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  <front>
    <journal-meta>
<journal-id journal-id-type="publisher">NPG</journal-id>
<journal-title-group>
<journal-title>Nonlinear Processes in Geophysics</journal-title>
<abbrev-journal-title abbrev-type="publisher">NPG</abbrev-journal-title>
<abbrev-journal-title abbrev-type="nlm-ta">Nonlin. Processes Geophys.</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1607-7946</issn>
<publisher><publisher-name>Copernicus Publications</publisher-name>
<publisher-loc>Göttingen, Germany</publisher-loc>
</publisher>
</journal-meta>

    <article-meta>
      <article-id pub-id-type="doi">10.5194/npg-24-553-2017</article-id><title-group><article-title>Non-Gaussian data assimilation of satellite-based leaf area index
observations with an individual-based dynamic<?xmltex \hack{\break}?> global vegetation model</article-title>
      </title-group><?xmltex \runningtitle{Non-Gaussian data assimilation of satellite-based LAI observations}?><?xmltex \runningauthor{H. Arakida et al.}?>
      <contrib-group>
        <contrib contrib-type="author" corresp="yes" rid="aff1">
          <name><surname>Arakida</surname><given-names>Hazuki</given-names></name>
          <email>hazuki.arakida@riken.jp</email>
        <ext-link>https://orcid.org/0000-0002-8570-7878</ext-link></contrib>
        <contrib contrib-type="author" corresp="yes" rid="aff1 aff2 aff3">
          <name><surname>Miyoshi</surname><given-names>Takemasa</given-names></name>
          <email>takemasa.miyoshi@riken.jp</email>
        <ext-link>https://orcid.org/0000-0003-3160-2525</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff4">
          <name><surname>Ise</surname><given-names>Takeshi</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1 aff5">
          <name><surname>Shima</surname><given-names>Shin-ichiro</given-names></name>
          
        <ext-link>https://orcid.org/0000-0001-5540-713X</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Kotsuki</surname><given-names>Shunji</given-names></name>
          
        </contrib>
        <aff id="aff1"><label>1</label><institution>RIKEN Advanced Institute for Computational Science, Kobe, 650-0047,
Japan</institution>
        </aff>
        <aff id="aff2"><label>2</label><institution>Department of Atmospheric and Oceanic Science, University of Maryland,
College Park, MD 20742, USA</institution>
        </aff>
        <aff id="aff3"><label>3</label><institution>Application Laboratory, Japan Agency for Marine-Earth Science and
Technology, Yokohama, 236-0001, Japan</institution>
        </aff>
        <aff id="aff4"><label>4</label><institution>Field Science Education and Research Center, Kyoto University, Kyoto,
606-8502, Japan</institution>
        </aff>
        <aff id="aff5"><label>5</label><institution>Graduate School of Simulation Studies, University of Hyogo, Kobe,
650-0047, Japan</institution>
        </aff>
      </contrib-group>
      <author-notes><corresp id="corr1">Hazuki Arakida (hazuki.arakida@riken.jp) and Takemasa Miyoshi
(takemasa.miyoshi@riken.jp)</corresp></author-notes><pub-date><day>6</day><month>September</month><year>2017</year></pub-date>
      
      <volume>24</volume>
      <issue>3</issue>
      <fpage>553</fpage><lpage>567</lpage>
      <history>
        <date date-type="received"><day>6</day><month>May</month><year>2016</year></date>
           <date date-type="rev-request"><day>25</day><month>May</month><year>2016</year></date>
           <date date-type="rev-recd"><day>20</day><month>April</month><year>2017</year></date>
           <date date-type="accepted"><day>24</day><month>July</month><year>2017</year></date>
      </history>
      <permissions>
<license license-type="open-access">
<license-p>This work is licensed under the Creative Commons Attribution 3.0 Unported License. To view a copy of this licence, visit <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/3.0/">https://creativecommons.org/licenses/by/3.0/</ext-link></license-p>
</license>
</permissions><self-uri xlink:href="https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017.html">This article is available from https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017.html</self-uri>
<self-uri xlink:href="https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017.pdf">The full text article is available as a PDF file from https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017.pdf</self-uri>


      <abstract>
    <p>We developed a data assimilation system based on a particle filter
approach with the spatially explicit individual-based dynamic global
vegetation model (SEIB-DGVM). We first performed an idealized observing
system simulation experiment to evaluate the impact of assimilating the leaf
area index (LAI) data every 4 days, simulating the satellite-based LAI.
Although we assimilated only LAI as a whole, the tree and grass LAIs were
estimated separately with high accuracy. Uncertain model parameters and
other state variables were also estimated accurately. Therefore, we extended
the experiment to the real world using the real Moderate Resolution Imaging
Spectroradiometer (MODIS) LAI data and obtained promising results.</p>
  </abstract>
    </article-meta>
  </front>
<body>
      

