment transferred to the <inline-formula><mml:math id="M100" display="inline"><mml:mi>j</mml:mi></mml:math></inline-formula>th compartment.
<inline-formula><mml:math id="M101" display="inline"><mml:mrow><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>b</mml:mi><mml:mrow><mml:mi>j</mml:mi><mml:mi>i</mml:mi></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn mathvariant="normal">0</mml:mn></mml:mrow></mml:math></inline-formula> when the exited C from the <inline-formula><mml:math id="M102" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th compartment
is fully transferred to all the other compartments, such as litterfall transferring from
plant to litter compartments, without C loss. <inline-formula><mml:math id="M103" display="inline"><mml:mrow><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>b</mml:mi><mml:mrow><mml:mi>j</mml:mi><mml:mi>i</mml:mi></mml:mrow></mml:msub><mml:mo>&lt;</mml:mo><mml:mn mathvariant="normal">0</mml:mn></mml:mrow></mml:math></inline-formula> when the exited C from the <inline-formula><mml:math id="M104" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th compartment is partly
transferred to the other compartments, such as litter or soil C
decomposition, with the rest lost to the atmosphere via respiration. The
denominator is the total amount of C loss from the ecosystem. The numerator
is the sum of products of respired C mass and C age.</p>
</sec>
<sec id="Ch1.S2.SS5">
  <title>Components of C transit time and their changes</title>
      <p id="d1e1977">Equation (4) can be reorganized as

                <disp-formula id="Ch1.E5" content-type="numbered"><mml:math id="M105" display="block"><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:msub><mml:mover accent="true"><mml:mi mathvariant="italic">τ</mml:mi><mml:mo mathvariant="normal">‾</mml:mo></mml:mover><mml:mi mathvariant="normal">ts</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced open="(" close=")"><mml:mi>t</mml:mi></mml:mfenced><mml:msub><mml:mi>f</mml:mi><mml:mrow><mml:mi mathvariant="normal">hr</mml:mi><mml:mo>,</mml:mo><mml:mi>i</mml:mi></mml:mrow></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mo>,</mml:mo></mml:mrow></mml:math></disp-formula>

          when we define the fraction of the total C loss from the <inline-formula><mml:math id="M106" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th compartment
(<inline-formula><mml:math id="M107" display="inline"><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mrow><mml:mi mathvariant="normal">hr</mml:mi><mml:mo>,</mml:mo><mml:mi>i</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>), hr represents heterotrophic respiration as

                <disp-formula id="Ch1.Ex2"><mml:math id="M108" display="block"><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:msub><mml:mi>f</mml:mi><mml:mrow><mml:mi mathvariant="normal">hr</mml:mi><mml:mo>,</mml:mo><mml:mi>i</mml:mi></mml:mrow></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mo>(</mml:mo><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>b</mml:mi><mml:mrow><mml:mi>j</mml:mi><mml:mi>i</mml:mi></mml:mrow></mml:msub><mml:mo>)</mml:mo><mml:msub><mml:mi>k</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow><mml:mrow><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo><mml:mo>(</mml:mo><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>b</mml:mi><mml:mrow><mml:mi>j</mml:mi><mml:mi>i</mml:mi></mml:mrow></mml:msub><mml:mo>)</mml:mo><mml:msub><mml:mi>k</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:mfrac></mml:mstyle><mml:mo>.</mml:mo></mml:mrow></mml:math></disp-formula>

          Equation (5) indicates that ecosystem C transit time consists of products of
two components: compartment C age (<inline-formula><mml:math id="M109" display="inline"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) and the fractional composition of
respired C (<inline-formula><mml:math id="M110" display="inline"><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mrow><mml:mi mathvariant="normal">hr</mml:mi><mml:mo>,</mml:mo><mml:mi>i</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>). Compartment C age as represented by Eq. (2)
changes due to C mixing with C in other compartments or external input.</p>
      <p id="d1e2230">According to Eq. (5), the change in ecosystem C transit time
<inline-formula><mml:math id="M111" display="inline"><mml:mrow><mml:msub><mml:mover accent="true"><mml:mi mathvariant="italic">τ</mml:mi><mml:mo mathvariant="normal">‾</mml:mo></mml:mover><mml:mi>R</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> can be attributed to the change in compartment C age
(change in C age structure) and the change in respired C composition as (see
the Supplement for details)

                <disp-formula specific-use="align" content-type="numbered"><mml:math id="M112" display="block"><mml:mtable displaystyle="true"><mml:mtr><mml:mtd><mml:mrow><mml:mstyle class="stylechange" displaystyle="true"/><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mover accent="true"><mml:mi mathvariant="italic">τ</mml:mi><mml:mo mathvariant="normal">‾</mml:mo></mml:mover><mml:mi mathvariant="normal">ts</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:mtd><mml:mtd><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:mo>=</mml:mo><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced close=")" open="("><mml:mi>t</mml:mi></mml:mfenced><mml:mi mathvariant="normal">Δ</mml:mi><mml:mfenced close=")" open="("><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mrow><mml:mi mathvariant="normal">hr</mml:mi><mml:mo>,</mml:mo><mml:mi>i</mml:mi></mml:mrow></mml:msub><mml:mfenced open="(" close=")"><mml:mi>t</mml:mi></mml:mfenced></mml:mrow></mml:mfenced><mml:mo>+</mml:mo><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>d</mml:mi></mml:munderover><mml:msub><mml:mi>f</mml:mi><mml:mrow><mml:mi mathvariant="normal">hr</mml:mi><mml:mo>,</mml:mo><mml:mi>i</mml:mi></mml:mrow></mml:msub><mml:mfenced close=")" open="("><mml:mi>t</mml:mi></mml:mfenced><mml:mi mathvariant="normal">Δ</mml:mi><mml:mfenced close=")" open="("><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced close=")" open="("><mml:mi>t</mml:mi></mml:mfenced></mml:mrow></mml:mfenced></mml:mrow></mml:mtd></mml:mtr><mml:mlabeledtr id="Ch1.E6"><mml:mtd/><mml:mtd><mml:mstyle displaystyle="true" class="stylechange"/></mml:mtd><mml:mtd><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:mo>+</mml:mo><mml:mspace linebreak="nobreak" width="0.25em"/><mml:mi>o</mml:mi><mml:mo>(</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced open="(" close=")"><mml:mi>t</mml:mi></mml:mfenced><mml:msub><mml:mi>f</mml:mi><mml:mrow><mml:mi mathvariant="normal">hr</mml:mi><mml:mo>,</mml:mo><mml:mi>i</mml:mi></mml:mrow></mml:msub><mml:mfenced open="(" close=")"><mml:mi>t</mml:mi></mml:mfenced><mml:mo>)</mml:mo><mml:mo>.</mml:mo></mml:mrow></mml:mtd></mml:mlabeledtr></mml:mtable></mml:math></disp-formula>

            The first term in Eq. (6) refers to C transit time change due to change in
respired C composition. If the fraction of respired C from fast-turnover pool
decreases, the mean ecosystem C transit time may increase because more
respired C comes from slow-turnover pools with older C ages. The second term
refers to C transit time change due to change in compartment C age structure.
Under elevated <inline-formula><mml:math id="M113" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, for example, young C enters a compartment
more than it leaves. C in the compartment becomes younger (i.e. young C
replenishment). Subsequently, mean ecosystem C transit time will decrease. The
third term refers to residuals that cannot be explained by the previous two
terms.</p>
      <p id="d1e2405">In this study, the C age dynamics and diagnostics represented by
Eqs. (1)–(6) are implemented into CABLE global simulations. During our
analysis of results, C age and transit time are averaged at different spatial
scales, i.e. grid cell scale, latitudinal scale, and global scale. Pool sizes
(<inline-formula><mml:math id="M114" display="inline"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>), pool ages (<inline-formula><mml:math id="M115" display="inline"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>), and pool-to-pool fluxes (<inline-formula><mml:math id="M116" display="inline"><mml:mrow><mml:msub><mml:mi>F</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>) in
Eqs. (1)–(6) uses the grid cell mean, latitudinal mean, and global mean of
each compartment to calculate three different scales of C age and transit
time. Therefore, the arithmetic average of C age or transit time at each grid
cell (e.g. Fig. 1a) does not equal the global average of C transit time
(e.g. Fig. 3a).</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F1" specific-use="star"><caption><p id="d1e2446">Global maps of <bold>(a)</bold> carbon transit time and
<bold>(b)</bold> mean carbon age are the average over 1901 to 1910 in each grid
cell.</p></caption>
          <?xmltex \igopts{width=483.69685pt}?><graphic xlink:href="https://bg.copernicus.org/articles/15/6559/2018/bg-15-6559-2018-f01.png"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F2"><caption><p id="d1e2464">Validation of simulated latitudinal variation pattern in ecosystem C
transit time. Comparison of the mean ecosystem C transit time from 1982 to
2005 as estimated in this study with the estimates from observation
(Carvalhais et al., 2014) and simulated C turnover time from CABLE. The grey
area indicates the uncertainty range of observation-based data. C transit
time theoretically equals C turnover time only at steady state. To ensure the
comparison is valid, we assumed, which is also assumed by some other
ecological studies (Trumbore, 2000), that ecosystem C cycle during the
data-covered period (1982 to 2005) is closed to the steady state. In
addition, the global C balance data also support our assumption. In the 1980s
and 1990s, global land C uptake from the Global Carbon Project (GCP) is about
0.8 GtC yr<inline-formula><mml:math id="M117" 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> with an uncertainty of 0.6 GtC yr<inline-formula><mml:math id="M118" 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>, which is
not significantly different from zero (Le Quéré et al., 2018).</p></caption>
          <?xmltex \igopts{width=170.716535pt}?><graphic xlink:href="https://bg.copernicus.org/articles/15/6559/2018/bg-15-6559-2018-f02.png"/>

        </fig>

</sec>
</sec>
<?pagebreak page6563?><sec id="Ch1.S3">
  <title>Results</title>
<sec id="Ch1.S3.SS1">
  <title>Global steady-state patterns of ecosystem C transit time</title>
      <p id="d1e2509">The global ecosystem C transit time at steady state generally shows a
latitudinal variation pattern (Fig. 1). The high values (greater than
70 years) are simulated not only in high-latitude regions, such as northern
Russia, northern Europe, and northern Canada but also in high-altitude
regions such as the Tibetan Plateau. Small values in C transit time (less than
30 years) are simulated in tropical rainforest, such as Amazon forest, Conga
forest, and Indonesian forest. Ecosystem C transit times in some grasslands
in middle-south Africa, south America, the southern Great Plains of the US, and
central northern Australia (savanna) are sometimes even smaller than those in
tropical forest. The spatial patterns of the mean ecosystem C age are quite
similar to the patterns of C transit time. However, the magnitude is
significantly higher than ecosystem C transit time. The mean ecosystem C age
ranges from 118 to 7952 years, whereas ecosystem C transit time ranges only
from 13 to 341 years.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F3"><caption><p id="d1e2514">CABLE simulates changes of global C transit time for each of the
three scenarios in <bold>(a)</bold> S1: climate warming scenario (red line), S2:
rising atmospheric [<inline-formula><mml:math id="M119" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario (green line), and S3: combined
climate warming and rising atmospheric [<inline-formula><mml:math id="M120" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario (blue line).
The changes in global ecosystem C transit time are separated into three
contributions based on Eq. (6): contribution from respired C composition
change, contribution from C age structure change, and residual
<bold>(b–d)</bold>.</p></caption>
          <?xmltex \igopts{width=241.848425pt}?><graphic xlink:href="https://bg.copernicus.org/articles/15/6559/2018/bg-15-6559-2018-f03.png"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F4" specific-use="star"><caption><p id="d1e2553">Global map of the change in C transit time in
<bold>(a)</bold> S1: climate warming scenario, <bold>(d)</bold> S2: rising
atmospheric [<inline-formula><mml:math id="M121" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario, and <bold>(g)</bold> S3: combined
climate warming and rising atmospheric [<inline-formula><mml:math id="M122" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario. In these
three scenarios, contribution from C age structure change and contribution
from respired C composition change are also estimated in relation to the
change in C transit time (S1: <bold>a</bold> and <bold>c</bold>; S2: <bold>e</bold> and
<bold>f</bold>; S3: <bold>h</bold> and <bold>i</bold>). The calculation of the contribution
from C age structure change and contribution from respired C contribution
change is based on Eq. (6). The positive contribution indicates the C age
structure change or composition change leads to C transit time change in
the same direction.</p></caption>
          <?xmltex \igopts{width=497.923228pt}?><graphic xlink:href="https://bg.copernicus.org/articles/15/6559/2018/bg-15-6559-2018-f04.png"/>