<sec id="Ch1.S1" sec-type="intro">
  <title>Introduction</title>
      <p>The terrestrial biosphere is an important part of the Earth system model
(ESM) to simulate the carbon and water cycles. However, terrestrial biosphere
models tend to have large uncertainties, for example, in phenology
(Richardson et al., 2012; Murray-Tortarolo et al., 2013) and in spatial
distributions of plant species (Cheaib et al., 2012). Recently, data
assimilation (DA) methods which incorporate observation data into models have
been applied to terrestrial biosphere models to reduce the uncertainties in
the state variables and model parameters (Luo et al., 2011; Peng et al.,
2011). Previous studies have successfully applied the ensemble Kalman filter
(e.g., Evensen, 2003; Williams et al., 2005; Quaife et al., 2008; Stöckli
et al., 2011) or adjoint method (e.g., Kaminski et al., 2013; Kato et al.,
2013) to the “static” vegetation models, but studies
with the “dynamic” global vegetation models (DGVMs) are still limited (Luo
et al., 2011; Peng et al., 2011), although Hartig et al. (2012) pointed out
the importance.</p>
      <p>The static vegetation models are time independent and do not include
the vegetation succession process (Peng, 2000). Alternatively, DGVMs include
the vegetation succession process and can simulate carbon and water cycle
changes linking to the vegetation shift under the changing climate.
Specifically, “individual-based” DGVMs simulate local interactions among
individual plants such as competitions for light and water, so that the model
can simulate the vegetation succession more explicitly (Smith et al., 2001;
Sato et al., 2007). Garreta et al. (2010) pioneered to apply DA to an
individual-based DGVM for paleoclimate, but no study has been published thus
far to assimilate fine timescale data from satellites and ground stations
using an individual-based DGVM. If the initial vegetation structure
and the model parameters of an individual-based DGVM are estimated
more accurately by assimilating the fine timescale data, the uncertainties
of the simulated future vegetation would be greatly reduced.</p>
      <p>This study explores the ability to assimilate frequent satellite-based leaf area index
(LAI) data with an individual-based DGVM known as the SEIB-DGVM,
which stands for the spatially explicit individual-based DGVM (Sato et al., 2007). We
developed a non-Gaussian ensemble DA system with the SEIB-DGVM based on a
particle filter approach. Although the particle filter is an existing,
well-known approach, this is the first attempt to apply it to an
individual-based DGVM with frequent LAI data. Therefore, we focus on
the methodological development in this study and perform a series of
numerical experiments at a single location with only a couple of plant
functional types (PFTs) as the first step. It would be numerically
straightforward to extend it to the global scale in future studies, since
the local-scale experiments can be performed in parallel for different
locations. In the present study, we first perform idealized simulation
experiments to investigate how well we can estimate the model parameters
associated with phenology by assimilating the LAI data every 4 days,
simulating the satellite-based LAI product from the Moderate Resolution
Imaging Spectroradiometer (MODIS) aboard the Terra and Aqua spacecrafts. We
also investigate to what extent assimilating the LAI data could improve the
estimates of the state variables such as GPP (gross primary production), RE
(ecosystem respiration), NEE (net ecosystem exchange), and biomass, the most
fundamental variables for carbon cycle and vegetation states. Sensitivities
to the filter settings such as the random perturbation sizes and particle
sizes are also investigated. Following the idealized experiments, we perform
an experiment using the real MODIS LAI observation data to see how well the
proposed approach performs in the real world.</p>
</sec>
<sec id="Ch1.S2">
  <title>Method</title>
<sec id="Ch1.S2.SS1">
  <title>SEIB-DGVM</title>
      <p>The SEIB-DGVM simulates establishment, growth, and decay of the individuals
of prescribed PFTs within a spatially explicit virtual forest (Sato et al.,
2007), forced by climate conditions such as air temperature, soil
temperature, cloudiness, precipitation, humidity, and winds. We used
version 2.71 (Sato and Ise, 2012) but with minimal modifications for DA. The
model simulates daily states, but the original model outputs were only once
per year. Outputs are needed for DA once every 4 days; thus, we modified
the model code to output the model states every 4 days. In addition, the
original model code assumed running for many years continuously, and the
initial seed for the random number generator was fixed. As a result, in this
study, we stop the model every 4 days, and the same seed is repeated every time when
we start the model. Therefore, we modified the model code to randomly
generate the seed for the random number generator every time when we initiate
the model. Other modifications are summarized in Appendix A.</p>
      <p><?xmltex \hack{\newpage}?>The size of the model state space is determined by the prognostic variables
for tree, grass, forest as a whole, and soil. Each individual tree has 13
prognostic variables such as biomass of root, leaf and trunk, and we assume
that up to 300 trees can exist in the forest area. Therefore, the number of
tree variables is less than or equal to 3900 (i.e., <inline-formula><mml:math id="M1" display="inline"><mml:mrow><mml:mn mathvariant="normal">300</mml:mn><mml:mo>×</mml:mo><mml:mn mathvariant="normal">13</mml:mn></mml:mrow></mml:math></inline-formula>). As for
grass, the forest area is divided into 30 by 30 grid cells, and each grid
cell has four variables such as biomass of root and leaf. Hence, the number of
grass variables is fixed at 3600 (i.e., <inline-formula><mml:math id="M2" display="inline"><mml:mrow><mml:mn mathvariant="normal">30</mml:mn><mml:mo>×</mml:mo><mml:mn mathvariant="normal">30</mml:mn><mml:mo>×</mml:mo><mml:mn mathvariant="normal">4</mml:mn></mml:mrow></mml:math></inline-formula>). In
addition, forest as a whole has eight prognostic variables such as snow and soil
carbon mass, and finally, soil moisture (one variable) is defined for 30 soil
layers. Therefore, the number of state variables is between 3638 (no tree,
i.e., 0 <inline-formula><mml:math id="M3" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> 3600 <inline-formula><mml:math id="M4" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> 8 <inline-formula><mml:math id="M5" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> 30) and 7538 (300 trees, i.e.,
3900 <inline-formula><mml:math id="M6" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> 3600 <inline-formula><mml:math id="M7" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> 8 <inline-formula><mml:math id="M8" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> 30).</p>
      <p>Among the various model outputs ranging from individual tree height to soil
water content (Sato et al., 2007, with updated information available from
the package of version 2.71), we focus on LAI because it is the key to the
vegetation model, and because previous studies show a promise in
assimilating satellite-based LAI data with a static vegetation model (Stöckli
et al., 2011) and a “non-individual-based” DGVM (Demarty et al., 2007). We extend the previous
studies to assimilate the LAI data with the individual-based DGVM.</p>
</sec>
<sec id="Ch1.S2.SS2">
  <title>Particle filter-based DA</title>
      <p>Individual-based DGVMs include highly non-linear processes such as occasional establishment
and death of individual plants. These processes produce and eliminate state
variables, and the phase space changes time to time. DA methods that have
been used in geophysical applications usually assume that the state
variables are defined uniquely for the given dynamical system and that the
phase space dimension stays the same. The widely used ensemble Kalman
filter, for example, finds the best linear combination of the ensemble with
optimal fit to the observations, but it is not trivial to define a linear
combination or even the ensemble mean for the variables missing in some
ensemble members. Therefore, it would not be trivial to apply the
widely used DA methods to individual-based DGVMs.</p>
      <p>Alternatively, particle filters run independent parallel simulations or
particles and represent the probability density function (PDF) explicitly by
assigning probability to each particle. Therefore, particle filters can
handle non-Gaussianity and non-linearity explicitly, and can be applied to
the individual-based DGVMs in a straightforward manner (e.g., Garreta et al., 2010) even
though the phase space dimension is different for each particle.</p>
      <p>Here, we adopt a particle filter approach known as sequential importance
resampling (SIR; Fig. 1) (Gordon et al., 1993). Although the method is not
efficient for large dimensional systems (e.g., Bickel et al., 2008; Snyder et
al., 2008, 2015; Snyder, 2012), we tested this well-known method as the first attempt
to construct the DA system with SEIB-DGVM. First, <inline-formula><mml:math id="M9" display="inline"><mml:mi>n</mml:mi></mml:math></inline-formula> parallel simulations
are performed, and each simulation is considered as a particle representing
the true state of the system with equal probability. Next, likelihood
<inline-formula><mml:math id="M10" display="inline"><mml:mrow><mml:msubsup><mml:mi>l</mml:mi><mml:mi>t</mml:mi><mml:mrow><mml:mo>(</mml:mo><mml:mi>i</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:msubsup></mml:mrow></mml:math></inline-formula> is calculated for each particle using the Gaussian likelihood
function:</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F1"><caption><p>Schematic showing the SIR
particle filter method. The size of the circles corresponds to the assigned
probability.</p></caption>
          <?xmltex \igopts{width=236.157874pt}?><graphic xlink:href="https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017-f01.pdf"/>

        </fig>

      <p><disp-formula specific-use="align" content-type="numbered"><mml:math id="M11" display="block"><mml:mtable displaystyle="true"><mml:mlabeledtr id="Ch1.E1"><mml:mtd/><mml:mtd><mml:mrow><mml:mstyle class="stylechange" displaystyle="true"/><mml:msubsup><mml:mi>l</mml:mi><mml:mi>t</mml:mi><mml:mrow><mml:mo>(</mml:mo><mml:mi>i</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:msubsup></mml:mrow></mml:mtd><mml:mtd><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:mo>=</mml:mo><mml:mi>p</mml:mi><mml:mfenced close=")" open="("><mml:msub><mml:mi>y</mml:mi><mml:mi>t</mml:mi></mml:msub><mml:mi mathvariant="normal">|</mml:mi><mml:msubsup><mml:mi>x</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mi mathvariant="normal">|</mml:mi><mml:mi>t</mml:mi><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mrow><mml:mo>(</mml:mo><mml:mi>i</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:msubsup></mml:mfenced><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mn mathvariant="normal">1</mml:mn><mml:msqrt><mml:mrow><mml:mn mathvariant="normal">2</mml:mn><mml:mi mathvariant="italic">π</mml:mi><mml:mo>⋅</mml:mo><mml:msup><mml:mi mathvariant="italic">σ</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:msqrt></mml:mfrac></mml:mstyle><mml:mi mathvariant="normal">exp</mml:mi><mml:mfenced close="}" open="{"><mml:mo>-</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:mo>(</mml:mo><mml:msub><mml:mi>y</mml:mi><mml:mi>t</mml:mi></mml:msub><mml:mo>-</mml:mo><mml:msubsup><mml:mi>x</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mi mathvariant="normal">|</mml:mi><mml:mi>t</mml:mi><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mrow><mml:mo>(</mml:mo><mml:mi>i</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:msubsup><mml:msup><mml:mo>)</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow><mml:mrow><mml:mn mathvariant="normal">2</mml:mn><mml:mo>⋅</mml:mo><mml:msup><mml:mi mathvariant="italic">σ</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:mfrac></mml:mstyle></mml:mfenced></mml:mrow></mml:mtd></mml:mlabeledtr><mml:mtr><mml:mtd><mml:mstyle class="stylechange" displaystyle="true"/></mml:mtd><mml:mtd><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:mtext>for</mml:mtext><mml:mspace width="0.25em" linebreak="nobreak"/><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn><mml:mo>,</mml:mo><mml:mi mathvariant="normal">…</mml:mi><mml:mo>,</mml:mo><mml:mi>n</mml:mi><mml:mo>.</mml:mo></mml:mrow></mml:mtd></mml:mtr></mml:mtable></mml:math></disp-formula>