        </fig>

      <p id="d1e2613">The global latitudinal pattern of C transit time in 1982–2005 is consistent
with the observation-based pattern of turnover time (Fig. 2). The latter is
estimated at each grid cell globally using the stock-over-flux method to divide
ecosystem C storage by GPP (Carvalhais et al.,
2014). The magnitude of the estimate is mostly within the uncertainty range
of the observation-based pattern. We compare estimated C transit time in
1982–2005 with the turnover time, partly to match modelled values with
contemporary observations and partly due to the fact that terrestrial C
cycle is still approximately at a quasi-steady state between 1982 and 2005.
Over the 1980s and 1990s, the annual average of global net land carbon sink
estimated from the Global Carbon Project (GCP) is about 0.8 GtC yr<inline-formula><mml:math id="M123" 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> with
an uncertainty of 0.6 GtC yr<inline-formula><mml:math id="M124" 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>. As a reference, the annual average of
net land carbon sink in the most recent decade (2007–2016) is 2.3 GtC yr<inline-formula><mml:math id="M125" 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>
with an uncertainty of 0.7 GtC yr<inline-formula><mml:math id="M126" 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> (Le Quéré et al., 2018). The
net change of global land carbon in the 1980s and 1990s is not that significant,
which indicates the land C cycle has not moved too far away from the steady
state. Moreover, the simulated latitudinal pattern of C transit time almost
overlaps with C turnover time, showing that C cycle is still near
the<?pagebreak page6564?> steady state at present day. Annual C turnover time theoretically equals C transit time when the C cycle is close to steady state (Sierra et al.,
2016).</p>
</sec>
<sec id="Ch1.S3.SS2">
  <title>Responses of global mean C transit time to climate change</title>
      <p id="d1e2670">In the 200-year simulation, mean global ecosystem C transit time increases by
11.8 years in response to climate warming (S1) and decreases by 5.6 years in
response to rising atmospheric [<inline-formula><mml:math id="M127" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] (S2) (Fig. 3a). When climate
warming and rising atmospheric [<inline-formula><mml:math id="M128" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] force together (S3), C transit
time decreases by 1.6 years. The increase in C transit time in S1 is not
significant in the 20th century but substantial in the 21st century.
Oppositely, the decrease in C transit time in S2 is steady before 2060 but
slows down afterward. Mean C transit time in S3 decreases but with a smaller
magnitude than that for S2 in the 21st century.</p>
      <p id="d1e2695">Across all three scenarios, the majority (over 93.4 %) of changes in C
transit time can be explained by two combined changes in compartment C age
structure and respired C composition. Changes in the compartment C age
structure and the respired C composition both significantly contribute to the
total change in global C transit time. However, the contribution fraction
varies among the three scenarios at different times. In the climate warming
scenario (S1), respired C composition changes contribute about 70 % of
the increase in C transit time in the 21st century (Fig. 3b). In the rising
atmospheric [<inline-formula><mml:math id="M129" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario (S2), respired C composition change and
C age structure change contribute equally (Fig. 3c). When coupling climate
warming and rising atmospheric [<inline-formula><mml:math id="M130" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] together in S3, respired C
composition change significantly contributes only in the middle of the 200-year
simulation (around the year 2000) and a little at the end of the 21st century. The
contribution of C age structure change to the change in C transit time
gradually increases.</p>
      <p id="d1e2720">The increase in C transit time in the climate warming scenario (S1) is the most
significant from low-latitude regions in South America and Africa (Fig. 4a).
Respired C composition change explains most of these regional changes
(Fig. 4c). The decrease in C transit time in the rising atmospheric [<inline-formula><mml:math id="M131" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>]
scenario (S2) is evenly simulated all over the world (Fig. 4d). Respired C
composition change also plays an important role in most regions except for
North Africa with little vegetation coverage. The C transit time in the combined
climate warming and rising atmospheric [<inline-formula><mml:math id="M132" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario (S3) mostly
decreases in the Northern Hemisphere but increases in some tropical grassland
regions in South America and Africa (Fig. 4g). In those regions where C
transit time decreases, compartment C age structure change due to fresh C
replenishment explains most of the change in C transit time.</p>
      <p id="d1e2745">Note that the response under combined effects (S3) is not a sum of those from
individual effects (S1 plus S2). The nonadditive response to climate warming
and rising atmospheric [<inline-formula><mml:math id="M133" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] is probably due to their interactions,
which have been commonly found in many ecological studies (Norby and Luo,
2004; Luo et al., 2008; Leuzinger et al., 2011; Campbell et al., 1997; Zhang
et al., 2016).</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F5"><caption><p id="d1e2762"><bold>(a)</bold> Changes of global C turnover time (stock over flux) in S1: climate warming scenario (red line), S2: rising
atmospheric [<inline-formula><mml:math id="M134" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario (green line), and S3: combined
climate warming and rising atmospheric [<inline-formula><mml:math id="M135" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario (blue line).
<bold>(b)</bold> The deviation of the change in C turnover time (<inline-formula><mml:math id="M136" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">to</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) is estimated relative to the change in C transit time
(<inline-formula><mml:math id="M137" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">ts</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>): (<inline-formula><mml:math id="M138" display="inline"><mml:mrow><mml:mfenced close="|" open="|"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">to</mml:mi></mml:msub></mml:mrow></mml:mfenced><mml:mo>-</mml:mo><mml:mfenced close="|" open="|"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">ts</mml:mi></mml:msub></mml:mrow></mml:mfenced><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>. Positive indicates more
change in C turnover time than C transit time. The grey line represents the
reference of no deviation. <bold>(c)</bold> The relative deviation of the change
in C turnover time in the years 2000 and 2100 is also estimated relative to the
change in C transit time: <inline-formula><mml:math id="M139" display="inline"><mml:mrow><mml:mstyle displaystyle="false"><mml:mfrac style="text"><mml:mrow><mml:mo>(</mml:mo><mml:mfenced open="|" close="|"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">to</mml:mi></mml:msub></mml:mrow></mml:mfenced><mml:mo>-</mml:mo><mml:mfenced open="|" close="|"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">ts</mml:mi></mml:msub></mml:mrow></mml:mfenced><mml:mo>)</mml:mo></mml:mrow><mml:mrow><mml:mfenced close="|" open="|"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">ts</mml:mi></mml:msub></mml:mrow></mml:mfenced></mml:mrow></mml:mfrac></mml:mstyle><mml:mo>×</mml:mo><mml:mn mathvariant="normal">100</mml:mn></mml:mrow></mml:math></inline-formula> %.</p></caption>
          <?xmltex \igopts{width=241.848425pt}?><graphic xlink:href="https://bg.copernicus.org/articles/15/6559/2018/bg-15-6559-2018-f05.png"/>