            Here, <inline-formula><mml:math id="M12" display="inline"><mml:mrow><mml:msubsup><mml:mi>x</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mi mathvariant="normal">|</mml:mi><mml:mi>t</mml:mi><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mrow><mml:mo>(</mml:mo><mml:mi>i</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:msubsup></mml:mrow></mml:math></inline-formula> denotes the simulated LAI of the <inline-formula><mml:math id="M13" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th particle
at time <inline-formula><mml:math id="M14" display="inline"><mml:mi>t</mml:mi></mml:math></inline-formula> from the previous time step <inline-formula><mml:math id="M15" display="inline"><mml:mrow><mml:mi>t</mml:mi><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula><mml:math id="M16" display="inline"><mml:mrow><mml:msub><mml:mi>y</mml:mi><mml:mi>t</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> the observed LAI at
time <inline-formula><mml:math id="M17" display="inline"><mml:mi>t</mml:mi></mml:math></inline-formula>, and <inline-formula><mml:math id="M18" display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> the observation error standard deviation. Since the
prior probability is uniform, Bayes' rule gives that the posterior
probability of the <inline-formula><mml:math id="M19" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th particle is proportional to <inline-formula><mml:math id="M20" display="inline"><mml:mrow><mml:msubsup><mml:mi>l</mml:mi><mml:mi>t</mml:mi><mml:mrow><mml:mo>(</mml:mo><mml:mi>i</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:msubsup></mml:mrow></mml:math></inline-formula>, i.e., the
particles closer to the observation have more probability. Next, we resample
the particles, so that each particle has equal probability. The particles
with more probability (larger <inline-formula><mml:math id="M21" display="inline"><mml:mrow><mml:msubsup><mml:mi>l</mml:mi><mml:mi>t</mml:mi><mml:mrow><mml:mo>(</mml:mo><mml:mi>i</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:msubsup><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> are duplicated, and the
particles with less probability (smaller <inline-formula><mml:math id="M22" display="inline"><mml:mrow><mml:msubsup><mml:mi>l</mml:mi><mml:mi>t</mml:mi><mml:mrow><mml:mo>(</mml:mo><mml:mi>i</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:msubsup><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> are removed. If <inline-formula><mml:math id="M23" display="inline"><mml:mi>n</mml:mi></mml:math></inline-formula>
is sufficiently large, we can evaluate the posterior PDF accurately. Each
resampled particle represents the true state of the system with equal
probability and acts as the initial particle for the next time step. This
Bayesian framework is repeated.</p>
</sec>
<sec id="Ch1.S2.SS3">
  <title>OSSE and the real-world experiment</title>
      <p>We first perform a series of idealized observing system simulation
experiments (OSSEs). The OSSE (e.g., Atlas, 1997) is a widely used approach
in meteorological DA to test the general performance of a DA system and to
evaluate the impact of specific observing systems. OSSE has the nature run,
which is usually generated by running a simulation for a certain period.
Observation data are simulated from the nature run by applying the
observation operator, i.e., converting the model variables to the observed
variables. Here, we add artificial random noise to simulate the observation
error. DA experiments are initiated from the state independent of the nature
run, and the simulated observations are assimilated. The resulting analyses
and subsequent forecasts are compared with the nature run to evaluate the
performance of DA. Once an OSSE is done, it is straightforward to extend the
OSSE to the real world by simply replacing the simulated observations with
the real-world observations.</p>
</sec>
</sec>
<sec id="Ch1.S3">
  <title>OSSE</title>
<sec id="Ch1.S3.SS1">
  <title>Experimental design</title>
      <p>To generate the nature run, the SEIB-DGVM was initialized with the bare
ground (i.e., no plant at the beginning) and was run for 107 years using the
climate forcing data from 2001 to 2010 available at the SEIB-DGVM
web page (<uri>http://seib-dgvm.com/</uri>). Here, the 10-year forcing data are
repeated for the 107-year simulation, and the last 7 years from years 101 to
107 use the actual climate forcing of 2001 to 2007; thus, we refer to years 101
to 107 as 2001 to 2007. The daily climate data were generated by the
procedure of Sato and Ise (2012) with updated information available at the
SEIB-DGVM web page, based on the monthly Climate Research Unit
observation-based data (CRU-TS3.22 0.5<inline-formula><mml:math id="M24" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> monthly climate time series)
(Harris et al., 2014) and the daily data from the National Centers for
Environmental Prediction (NCEP)/National Center for Atmospheric Research
(NCAR) reanalysis (Kalnay et al., 1996). We chose the study area at one of
the AsiaFlux sites, the Siberia Yakutsk larch forest site at Spasskaya Pad,
the middle basin of the Lena River (62<inline-formula><mml:math id="M25" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>15<inline-formula><mml:math id="M26" display="inline"><mml:msup><mml:mi/><mml:mo>′</mml:mo></mml:msup></mml:math></inline-formula>18<inline-formula><mml:math id="M27" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>′</mml:mo><mml:mo>′</mml:mo></mml:mrow></mml:msup></mml:math></inline-formula> N,
129<inline-formula><mml:math id="M28" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>14<inline-formula><mml:math id="M29" display="inline"><mml:msup><mml:mi/><mml:mo>′</mml:mo></mml:msup></mml:math></inline-formula>29<inline-formula><mml:math id="M30" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>′</mml:mo><mml:mo>′</mml:mo></mml:mrow></mml:msup></mml:math></inline-formula> E). The observed climate data at this site were not
directly used in this study, but these data may have been included in the
NCEP/NCAR reanalysis. Field-observed carbon flux data are available as the
ground truth to verify the DA results at this site. Forced by the climate
data, the SEIB-DGVM simulates the vegetation shifts from the bare ground to a
grassland, and then to a forest. The two PFTs, the boreal deciduous needleleaf trees and C3 grass, are the dominant PFTs in this study area.
Therefore, we do not consider the other PFTs in this study following Sato et
al. (2010). We call these two PFTs simply “tree” and “grass”.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F2" specific-use="star"><caption><p>Schematic illustrations of the nature run, observations, and model
parameter sensitivities. <bold>(a)</bold> Time series of LAI (m<inline-formula><mml:math id="M31" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:math></inline-formula> m<inline-formula><mml:math id="M32" display="inline"><mml:mrow><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>
for the nature run (black), simulated observations (red dots), and their
error standard deviations (SD, red error bars). <bold>(b)</bold> Time series of
LAI with different Pmax and Dor values. The perturbed parameters (Pmax and
Dor for tree and grass) cause differences between the particles.</p></caption>
          <?xmltex \igopts{width=327.206693pt}?><graphic xlink:href="https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017-f02.pdf"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F3" specific-use="star"><caption><p>Time series of LAI for <bold>(a)</bold> tree plus grass, <bold>(b)</bold> tree, and <bold>(c)</bold> grass
for an experiment without DA (NODA, left) and an experiment with DA
(TEST, right). Dark and light gray areas indicate the
quartiles and 1–99 % quantiles of the particles as shown in the legend. Thick
black curves indicate the medians. Blue dots with error bars indicate the
observations and their error standard deviations, and red lines indicate the
nature run.</p></caption>
          <?xmltex \igopts{width=389.802756pt}?><graphic xlink:href="https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017-f03.pdf"/>

        </fig>

      <p>The nature run (Fig. 2a) was performed with the “true” parameter values
Pmax <inline-formula><mml:math id="M33" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 15 <inline-formula><mml:math id="M34" display="inline"><mml:mi mathvariant="normal">µ</mml:mi></mml:math></inline-formula>molCO<inline-formula><mml:math id="M35" display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> m<inline-formula><mml:math id="M36" 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> s<inline-formula><mml:math id="M37" 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> and
Dor <inline-formula><mml:math id="M38" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 230 DOY (day of year) for tree and
Pmax <inline-formula><mml:math id="M39" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 9 <inline-formula><mml:math id="M40" display="inline"><mml:mi mathvariant="normal">µ</mml:mi></mml:math></inline-formula>molCO<inline-formula><mml:math id="M41" display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> m<inline-formula><mml:math id="M42" 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> s<inline-formula><mml:math id="M43" 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> and
Dor <inline-formula><mml:math id="M44" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 270 DOY for grass, where Pmax and Dor stand for the maximum
photosynthesis rate and the start date of the dormancy, respectively
(Fig. 2b). Hereafter, we omit the units for Pmax
(<inline-formula><mml:math id="M45" display="inline"><mml:mi mathvariant="normal">µ</mml:mi></mml:math></inline-formula>molCO<inline-formula><mml:math id="M46" display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> m<inline-formula><mml:math id="M47" 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> s<inline-formula><mml:math id="M48" display="inline"><mml:mrow><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> and Dor (DOY) for simplicity. The
LAI observations for the last 4 years from 2004 to 2007 were created by
adding independent Gaussian random noise to the LAI values from the nature
run (Fig. 2a) every 4 days, simulating the MODIS LAI product. Here, the
observation error standard deviation was given by 10 % of the nature run
LAI value. The observed LAI values of less than 0.5 were not used for DA because the MODIS
data for the real-world experiment did not include LAI values of less than 0.5. There are
too few data with real MODIS LAI values of less than 0.5, and we assign the missing value
in preprocessing. Since the LAI is observed only for values of 0.5 or larger, the LAI
observation exists only in the summer season.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F4" specific-use="star"><caption><p>Similar to Fig. 3 but for the model parameters: <bold>(a)</bold> Pmax for
tree, <bold>(b)</bold> Pmax for grass, <bold>(c)</bold> Dor for tree, and <bold>(d)</bold> Dor for grass. There is no
observation for these parameters.</p></caption>
          <?xmltex \igopts{width=389.802756pt}?><graphic xlink:href="https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017-f04.pdf"/>

        </fig>

      <p>Next, 8000 particles (parallel simulations) were generated with uniformly
perturbed parameters: Pmax <inline-formula><mml:math id="M49" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> [0, 60] for tree, Pmax <inline-formula><mml:math id="M50" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> [0, 15] for
grass, and Dor <inline-formula><mml:math id="M51" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> [200, 300] for both. Here, [<inline-formula><mml:math id="M52" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula>, <inline-formula><mml:math id="M53" display="inline"><mml:mi>b</mml:mi></mml:math></inline-formula>] denotes random draws
from the uniform distribution between <inline-formula><mml:math id="M54" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> and <inline-formula><mml:math id="M55" display="inline"><mml:mi>b</mml:mi></mml:math></inline-formula>. These initial perturbation
sizes are based on the previous studies (Kolari et., 2006; Zeng et al., 2011;
Zhao et al., 2015; Takagi et al., 2015). We ran 8000 parallel simulations for
103 years for spin-up from the bare ground using the same climate forcing
data as the nature run. In the course of the vegetation succession, these
randomly perturbed parameter sets result in a variety of LAI simulations
(Fig. 2b).</p>
      <p>The 8000 particles at the end of the 103-year spin-up runs are used as the
initial conditions for DA. The simulated LAI observations are assimilated
every 4 days. The nature run and particle filter use the same climate forcing
data, so that the difference comes from the model parameter values. The
particles continue to be the free runs until the first LAI observation is
assimilated in the summer season. The state variables and model parameters
are estimated together at DA, and the model systematic errors associated with
the model parameters are corrected by DA with parameter estimation. No
explicit bias correction is applied. To avoid the exact duplications after
resampling, the model parameters Pmax and Dor are randomly perturbed for the
duplicated particles. The random perturbations avoid particle degeneracy,
which usually causes filter divergence. After some tuning, we found proper
perturbation sizes that work for stable filtering without causing particle
degeneracy, especially for biomass which is found to be the most sensitive to
the perturbation sizes. Here, random draws [<inline-formula><mml:math id="M56" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>4, 4] are added to Pmax for
tree and to Dor for both tree and grass, and [<inline-formula><mml:math id="M57" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>1, 1] are added to Pmax for
grass because the initial Pmax perturbation size for grass is a quarter of
that of tree. The sensitivity to the resampling perturbation sizes will be
discussed in the next section. In the case that these perturbed parameters exceed
the corresponding initial parameter range, the excess value was bounced back
from the limits. To assess the impact of DA, we also perform an experiment
without DA (“NODA” hereafter) and compare it to the experiment with DA
(“TEST” hereafter).</p>
</sec>
<sec id="Ch1.S3.SS2">
  <title>Results</title>
      <p>Figure 3 shows the time series of LAI for NODA (left) and TEST (right). The
observations (Fig. 3a, blue dots with error bars) cannot distinguish the tree
and grass, but the model simulates LAIs for tree and grass separately
(Fig. 3b, c). Although the particles without DA are widely spread (left, gray
areas), DA makes the particles much narrower (right) and consistent with the
nature run (red curves). With DA, the median of the particles for tree is
almost identical to the nature run for the entire 4 years (Fig. 3b, right).
As for grass, the median of the particles is also very close to the nature
run with DA, but in the first 3 years the dormancy period is delayed
(Fig. 3c, right).</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F5" specific-use="star"><caption><p>Similar to Figs. 3 and 4 but for unobserved model variables:
<bold>(a)</bold> GPP, <bold>(b)</bold> RE, <bold>(c)</bold> NEE, and <bold>(d)</bold> biomass.</p></caption>
          <?xmltex \igopts{width=389.802756pt}?><graphic xlink:href="https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017-f05.pdf"/>

        </fig>

      <p>The model parameters are estimated accurately (Fig. 4). There is no direct
observation of these parameters, so that the estimations are purely due to DA
of the LAI observations. Although the particles of the NODA experiment are
uniformly distributed (Fig. 4, left), DA makes the particles close to the
true parameters (Fig. 4, right). Since we assimilated the LAI only when it was 0.5
or larger, DA has an impact only in the summer season when the leaves grow.
It takes 1–4 years until the true values fall within the quartiles of the
particles. The Pmax estimates for both tree and grass show occasional jumps
but tend to stay around the true values (Fig. 4a, b). Dor for tree seems the
most accurate and stable after the dormancy period of the first year (Fig. 4c).
Dor for grass takes the longest; the estimation is not accurate until the
dormancy of the fourth year (Fig. 4d). This may be related to the previous
results showing the erroneous estimates of the grass LAI near the dormancy
period in the first 3 years (Fig. 3c). The systematic errors in NODA come
from the uncertain parameter settings. TEST can estimate the parameters
through DA and can reduce the systematic errors. This is different from the
bias-correction strategy of the first guess.</p>
      <p>Other model variables such as GPP, RE, NEE, and biomass show large
improvements (Fig. 5). Although the particles of the NODA experiment are
widely spread, DA with only LAI observations greatly reduces the
uncertainties for the four variables, and the estimations are generally
reasonable.</p>
</sec>
</sec>
<sec id="Ch1.S4">
  <title>Sensitivity experiments for OSSE</title>
<sec id="Ch1.S4.SS1">
  <title>Sensitivity to the nature run</title>
      <p>To investigate the sensitivity to the choice of the nature run, we performed
two additional OSSEs, which we call “OSSE2” and “OSSE3”, by generating
different nature runs with different parameter sets (Table 1). The random
numbers for the observation errors are also different. The other settings
follow the TEST experiment.</p>