        </fig>

</sec>
<sec id="Ch1.S3.SS3">
  <title>Global C turnover time and its deviation</title>
      <p id="d1e2909">Similar to the changes in C transit time, the global C turnover time
increases with climate warming and decreases with rising atmospheric
[<inline-formula><mml:math id="M140" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] (Fig. 5a). However, the magnitude substantially differs
between these two methods (Figs. 3a,<?pagebreak page6565?> 5a). In response to climate warming
(S1), global ecosystem C turnover time increases by only 2.4 years at the end of
the simulation, which is only one-fifth of the increase in C transit time
(11.8 years). In response to rising atmospheric [<inline-formula><mml:math id="M141" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] (S2), global
C turnover time decreases by 3.7 years, whereas C transit time decreases by
5.6 years. In response to the coupled scenario (S3) in which climate warming and
rising atmospheric [<inline-formula><mml:math id="M142" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] force together, global ecosystem C
turnover time decreases by 4.5 years, while C transit time decreases by only
1.6 years.</p>
      <p id="d1e2945">In 1901, the global C turnover time is about 0.5 year longer than the C
transit time (Figs. 3a, 5a). Theoretically, C turnover time equals transit
time when land C cycle is at steady state. The offset at the initial state of
simulations probably results from C seasonal cycles, which are not at steady
state. The underestimates of the change in C turnover time relative to C
transit time increase in the climate warming scenario (S1) by up to 9.4 years in
the end of the 21st century, which is 79.6 % of the total increase in C
transit time (Fig. 5b). In the rising atmospheric [<inline-formula><mml:math id="M143" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario (S2),
the deviation constantly grows to about 1.9 years, 27.7 % of the
underestimated decrease in C turnover time. In the climate warming and rising
atmospheric [<inline-formula><mml:math id="M144" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario (S3), the change in C turnover time is
overestimated by 2.9 years or 181.1 % relative to the change in C
transit time in 2100 (Fig. 5b, c).</p>

      <?xmltex \floatpos{p}?><fig id="Ch1.F6" specific-use="star"><caption><p id="d1e2972"><bold>(a)</bold> Latitudinal variation in C transit time
(<inline-formula><mml:math id="M145" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">transit</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) at steady state and <bold>(b, c)</bold> its change
are compared to <bold>(d–f)</bold> C turnover time (<inline-formula><mml:math id="M146" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">to</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>).
The changes between the 2090s and 1900s are estimated by <bold>(c, f)</bold> absolute value: <inline-formula><mml:math id="M147" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:mi mathvariant="italic">τ</mml:mi><mml:mo>=</mml:mo><mml:mo>(</mml:mo><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mrow><mml:mn mathvariant="normal">2090</mml:mn><mml:mi mathvariant="normal">s</mml:mi></mml:mrow></mml:msub><mml:mo>-</mml:mo><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mrow><mml:mn mathvariant="normal">1900</mml:mn><mml:mi mathvariant="normal">s</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>) and by <bold>(b, e)</bold> relative value: <inline-formula><mml:math id="M148" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">r</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mstyle displaystyle="false"><mml:mfrac style="text"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:mi mathvariant="italic">τ</mml:mi></mml:mrow><mml:mrow><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mrow><mml:mn mathvariant="normal">1900</mml:mn><mml:mi mathvariant="normal">s</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:mfrac></mml:mstyle></mml:mrow></mml:math></inline-formula>. <bold>(g)</bold> The deviation of C turnover time relative
to C transit time is estimated by <inline-formula><mml:math id="M149" display="inline"><mml:mrow><mml:mo>(</mml:mo><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">to</mml:mi></mml:msub><mml:mo>-</mml:mo><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">ts</mml:mi></mml:msub><mml:mo>)</mml:mo><mml:mo>/</mml:mo><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">ts</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> at steady state. Relative to C
transit time, the deviation of the change in C turnover time is estimated by
<bold>(h)</bold> absolute deviation (<inline-formula><mml:math id="M150" display="inline"><mml:mrow><mml:mfenced open="|" close="|"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">to</mml:mi></mml:msub></mml:mrow></mml:mfenced><mml:mo>-</mml:mo><mml:mfenced close="|" open="|"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">ts</mml:mi></mml:msub></mml:mrow></mml:mfenced><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> and
<bold>(i)</bold> relative deviation in <inline-formula><mml:math id="M151" display="inline"><mml:mstyle displaystyle="false"><mml:mfrac style="text"><mml:mrow><mml:mo>(</mml:mo><mml:mfenced close="|" open="|"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">to</mml:mi></mml:msub></mml:mrow></mml:mfenced><mml:mo>-</mml:mo><mml:mfenced close="|" open="|"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">ts</mml:mi></mml:msub></mml:mrow></mml:mfenced><mml:mo>)</mml:mo></mml:mrow><mml:mrow><mml:mfenced open="|" close="|"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi mathvariant="italic">τ</mml:mi><mml:mi mathvariant="normal">ts</mml:mi></mml:msub></mml:mrow></mml:mfenced></mml:mrow></mml:mfrac></mml:mstyle></mml:math></inline-formula>. All variables are
compared in S1: only climate warming scenario (red line),
S2: rising atmospheric [<inline-formula><mml:math id="M152" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario (green line), and S3:
combined climate warming and rising atmospheric [<inline-formula><mml:math id="M153" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>]
scenario (blue line). Grey lines in <bold>(b)</bold>, <bold>(c)</bold>, <bold>(e)</bold>,
and <bold>(f)</bold> represent the reference lines of no change and those
in <bold>(h)</bold> and <bold>(i)</bold> represent the reference line of no deviation.</p></caption>
          <?xmltex \igopts{width=497.923228pt}?><graphic xlink:href="https://bg.copernicus.org/articles/15/6559/2018/bg-15-6559-2018-f06.png"/>