<?xmltex \floatpos{t}?><table-wrap id="Ch1.T1"><caption><p>Parameter settings for TEST, OSSE2 and OSSE3.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="5">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="right"/>
     <oasis:colspec colnum="3" colname="col3" align="right"/>
     <oasis:colspec colnum="4" colname="col4" align="right"/>
     <oasis:colspec colnum="5" colname="col5" align="right"/>
     <oasis:thead>
       <oasis:row>  
         <oasis:entry colname="col1">OSSEs</oasis:entry>  
         <oasis:entry colname="col2">Pmax</oasis:entry>  
         <oasis:entry colname="col3">Pmax</oasis:entry>  
         <oasis:entry colname="col4">Dor</oasis:entry>  
         <oasis:entry colname="col5">Dor</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">for</oasis:entry>  
         <oasis:entry colname="col3">for</oasis:entry>  
         <oasis:entry colname="col4">for</oasis:entry>  
         <oasis:entry colname="col5">for</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">tree</oasis:entry>  
         <oasis:entry colname="col3">grass</oasis:entry>  
         <oasis:entry colname="col4">tree</oasis:entry>  
         <oasis:entry colname="col5">grass</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>  
         <oasis:entry colname="col1">TEST</oasis:entry>  
         <oasis:entry colname="col2">15</oasis:entry>  
         <oasis:entry colname="col3">9</oasis:entry>  
         <oasis:entry colname="col4">230</oasis:entry>  
         <oasis:entry colname="col5">270</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">OSSE2</oasis:entry>  
         <oasis:entry colname="col2">20</oasis:entry>  
         <oasis:entry colname="col3">12</oasis:entry>  
         <oasis:entry colname="col4">220</oasis:entry>  
         <oasis:entry colname="col5">260</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">OSSE3</oasis:entry>  
         <oasis:entry colname="col2">25</oasis:entry>  
         <oasis:entry colname="col3">7</oasis:entry>  
         <oasis:entry colname="col4">210</oasis:entry>  
         <oasis:entry colname="col5">280</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

      <p>The results show that both OSSE2 and OSSE3 perform well in general. Namely,
the LAI and parameters are estimated generally well (Fig. 6). We find the
main difference between OSSE2 and OSSE3 in the parameters for grass (Fig. 6c,
e). OSSE3 shows significantly larger uncertainties for the parameters for
grass. In OSSE2, the Pmax value for grass is larger and produces more grass
LAI. Since grass starts to grow earlier and stays longer than tree, it is
critical to have LAI observations near the emerging and falling periods for
estimating the grass parameters. Due to the larger Pmax value for grass in
OSSE2, LAI can be observed with the observing threshold of LAI of 0.5 near
the emerging and falling periods. By contrast, in OSSE3, the Pmax value for
grass is smaller, and the small grass LAI of less than 0.5 cannot be observed. We
can see this in the LAI time series (Fig. 6a, right) near the tails in the
spring and fall seasons every year. The uncertainties of LAI are not reduced
year by year, corresponding to the large uncertainties of the grass
parameters. In the summer, LAI becomes larger mostly due to trees, so that
the tree parameters can be estimated well.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F6" specific-use="star"><caption><p>Similar to Figs. 3 and 4 but for OSSE2 (left) and OSSE3 (right).
<bold>(a)</bold> Time series of LAI for tree plus grass, <bold>(b)</bold> Pmax for tree, <bold>(c)</bold> Pmax for
grass, <bold>(d)</bold> Dor for tree, and <bold>(e)</bold> Dor for grass.</p></caption>
          <?xmltex \igopts{width=389.802756pt}?><graphic xlink:href="https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017-f06.pdf"/>

        </fig>

</sec>
<sec id="Ch1.S4.SS2">
  <title>Sensitivity to the initial perturbation size</title>
      <p>Here, we investigate the sensitivity to the initial perturbation sizes with
particle sizes ranging from 1000 to 16 000. Table 2 shows the three initial
perturbation settings: small, moderate, and large. For the TEST experiment,
the moderate initial perturbation sizes were used. We perform additional
sensitivity experiments with the small and large initial perturbation sizes.
Except for the initial perturbation sizes and the particle size, the
experiments follow the TEST experiment.</p>

<?xmltex \floatpos{t}?><table-wrap id="Ch1.T2"><caption><p>Initial perturbation settings.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="5">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="right"/>
     <oasis:colspec colnum="3" colname="col3" align="right"/>
     <oasis:colspec colnum="4" colname="col4" align="right"/>
     <oasis:colspec colnum="5" colname="col5" align="right"/>
     <oasis:thead>
       <oasis:row>  
         <oasis:entry colname="col1">Initial</oasis:entry>  
         <oasis:entry colname="col2">Pmax</oasis:entry>  
         <oasis:entry colname="col3">Pmax</oasis:entry>  
         <oasis:entry colname="col4">Dor</oasis:entry>  
         <oasis:entry colname="col5">Dor</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">perturbation</oasis:entry>  
         <oasis:entry colname="col2">for</oasis:entry>  
         <oasis:entry colname="col3">for</oasis:entry>  
         <oasis:entry colname="col4">for</oasis:entry>  
         <oasis:entry colname="col5">for</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">sizes</oasis:entry>  
         <oasis:entry colname="col2">tree</oasis:entry>  
         <oasis:entry colname="col3">grass</oasis:entry>  
         <oasis:entry colname="col4">tree</oasis:entry>  
         <oasis:entry colname="col5">grass</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>  
         <oasis:entry colname="col1">Small</oasis:entry>  
         <oasis:entry colname="col2">[0, 20]</oasis:entry>  
         <oasis:entry colname="col3">[0, 10]</oasis:entry>  
         <oasis:entry colname="col4">[200, 250]</oasis:entry>  
         <oasis:entry colname="col5">[250, 300]</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Moderate</oasis:entry>  
         <oasis:entry colname="col2">[0, 60]</oasis:entry>  
         <oasis:entry colname="col3">[0, 15]</oasis:entry>  
         <oasis:entry colname="col4">[200, 300]</oasis:entry>  
         <oasis:entry colname="col5">[200, 300]</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Large</oasis:entry>  
         <oasis:entry colname="col2">[0, 120]</oasis:entry>  
         <oasis:entry colname="col3">[0, 30]</oasis:entry>  
         <oasis:entry colname="col4">[150, 350]</oasis:entry>  
         <oasis:entry colname="col5">[150, 350]</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

<?xmltex \floatpos{p}?><table-wrap id="Ch1.T3" specific-use="star" orientation="landscape"><caption><p>Results for the sensitivity experiments on the initial perturbation
size: <bold>(a)</bold> mean absolute error (MAE) and <bold>(b)</bold> the widths of
the 1–99 % quantiles, averaged over a year in 2007. Italic font shows
the filter divergence. Bold letters show the TEST experiment (8000 particles
with moderate initial perturbations).</p></caption><oasis:table frame="topbot"><?xmltex \begin{scaleboxenv}{.90}[.90]?><oasis:tgroup cols="12">
     <oasis:colspec colnum="1" colname="col1" align="center"/>
     <oasis:colspec colnum="2" colname="col2" align="left"/>
     <oasis:colspec colnum="3" colname="col3" align="left"/>
     <oasis:colspec colnum="4" colname="col4" align="right"/>
     <oasis:colspec colnum="5" colname="col5" align="right"/>
     <oasis:colspec colnum="6" colname="col6" align="right"/>
     <oasis:colspec colnum="7" colname="col7" align="right"/>
     <oasis:colspec colnum="8" colname="col8" align="right"/>
     <oasis:colspec colnum="9" colname="col9" align="right"/>
     <oasis:colspec colnum="10" colname="col10" align="right"/>
     <oasis:colspec colnum="11" colname="col11" align="right"/>
     <oasis:colspec colnum="12" colname="col12" align="right"/>
     <oasis:thead>
       <oasis:row>  
         <oasis:entry colname="col1"><bold>(a)</bold></oasis:entry>  
         <oasis:entry colname="col2">Particle</oasis:entry>  
         <oasis:entry colname="col3">Initial</oasis:entry>  
         <oasis:entry colname="col4">Pmax for</oasis:entry>  
         <oasis:entry colname="col5">Pmax for</oasis:entry>  
         <oasis:entry colname="col6">Dor  for</oasis:entry>  
         <oasis:entry colname="col7">Dor for</oasis:entry>  
         <oasis:entry colname="col8">LAI</oasis:entry>  
         <oasis:entry colname="col9">Biomass</oasis:entry>  
         <oasis:entry colname="col10">GPP</oasis:entry>  
         <oasis:entry colname="col11">RE</oasis:entry>  
         <oasis:entry colname="col12">NEE</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">sizes</oasis:entry>  
         <oasis:entry colname="col3">perturbation sizes</oasis:entry>  
         <oasis:entry colname="col4">tree</oasis:entry>  
         <oasis:entry colname="col5">grass</oasis:entry>  
         <oasis:entry colname="col6">tree</oasis:entry>  
         <oasis:entry colname="col7">grass</oasis:entry>  
         <oasis:entry colname="col8">(m<inline-formula><mml:math id="M58" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:math></inline-formula> m<inline-formula><mml:math id="M59" 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>)</oasis:entry>  
         <oasis:entry colname="col9">(MgC ha<inline-formula><mml:math id="M60" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>  
         <oasis:entry colname="col10">(MgC ha<inline-formula><mml:math id="M61" 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> day<inline-formula><mml:math id="M62" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>  
         <oasis:entry colname="col11">(MgC ha<inline-formula><mml:math id="M63" 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> day<inline-formula><mml:math id="M64" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>  
         <oasis:entry colname="col12">(MgC ha<inline-formula><mml:math id="M65" 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> day<inline-formula><mml:math id="M66" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">1000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">0.76</oasis:entry>  
         <oasis:entry colname="col5">0.75</oasis:entry>  
         <oasis:entry colname="col6">0.43</oasis:entry>  
         <oasis:entry colname="col7">3.68</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9"><italic>2.46</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">3.49</oasis:entry>  
         <oasis:entry colname="col5">1.59</oasis:entry>  
         <oasis:entry colname="col6">0.64</oasis:entry>  
         <oasis:entry colname="col7">0.85</oasis:entry>  
         <oasis:entry colname="col8">0.02</oasis:entry>  
         <oasis:entry colname="col9"><italic>4.75</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4"><italic>9.18</italic></oasis:entry>  
         <oasis:entry colname="col5">0.59</oasis:entry>  
         <oasis:entry colname="col6"><italic>33.41</italic></oasis:entry>  
         <oasis:entry colname="col7"><italic>35.70</italic></oasis:entry>  
         <oasis:entry colname="col8"><italic>0.10</italic></oasis:entry>  
         <oasis:entry colname="col9"><italic>0.21</italic></oasis:entry>  
         <oasis:entry colname="col10"><italic>0.007</italic></oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12"><italic>0.006</italic></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">2000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">0.66</oasis:entry>  
         <oasis:entry colname="col5">0.52</oasis:entry>  
         <oasis:entry colname="col6">0.51</oasis:entry>  
         <oasis:entry colname="col7">3.83</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9"><italic>1.36</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">3.49</oasis:entry>  
         <oasis:entry colname="col5">2.05</oasis:entry>  
         <oasis:entry colname="col6">0.71</oasis:entry>  
         <oasis:entry colname="col7">3.11</oasis:entry>  
         <oasis:entry colname="col8">0.04</oasis:entry>  
         <oasis:entry colname="col9"><italic>3.29</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">3.30</oasis:entry>  
         <oasis:entry colname="col5">1.21</oasis:entry>  
         <oasis:entry colname="col6">1.01</oasis:entry>  
         <oasis:entry colname="col7"><italic>59.52</italic></oasis:entry>  
         <oasis:entry colname="col8">0.10</oasis:entry>  
         <oasis:entry colname="col9"><italic>9.22</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">4000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">0.97</oasis:entry>  
         <oasis:entry colname="col5">0.26</oasis:entry>  
         <oasis:entry colname="col6">0.79</oasis:entry>  
         <oasis:entry colname="col7">2.38</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9">1.16</oasis:entry>  
         <oasis:entry colname="col10">0.002</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">3.55</oasis:entry>  
         <oasis:entry colname="col5">1.20</oasis:entry>  
         <oasis:entry colname="col6">0.63</oasis:entry>  
         <oasis:entry colname="col7">3.02</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9"><italic>2.80</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">3.46</oasis:entry>  
         <oasis:entry colname="col5">0.69</oasis:entry>  
         <oasis:entry colname="col6">1.23</oasis:entry>  
         <oasis:entry colname="col7">2.46</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.39</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">8000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">0.69</oasis:entry>  
         <oasis:entry colname="col5">0.30</oasis:entry>  
         <oasis:entry colname="col6">0.78</oasis:entry>  
         <oasis:entry colname="col7">2.30</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9">0.24</oasis:entry>  
         <oasis:entry colname="col10">0.002</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"><bold>Moderate</bold></oasis:entry>  
         <oasis:entry colname="col4"><bold>2.88</bold></oasis:entry>  
         <oasis:entry colname="col5"><bold>1.10</bold></oasis:entry>  
         <oasis:entry colname="col6"><bold>0.73</bold></oasis:entry>  
         <oasis:entry colname="col7"><bold>7.83</bold></oasis:entry>  
         <oasis:entry colname="col8"><bold>0.03</bold></oasis:entry>  
         <oasis:entry colname="col9"><bold>3.72</bold></oasis:entry>  
         <oasis:entry colname="col10"><bold>0.003</bold></oasis:entry>  
         <oasis:entry colname="col11"><bold>0.001</bold></oasis:entry>  
         <oasis:entry colname="col12"><bold>0.003</bold></oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">3.36</oasis:entry>  
         <oasis:entry colname="col5">0.99</oasis:entry>  
         <oasis:entry colname="col6">1.18</oasis:entry>  
         <oasis:entry colname="col7">3.20</oasis:entry>  
         <oasis:entry colname="col8">0.04</oasis:entry>  
         <oasis:entry colname="col9"><italic>1.60</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">16 000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">2.92</oasis:entry>  
         <oasis:entry colname="col5">1.01</oasis:entry>  
         <oasis:entry colname="col6">0.72</oasis:entry>  
         <oasis:entry colname="col7">6.91</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9">0.80</oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">3.22</oasis:entry>  
         <oasis:entry colname="col5">1.13</oasis:entry>  
         <oasis:entry colname="col6">0.65</oasis:entry>  
         <oasis:entry colname="col7">2.16</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9">3.08</oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">3.02</oasis:entry>  
         <oasis:entry colname="col5">1.03</oasis:entry>  
         <oasis:entry colname="col6">0.39</oasis:entry>  
         <oasis:entry colname="col7">6.26</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9"><italic>1.72</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"><bold>(b)</bold></oasis:entry>  
         <oasis:entry colname="col2">Particle</oasis:entry>  
         <oasis:entry colname="col3">Initial</oasis:entry>  
         <oasis:entry colname="col4">Pmax for</oasis:entry>  
         <oasis:entry colname="col5">Pmax for</oasis:entry>  
         <oasis:entry colname="col6">Dor  for</oasis:entry>  
         <oasis:entry colname="col7">Dor for</oasis:entry>  
         <oasis:entry colname="col8">LAI</oasis:entry>  
         <oasis:entry colname="col9">Biomass</oasis:entry>  
         <oasis:entry colname="col10">GPP</oasis:entry>  
         <oasis:entry colname="col11">RE</oasis:entry>  
         <oasis:entry colname="col12">NEE</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">sizes</oasis:entry>  
         <oasis:entry colname="col3">perturbation sizes</oasis:entry>  
         <oasis:entry colname="col4">tree</oasis:entry>  
         <oasis:entry colname="col5">grass</oasis:entry>  
         <oasis:entry colname="col6">tree</oasis:entry>  
         <oasis:entry colname="col7">grass</oasis:entry>  
         <oasis:entry colname="col8">(m<inline-formula><mml:math id="M67" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:math></inline-formula> m<inline-formula><mml:math id="M68" 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>)</oasis:entry>  
         <oasis:entry colname="col9">(MgC ha<inline-formula><mml:math id="M69" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>  
         <oasis:entry colname="col10">(MgC ha<inline-formula><mml:math id="M70" 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> day<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>)</oasis:entry>  
         <oasis:entry colname="col11">(MgC ha<inline-formula><mml:math id="M72" 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> day<inline-formula><mml:math id="M73" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>  
         <oasis:entry colname="col12">(MgC ha<inline-formula><mml:math id="M74" 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> day<inline-formula><mml:math id="M75" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">1000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">14.93</oasis:entry>  
         <oasis:entry colname="col5">5.10</oasis:entry>  
         <oasis:entry colname="col6">23.44</oasis:entry>  
         <oasis:entry colname="col7">28.26</oasis:entry>  
         <oasis:entry colname="col8">0.17</oasis:entry>  
         <oasis:entry colname="col9"><italic>4.63</italic></oasis:entry>  
         <oasis:entry colname="col10">0.013</oasis:entry>  
         <oasis:entry colname="col11">0.005</oasis:entry>  
         <oasis:entry colname="col12">0.012</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">24.18</oasis:entry>  
         <oasis:entry colname="col5">8.02</oasis:entry>  
         <oasis:entry colname="col6">23.60</oasis:entry>  
         <oasis:entry colname="col7">26.80</oasis:entry>  
         <oasis:entry colname="col8">0.18</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.42</italic></oasis:entry>  
         <oasis:entry colname="col10">0.017</oasis:entry>  
         <oasis:entry colname="col11">0.006</oasis:entry>  
         <oasis:entry colname="col12">0.014</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4"><italic>17.92</italic></oasis:entry>  
         <oasis:entry colname="col5">7.47</oasis:entry>  
         <oasis:entry colname="col6"><italic>19.61</italic></oasis:entry>  
         <oasis:entry colname="col7"><italic>26.97</italic></oasis:entry>  
         <oasis:entry colname="col8"><italic>0.11</italic></oasis:entry>  
         <oasis:entry colname="col9"><italic>0.29</italic></oasis:entry>  
         <oasis:entry colname="col10"><italic>0.010</italic></oasis:entry>  
         <oasis:entry colname="col11">0.005</oasis:entry>  
         <oasis:entry colname="col12"><italic>0.009</italic></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">2000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">16.68</oasis:entry>  
         <oasis:entry colname="col5">5.36</oasis:entry>  
         <oasis:entry colname="col6">27.77</oasis:entry>  
         <oasis:entry colname="col7">30.34</oasis:entry>  
         <oasis:entry colname="col8">0.18</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.93</italic></oasis:entry>  
         <oasis:entry colname="col10">0.014</oasis:entry>  
         <oasis:entry colname="col11">0.005</oasis:entry>  
         <oasis:entry colname="col12">0.012</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">24.08</oasis:entry>  
         <oasis:entry colname="col5">9.05</oasis:entry>  
         <oasis:entry colname="col6">26.24</oasis:entry>  
         <oasis:entry colname="col7">29.50</oasis:entry>  
         <oasis:entry colname="col8">0.20</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.63</italic></oasis:entry>  
         <oasis:entry colname="col10">0.018</oasis:entry>  
         <oasis:entry colname="col11">0.007</oasis:entry>  
         <oasis:entry colname="col12">0.015</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">27.30</oasis:entry>  
         <oasis:entry colname="col5">9.23</oasis:entry>  
         <oasis:entry colname="col6">25.97</oasis:entry>  
         <oasis:entry colname="col7"><italic>54.89</italic></oasis:entry>  
         <oasis:entry colname="col8">0.24</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.62</italic></oasis:entry>  
         <oasis:entry colname="col10">0.017</oasis:entry>  
         <oasis:entry colname="col11">0.006</oasis:entry>  
         <oasis:entry colname="col12">0.014</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">4000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">15.72</oasis:entry>  
         <oasis:entry colname="col5">4.60</oasis:entry>  
         <oasis:entry colname="col6">24.18</oasis:entry>  
         <oasis:entry colname="col7">28.76</oasis:entry>  
         <oasis:entry colname="col8">0.14</oasis:entry>  
         <oasis:entry colname="col9">4.89</oasis:entry>  
         <oasis:entry colname="col10">0.012</oasis:entry>  
         <oasis:entry colname="col11">0.005</oasis:entry>  
         <oasis:entry colname="col12">0.011</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">27.11</oasis:entry>  
         <oasis:entry colname="col5">8.62</oasis:entry>  
         <oasis:entry colname="col6">27.73</oasis:entry>  
         <oasis:entry colname="col7">28.68</oasis:entry>  
         <oasis:entry colname="col8">0.20</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.70</italic></oasis:entry>  
         <oasis:entry colname="col10">0.018</oasis:entry>  
         <oasis:entry colname="col11">0.007</oasis:entry>  
         <oasis:entry colname="col12">0.015</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">27.07</oasis:entry>  
         <oasis:entry colname="col5">8.12</oasis:entry>  
         <oasis:entry colname="col6">27.59</oasis:entry>  
         <oasis:entry colname="col7">29.29</oasis:entry>  
         <oasis:entry colname="col8">0.19</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.60</italic></oasis:entry>  
         <oasis:entry colname="col10">0.016</oasis:entry>  
         <oasis:entry colname="col11">0.006</oasis:entry>  
         <oasis:entry colname="col12">0.014</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">8000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">16.02</oasis:entry>  
         <oasis:entry colname="col5">4.50</oasis:entry>  
         <oasis:entry colname="col6">25.38</oasis:entry>  
         <oasis:entry colname="col7">30.14</oasis:entry>  
         <oasis:entry colname="col8">0.15</oasis:entry>  
         <oasis:entry colname="col9">7.27</oasis:entry>  
         <oasis:entry colname="col10">0.012</oasis:entry>  
         <oasis:entry colname="col11">0.005</oasis:entry>  
         <oasis:entry colname="col12">0.011</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"><bold>Moderate</bold></oasis:entry>  
         <oasis:entry colname="col4"><bold>28.32</bold></oasis:entry>  
         <oasis:entry colname="col5"><bold>9.29</bold></oasis:entry>  
         <oasis:entry colname="col6"><bold>26.23</bold></oasis:entry>  
         <oasis:entry colname="col7"><bold>33.99</bold></oasis:entry>  
         <oasis:entry colname="col8"><bold>0.20</bold></oasis:entry>  
         <oasis:entry colname="col9"><bold>11.40</bold></oasis:entry>  
         <oasis:entry colname="col10"><bold>0.017</bold></oasis:entry>  
         <oasis:entry colname="col11"><bold>0.008</bold></oasis:entry>  
         <oasis:entry colname="col12"><bold>0.015</bold></oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">27.47</oasis:entry>  
         <oasis:entry colname="col5">9.37</oasis:entry>  
         <oasis:entry colname="col6">26.60</oasis:entry>  
         <oasis:entry colname="col7">31.75</oasis:entry>  
         <oasis:entry colname="col8">0.21</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.71</italic></oasis:entry>  
         <oasis:entry colname="col10">0.017</oasis:entry>  
         <oasis:entry colname="col11">0.007</oasis:entry>  
         <oasis:entry colname="col12">0.014</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">16 000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">27.66</oasis:entry>  
         <oasis:entry colname="col5">9.28</oasis:entry>  
         <oasis:entry colname="col6">27.18</oasis:entry>  
         <oasis:entry colname="col7">48.79</oasis:entry>  
         <oasis:entry colname="col8">0.22</oasis:entry>  
         <oasis:entry colname="col9">8.44</oasis:entry>  
         <oasis:entry colname="col10">0.017</oasis:entry>  
         <oasis:entry colname="col11">0.007</oasis:entry>  
         <oasis:entry colname="col12">0.015</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">28.47</oasis:entry>  
         <oasis:entry colname="col5">8.85</oasis:entry>  
         <oasis:entry colname="col6">27.86</oasis:entry>  
         <oasis:entry colname="col7">31.91</oasis:entry>  
         <oasis:entry colname="col8">0.21</oasis:entry>  
         <oasis:entry colname="col9">6.53</oasis:entry>  
         <oasis:entry colname="col10">0.017</oasis:entry>  
         <oasis:entry colname="col11">0.008</oasis:entry>  
         <oasis:entry colname="col12">0.015</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">28.76</oasis:entry>  
         <oasis:entry colname="col5">8.93</oasis:entry>  
         <oasis:entry colname="col6">25.77</oasis:entry>  
         <oasis:entry colname="col7">47.88</oasis:entry>  
         <oasis:entry colname="col8">0.21</oasis:entry>  
         <oasis:entry colname="col9"><italic>2.08</italic></oasis:entry>  
         <oasis:entry colname="col10">0.017</oasis:entry>  
         <oasis:entry colname="col11">0.006</oasis:entry>  
         <oasis:entry colname="col12">0.014</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup><?xmltex \end{scaleboxenv}?></oasis:table></table-wrap>