        </fig>

</sec>
<sec id="Ch1.S3.SS4">
  <title>Latitudinal variation in C turnover time and its deviation</title>
      <p id="d1e3237">Latitudinal patterns in C transit time and C turnover time at the initial
state in 1900 are nearly the same. Steady-state estimates are from
20 years in low latitudes and 100 years in high latitudes (Fig. 6a, d). However,
significant deviation still exists in high latitudes (north of 60<inline-formula><mml:math id="M154" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> N
and south of 50<inline-formula><mml:math id="M155" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> S) (Fig. 6g) because seasonal soil freeze–thaw
processes in this region lead to a strong seasonal cycle of soil
decomposition and violate the steady-state assumption of the C turnover time.
The underestimates of C turnover time can be up to 10 years in high-latitude
regions, which is about 8 % of C transit time. In other areas, deviation
of turnover time is less than 0.5 years.</p>
      <p id="d1e3258">Changes in C turnover time and C transit time deviate in different regions in
response to climate warming (S1) (Fig. 6b, c, e, f). In temperate and
tropical regions, C<?pagebreak page6566?> transit time significantly increases, while C turnover
time also increases but at a much smaller magnitude. In the tropics, C transit
time increases by 13 years in 2100, up to 60 % of the initial value in
1900, whereas C turnover time increases by only 2 years. In the high-latitude
region, C transit time slightly decreases (Fig. 6b and c) but C turnover time
significantly decreases by several decades in the high latitudes (Fig. 6f). In
some regions between 40 and 60<inline-formula><mml:math id="M156" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> N, C transit time increases but
turnover time decreases in response to climate warming. C turnover time
overall changes less than C transit time in the S1 scenario. Warming-induced
changes in C turnover time are underestimated by 5 % at the high latitudes
of the Southern Hemisphere and up to 50 % at the low latitudes (Fig. 6h), which
ranges from 2 to 29 years (Fig. 6i).</p>
      <p id="d1e3270">In response to rising atmospheric [<inline-formula><mml:math id="M157" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] (S2), both C turnover time
and transit time decrease. The magnitude of changes for both of them is
generally greater at the mid-latitudes than at either low or high
latitudes (Fig. 6b, e). At most latitudes, C turnover time decreases less
than C transit time, leading to a positive deviation (Fig. 6h, i). The
deviation of the change is higher in low than high latitudes. In response
to rising atmospheric [<inline-formula><mml:math id="M158" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>], the underestimate of the decrease in C
turnover time is at most 2 years in absolute deviation or 10 % in
relative deviation (Fig. 6h, i).</p>
      <p id="d1e3295">In the climate warming and rising atmospheric [<inline-formula><mml:math id="M159" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario (S3), C
turnover time and C transit time decrease at most of the latitudinal regions
except for some tropic areas (Fig. b, 6c, e, f). The decrease in C turnover
time is more than that in C transit time (Fig. 6h, i). Especially in high
latitudes, the difference in changes is much more significant. C turnover
time is reduced by up to 3 decades (Fig. 6f) or 35 % (Fig. 6e),
whereas C transit time shows nearly no relative changes. Deviation
in these areas can be up to 27 years (Fig. 6i).</p>
</sec>
</sec>
<sec id="Ch1.S4">
  <title>Discussion</title>
<sec id="Ch1.S4.SS1">
  <title>C transit time and its two components</title>
      <p id="d1e3321">Changes in C transit time can be explained by its two components: the
respired C composition and compartment C age structure. The first component
is to account for different contributions of respired C from different pools
to total ecosystem C loss. Previous studies have demonstrated that pathways
of respiring C from multiple compartments are variably controlled by the global
change factor (Luo et al., 2001). Results from this study provide more
spatial details about where C transit time changes due to respired C
composition change. For example, over 80 % of the increase in C transit
time under warming is explained by respired C composition change in the South
American grassland region (Fig. 4a). In contrast, change in respired C
composition only accounts for approximately 10 % of the increase in C
transit time under warming in the boreal and high-latitude regions of North
America.</p>
      <?pagebreak page6568?><p id="d1e3324">The second component is the C age structure, primarily from change in mean C
age of the individual pool modified by the relative fraction of each pool. In the coupled
climate warming with rising atmospheric [<inline-formula><mml:math id="M160" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenario (S3), C age
structure change primarily contributes to the C transit time response in most
global regions in 2100 (Fig. 4h). In this scenario, mean ecosystem C transit
time decreases by 1.6 years. The decrease in C transit time results from
increased young C uptake with rising atmospheric [<inline-formula><mml:math id="M161" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>], which
is more than the increased young C loss with warming. A previous study
has also shown that models with multiple pools usually have a more
heterogeneous C age structure and thus can store extremely old C than a
single pool model (Manzoni et al., 2009).</p>
</sec>
<sec id="Ch1.S4.SS2">
  <title>Deviation arising from estimated C turnover time</title>
      <p id="d1e3355">C turnover time has been widely used to quantify the ecosystem C cycle partly
because both ecosystem C storage and C flux can be easily measured (Sanderman
et al., 2003; Chen et al., 2013; Carvalhais et al., 2014; McCulley et al.,
2004; Raich and Schlesinger, 1992; Yan et al., 2017). The C turnover time has
been theoretically shown to equal C transit time at steady state but they
deviate under non-steady states (Sierra et al., 2016). This study illustrates
how much deviation occurs between C transit time and C turnover time in
response to three scenarios of climate change. Our results show that even at
initial steady state, global ecosystem C turnover time is slightly greater
than C transit time by 3 %. This is because the steady state reached by
spin-up does not mean the terrestrial C cycle system is completely at
equilibrium. Seasonal variations in ecosystem C uptake and turnover still
lead to periodical oscillation of the terrestrial C cycle.</p>
      <p id="d1e3358">The deviation between C transit time and turnover time also indicates to what
extent that turnover time can properly represent time characteristics in the C
cycle. In the climate warming and rising atmospheric [<inline-formula><mml:math id="M162" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>]
scenario (S3), the deviation does not increase significantly until 2050. The
modelled latitudinal pattern of present-day C transit time matches the C
turnover time estimated from observations well (Carvalhais et al., 2014).
This
indicates that the stock-over-flux estimates are still useful at present day.
However, the deviation between C transit time and turnover time remarkably
increases after 2050 (Fig. 5b). Then it requires caution when we use the C
turnover time for estimating C sequestration in multiple compartmental
ecosystems.</p>
      <p id="d1e3372">In transient state, the changes in C transit time and C turnover time differ
the most in the climate warming scenario (S1). Tropical and high-latitude regions
contribute the most to the deviation (Fig. 6h, i). In tropical and
subtropical regions, C transit time increases by about 60 % (Fig. 6b)
while C turnover time increases by 20 % or less (Fig. 6e). The great
difference between changes in C transit time and turnover time is due to
their different assumptions. In response to climate warming, composition
change in respired C contributes most to the change in C transit time in
tropical regions. However, C turnover time assumes all ecosystem C as
one homogenous pool, even if both plant and soil C can be extremely
heterogeneous. This homogeneity assumption ignores the composition changes in
respired C, which causes up to 80 % of change in C transit time.</p>
      <p id="d1e3375">In high-latitude regions, C transit time slightly decreases by up to
10 %, whereas C turnover time considerably decreases by over 30 % in
response to climate warming. Warming significantly increases soil respiration
due to permafrost thaw, whereas the change in permafrost ecosystem C pool
size is relatively small. Thus, C turnover time significantly decreases. C
transit time slowly responds to climate warming because the young C input
added to permafrost ecosystems is relatively small compared to large C storage
in this area and C age structure does not change much. These big deviations
between C turnover time and C transit time in tropical and permafrost regions
suggest that future C cycle analysis based on turnover time likely leads to
strong deviations as it does not represent transient C dynamics in multi-pool
ecosystems.</p>
</sec>
<sec id="Ch1.S4.SS3">
  <title>C transit time versus turnover time under other global change
scenarios</title>
      <p id="d1e3384">This study has illustrated how C transit time and turnover time deviate under
climate warming and rising atmospheric [<inline-formula><mml:math id="M163" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] scenarios. Those
deviations may become even bigger under other global change scenarios. For
example, land use change and fire can drive ecosystems out of steady state to
be at disequilibrium (Luo and Weng, 2011). Clear-cutting of forest or forest fire
removes at least the aboveground wood C pools and thus greatly changes both
the total C stock and NPP, leading to a large change in C turnover time (Wang
et al., 1999; Zhou and Luo, 2008). Clear-cutting of forest or forest fire also
changes age structure and composition of respired C from different pools
within the ecosystem, resulting in change in C transit time. Such a
disturbance usually drives the ecosystem to a stronger degree of disequilibrium
than climate change does. The deviation between turnover time and transit
time should be bigger under a severe disturbance than climate change since