      <p>Table 3 shows the mean absolute errors (MAEs) and the widths of the
1–99 % quantiles, respectively, averaged over a year in 2007. We
consider that the filter diverges when the MAE is larger than the half width
of the 1–99 % quantiles, as shown by the italic font in the tables. The
results show that the filter diverges for biomass in 10 out of 15
experiments. The five experiments that do not diverge are (4000; small), (8000;
small), (16 000; small), (8000; moderate) <inline-formula><mml:math id="M76" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> TEST, and (16 000;
moderate), where ( ; ) denotes (particle size; initial perturbation sizes).
The (1000; large) experiment causes filter divergence for most variables and parameters.
The (2000; large) experiment shows filter divergence for Dor for grass in addition to
biomass. Sampling a wider interval with a smaller particle size generally
reduces the particle density, or the effective number of the particles, so
that the results seem to be reasonable.</p>
</sec>
<sec id="Ch1.S4.SS3">
  <title>Sensitivity to the resampling perturbation size</title>
      <p>Here, we investigate the sensitivity to the resampling perturbation sizes with
particle sizes ranging from 500 to 16 000, in a similar way as the previous
subsection. Resampling perturbations add random perturbations to Pmax and Dor
when resampling and avoid particle degeneracy. Table 4 shows the three
resampling perturbation settings: small, moderate, and large. For the TEST
experiment, the moderate resampling perturbation sizes were used.</p>