our results have indicated that C transit time and turnover time deviates
more significantly when an ecosystem is further away from equilibrium
(Fig. 5).</p>
      <p id="d1e3398">In contrast to the static vegetation distribution used in CABLE, natural
vegetation distribution may change over time in the real world. C transit
time and turnover time may further deviate under natural vegetation dynamics.
However, whether forest will expand or die back in a future warming world is
still quite unknown. Previous studies come to various conclusions due to their focus
on different areas with different methods (Masek, 2001; Soja et al., 2007;
Cox et al., 2004, 2013). Nevertheless, most bioclimatic models consistently
suggest temperate and boreal biomes rapidly increase in area under warming
(Kirilenko and Solomon, 1998). If the forest species, which stores more C in
slow-turnover tissue, takes over the grass species, which stores more C in
fast-turnover tissue, the expansion of forest may increase C transit time
significantly. However, C turnover time, by lumping all different C
compartments together, may underestimate such changes.</p>
      <p id="d1e3401">In the real world, the land C cycle is always at dynamic disequilibrium due to
cyclic environmental conditions (e.g. diurnal, seasonal, and interannual
variability), directional global change (e.g. climate warming, rising
atmospheric <inline-formula><mml:math id="M164" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> concentration, altered precipitation, and nitrogen
deposition), recursive disturbance–recovery cycles, shifted<?pagebreak page6569?> climatic and
disturbance regimes, and vegetation changes (Luo and Weng, 2011). Thus, the
estimated C turnover time is expected to differ from the C transit time at
any time point and at any spatial location. The degree of deviation between C
turnover time and transit time may vary.</p>
      <p id="d1e3415">In addition to various agents that cause an ecosystem to be at disequilibrium,
deviation between estimated C transit time and turnover time also depends on
model structure. Vertically resolved soil C models, for example, include
vertical C mixing and depth-dependent C decomposition rates (Koven et al.,
2013; Huang et al., 2018). Representation of vertically resolved processes
likely increases soil heterogeneity. When warming induces deep soil thaw and
increases deep soil decomposition, the fraction of respired C from the deep layer
with old C increases. The C transit time together with a vertically
resolved model may substantially increase, whereas C turnover time, which
implicitly assumes the ecosystem to be one homogeneous pool, may not respond much.</p>
</sec>
<sec id="Ch1.S4.SS4">
  <title>Estimation of C transit time in the real world</title>
      <p id="d1e3424">Previous studies have argued that C transit time is conceptually sounder than
C turnover time (Rasmussen et al., 2016; Sierra et al., 2016). In this study,
we have shown that the C turnover time can substantially deviate from the
transit time in response to climate change and other environmental changes.
However, C turnover time can be easily calculated from C stock over flux,
both of which can be easily measured. In contrast, C transit time cannot be
easily estimated from field measurements. Equation (5) indicates that we need
data from measurement of mean C ages (<inline-formula><mml:math id="M165" display="inline"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) and fractional composition of
respired C (<inline-formula><mml:math id="M166" display="inline"><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mrow><mml:mi mathvariant="normal">hr</mml:mi><mml:mo>,</mml:mo><mml:mi>i</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>) in individual C pools in order to calculate
mean ecosystem C transit time (<inline-formula><mml:math id="M167" display="inline"><mml:mrow><mml:msub><mml:mover accent="true"><mml:mi mathvariant="italic">τ</mml:mi><mml:mo mathvariant="normal">‾</mml:mo></mml:mover><mml:mi mathvariant="normal">ts</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>). Neither
<inline-formula><mml:math id="M168" display="inline"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> nor <inline-formula><mml:math id="M169" display="inline"><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mrow><mml:mi mathvariant="normal">hr</mml:mi><mml:mo>,</mml:mo><mml:mi>i</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> have been widely measured in field. Thus, our
research community faces a tremendous challenge to estimate a conceptually
sound and scientifically important parameter.</p>
      <p id="d1e3495">In the past, radiocarbon <inline-formula><mml:math id="M170" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">14</mml:mn></mml:msup></mml:math></inline-formula>C has been used to quantify mean C ages of
various litter and soil pools (Gaudinski et al., 2000). Measured soil
respiration in response to elevated <inline-formula><mml:math id="M171" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> treatment in Duke Forest
has been decomposed to various fractional composition using a deconvolution
method or inverse analysis (Luo et al., 2001). It appears that estimation of
C transit time in real-world ecosystems requires measurement of isotope
signatures in different litter and soil fractions together with measurement
of respiration from soil surface and soil components. Those measurements,
together with many other datasets, may need to be analysed to estimate mean C
ages, fractional composition of respired C in individual C pools, and
then mean ecosystem C transit time (<inline-formula><mml:math id="M172" display="inline"><mml:mrow><mml:msub><mml:mover accent="true"><mml:mi mathvariant="italic">τ</mml:mi><mml:mo mathvariant="normal">‾</mml:mo></mml:mover><mml:mi mathvariant="normal">ts</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) using some
innovative methods, such as data assimilation.</p>
      <p id="d1e3532">Estimating C transit times in the real world can help constrain projections
in land C sequestration because internal timescales of the carbon cycle are a
major source of model uncertainty (Friend et al., 2014; He et al., 2016). Our
study has shown that the change in C transit time can be separated into two
components, C composition change and C age change. Assessment of the two
components would provide additional constraints on model projections. Many of
the ecosystem pools, such as leaf C, wood C, root C pool, litter C pools, and
soil C, can be measured separately. They provide plenty of information to
constrain ecosystem C composition change. Isotope data from each of those
ecosystem components also can offer information to constrain the mean compartment
age. Although discrete soil C pools may not be easy to separate, many
datasets from field and laboratory measurements have been used to constrain
multi-pool soil carbon models by using data assimilation techniques (Liang et
al., 2018; Xu et al., 2006). To constrain C transit time through its two
components with observation, modelled C cycle and land C sequestration can be
significantly improved.</p>
</sec>
</sec>
<sec id="Ch1.S5" sec-type="conclusions">
  <title>Conclusions</title>
      <p id="d1e3543">This study explores how global ecosystem C transit time deviates from the
turnover time under climate warming and rising atmospheric [<inline-formula><mml:math id="M173" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>].
Although both global ecosystem C transit time and turnover time increase in
response to climate warming and decrease in response to rising atmospheric
[<inline-formula><mml:math id="M174" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>], their deviations increase with time in all three climate
change scenarios. In 2100, the deviations are high in tropical regions under
the
climate warming scenario (S1) and rising atmospheric [<inline-formula><mml:math id="M175" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>]
scenario (S2) and in high-latitude regions under S1 and the combined change
scenario (S3). Knowledge about the deviation between C transit time and
turnover time in different regions under different scenarios (warming and
[<inline-formula><mml:math id="M176" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>] rising) is useful for us to understand time characteristics of
the ecosystem carbon dynamics. When we lump all pools and fluxes together to
calculate turnover time by stock over flux, the time characteristic is
different from that of transit time when individual pools and fluxes are
considered within a networked compartmental system. Thus, our results provide
information on how turnover time in the future could deviate from transit
time in specific regions and natural ecosystems under different climate
change scenarios.</p>
      <p id="d1e3590">The changes in C transit time result from both the C age structure changes
and composition changes in respired C in multi-pool ecosystems. The C age
structure changes mainly depend on young C replenishment from external C
input. The composition change is due to differential responses of various C
pools to climate warming and rising atmospheric [<inline-formula><mml:math id="M177" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>]. However, C
turnover time assumes the ecosystem to be one homogeneous pool, and it does not
account for changes in age structure and contribution fractions of different
pools to ecosystem respiration. Thus, C transit time is a better parameter
than C turnover time to characterize the C cycle in multi-pool ecosystems,
especially when they are at transient states.</p>
      <?pagebreak page6570?><p id="d1e3604"><?xmltex \hack{\newpage}?>However, C transit time cannot be easily measured because it requires
information of the C age structure and composition of respired C. Both of
them are usually not measurable in field studies. Radiocarbon <inline-formula><mml:math id="M178" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">14</mml:mn></mml:msup></mml:math></inline-formula>C
measurement in the field has the potential to offer information on mean C
ages in various pools. It is not easy, either, to estimate contribution
fractions of different pools from measured ecosystem or soil respiration to
respired C. We may have to combine compartment models with different types
of measurements via data assimilation techniques to estimate both age
structure and composition of respired C before we can estimate ecosystem C
transit time.</p>
</sec>

      
      </body>
    <back><notes notes-type="dataavailability">

      <p id="d1e3621">The CABLE code is available online
<uri>https://trac.nci.org.au/trac/cable</uri> (Kowalczyk et al., 2006; Wang et
al., 2010, 2011). C age and transit time data are generated using the NCAR
Command Language (NCL) script, which is available online
<uri>http://www2.nau.edu/luo-lab/download/Lu_2018_Biogeosciences.php</uri>.</p>
  </notes><app-group>
        <supplementary-material position="anchor"><p id="d1e3630">The Supplement related to this article is available online at: <inline-supplementary-material xlink:href="https://doi.org/10.5194/bg-15-6559-2018-supplement" xlink:title="pdf">https://doi.org/10.5194/bg-15-6559-2018-supplement</inline-supplementary-material>.</p></supplementary-material>
        </app-group><notes notes-type="authorcontribution">

      <p id="d1e3636">XL, YW, and YL designed the study. XL
conducted the model simulations. XL, YW, YL, and LJ analysed the results
together. All co-authors were involved in writing the paper
and contributed to the study with feedback and critique.</p>
  </notes><notes notes-type="competinginterests">

      <p id="d1e3642">The authors declare that they have no conflict of
interest.</p>
  </notes><ack><title>Acknowledgements</title><p id="d1e3648">This research was financially supported by the postdoctoral fellowship from
the CSIRO Office of Chief Executive to Xingjie Lu, U.S. Department of Energy
grants DE-SC0008270 and DE-SC0014085, and US National Science Foundation (NSF)
grants EF-1807529 and OIA-1301789 to Yiqi Luo EcoLab.<?xmltex \hack{\newline\newline}?>
Edited by: Alexey V. Eliseev <?xmltex \hack{\newline}?>Reviewed by: two anonymous
referees</p></ack><ref-list>
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    <!--<article-title-html>Ecosystem carbon transit versus turnover times in response to climate warming and rising atmospheric CO<sub>2</sub> concentration</article-title-html>
<abstract-html><p>Ecosystem carbon (C) transit time is a critical diagnostic parameter to
characterize land C sequestration. This parameter has different variants in
the literature, including a commonly used turnover time. However, we know
little about how different transit time and turnover time are in representing
carbon cycling through multiple compartments under a non-steady state. In this
study, we estimate both C turnover time as defined by the conventional
stock over flux and mean C transit time as defined by the mean age of C mass
leaving the system. We incorporate them into the Community
Atmosphere Biosphere Land Exchange (CABLE) model to estimate C turnover time
and transit time in response to climate warming and rising
atmospheric [CO<sub>2</sub>]. Modelling analysis shows that both C turnover
time and transit time increase with climate warming but decrease with rising
atmospheric [CO<sub>2</sub>]. Warming increases C turnover time by 2.4 years
and transit time by 11.8 years in 2100 relative to that at steady state in
1901. During the same period, rising atmospheric [CO<sub>2</sub>] decreases C
turnover time by 3.8 years and transit time by 5.5 years. Our analysis shows
that 65&thinsp;% of the increase in global mean C transit time with climate
warming results from the depletion of fast-turnover C pool. The remaining
35&thinsp;% increase results from accompanied changes in compartment C age
structures. Similarly, the decrease in mean C transit time with rising
atmospheric [CO<sub>2</sub>] results approximately equally from replenishment
of C into fast-turnover C pool and subsequent decrease in compartment C age
structure. Greatly different from the transit time, the turnover time, which
does not account for changes in either C age structure or composition of
respired C, underestimated impacts of warming and rising atmospheric
[CO<sub>2</sub>] on C diagnostic time and potentially led to deviations in
estimating land C sequestration in multi-compartmental ecosystems.</p></abstract-html>
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