<?xmltex \floatpos{t}?><table-wrap id="Ch1.T4"><caption><p>Resampling perturbation settings.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="5">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="right"/>
     <oasis:colspec colnum="3" colname="col3" align="right"/>
     <oasis:colspec colnum="4" colname="col4" align="right"/>
     <oasis:colspec colnum="5" colname="col5" align="right"/>
     <oasis:thead>
       <oasis:row>  
         <oasis:entry colname="col1">Resampling</oasis:entry>  
         <oasis:entry colname="col2">Pmax</oasis:entry>  
         <oasis:entry colname="col3">Pmax</oasis:entry>  
         <oasis:entry colname="col4">Dor</oasis:entry>  
         <oasis:entry colname="col5">Dor</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">perturbation</oasis:entry>  
         <oasis:entry colname="col2">for</oasis:entry>  
         <oasis:entry colname="col3">for</oasis:entry>  
         <oasis:entry colname="col4">for</oasis:entry>  
         <oasis:entry colname="col5">for</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">sizes</oasis:entry>  
         <oasis:entry colname="col2">tree</oasis:entry>  
         <oasis:entry colname="col3">grass</oasis:entry>  
         <oasis:entry colname="col4">tree</oasis:entry>  
         <oasis:entry colname="col5">grass</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>  
         <oasis:entry colname="col1">Small</oasis:entry>  
         <oasis:entry colname="col2">[<inline-formula><mml:math id="M77" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>2, 2]</oasis:entry>  
         <oasis:entry colname="col3">[<inline-formula><mml:math id="M78" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>0.5, 0.5]</oasis:entry>  
         <oasis:entry colname="col4">[<inline-formula><mml:math id="M79" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>2, 2]</oasis:entry>  
         <oasis:entry colname="col5">[<inline-formula><mml:math id="M80" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>2, 2]</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Moderate</oasis:entry>  
         <oasis:entry colname="col2">[<inline-formula><mml:math id="M81" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>4, 4]</oasis:entry>  
         <oasis:entry colname="col3">[<inline-formula><mml:math id="M82" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>1, 1]</oasis:entry>  
         <oasis:entry colname="col4">[<inline-formula><mml:math id="M83" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>4, 4]</oasis:entry>  
         <oasis:entry colname="col5">[<inline-formula><mml:math id="M84" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>4, 4]</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Large</oasis:entry>  
         <oasis:entry colname="col2">[<inline-formula><mml:math id="M85" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>8, 8]</oasis:entry>  
         <oasis:entry colname="col3">[<inline-formula><mml:math id="M86" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>2, 2]</oasis:entry>  
         <oasis:entry colname="col4">[<inline-formula><mml:math id="M87" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>8, 8]</oasis:entry>  
         <oasis:entry colname="col5">[<inline-formula><mml:math id="M88" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>8, 8]</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

<?xmltex \floatpos{p}?><table-wrap id="Ch1.T5" specific-use="star" orientation="landscape"><caption><p>Similar to Table 3 but for the sensitivity experiments on the
resampling perturbation sizes.</p></caption><oasis:table frame="topbot"><?xmltex \begin{scaleboxenv}{.90}[.90]?><oasis:tgroup cols="12">
     <oasis:colspec colnum="1" colname="col1" align="center"/>
     <oasis:colspec colnum="2" colname="col2" align="left"/>
     <oasis:colspec colnum="3" colname="col3" align="right"/>
     <oasis:colspec colnum="4" colname="col4" align="right"/>
     <oasis:colspec colnum="5" colname="col5" align="right"/>
     <oasis:colspec colnum="6" colname="col6" align="right"/>
     <oasis:colspec colnum="7" colname="col7" align="right"/>
     <oasis:colspec colnum="8" colname="col8" align="right"/>
     <oasis:colspec colnum="9" colname="col9" align="right"/>
     <oasis:colspec colnum="10" colname="col10" align="right"/>
     <oasis:colspec colnum="11" colname="col11" align="right"/>
     <oasis:colspec colnum="12" colname="col12" align="right"/>
     <oasis:thead>
       <oasis:row>  
         <oasis:entry colname="col1"><bold>(a)</bold></oasis:entry>  
         <oasis:entry colname="col2">Particle</oasis:entry>  
         <oasis:entry colname="col3">Resampling</oasis:entry>  
         <oasis:entry colname="col4">Pmax for</oasis:entry>  
         <oasis:entry colname="col5">Pmax for</oasis:entry>  
         <oasis:entry colname="col6">Dor for</oasis:entry>  
         <oasis:entry colname="col7">Dor for</oasis:entry>  
         <oasis:entry colname="col8">LAI</oasis:entry>  
         <oasis:entry colname="col9">Biomass</oasis:entry>  
         <oasis:entry colname="col10">GPP</oasis:entry>  
         <oasis:entry colname="col11">RE</oasis:entry>  
         <oasis:entry colname="col12">NEE</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">sizes</oasis:entry>  
         <oasis:entry colname="col3">perturbation sizes</oasis:entry>  
         <oasis:entry colname="col4">tree</oasis:entry>  
         <oasis:entry colname="col5">grass</oasis:entry>  
         <oasis:entry colname="col6">tree</oasis:entry>  
         <oasis:entry colname="col7">grass</oasis:entry>  
         <oasis:entry colname="col8">(m<inline-formula><mml:math id="M89" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:math></inline-formula> m<inline-formula><mml:math id="M90" 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>)</oasis:entry>  
         <oasis:entry colname="col9">(MgC ha<inline-formula><mml:math id="M91" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>  
         <oasis:entry colname="col10">(MgC ha<inline-formula><mml:math id="M92" 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> day<inline-formula><mml:math id="M93" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>  
         <oasis:entry colname="col11">(MgC ha<inline-formula><mml:math id="M94" 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> day<inline-formula><mml:math id="M95" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>  
         <oasis:entry colname="col12">(MgC ha<inline-formula><mml:math id="M96" 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> day<inline-formula><mml:math id="M97" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">500</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">2.59</oasis:entry>  
         <oasis:entry colname="col5">1.68</oasis:entry>  
         <oasis:entry colname="col6">1.74</oasis:entry>  
         <oasis:entry colname="col7"><italic>15.74</italic></oasis:entry>  
         <oasis:entry colname="col8"><italic>0.08</italic></oasis:entry>  
         <oasis:entry colname="col9"><italic>1.47</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">2.28</oasis:entry>  
         <oasis:entry colname="col5">0.66</oasis:entry>  
         <oasis:entry colname="col6">1.34</oasis:entry>  
         <oasis:entry colname="col7">5.57</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9"><italic>1.43</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">10.13</oasis:entry>  
         <oasis:entry colname="col5">1.25</oasis:entry>  
         <oasis:entry colname="col6">2.55</oasis:entry>  
         <oasis:entry colname="col7">5.29</oasis:entry>  
         <oasis:entry colname="col8">0.04</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.35</italic></oasis:entry>  
         <oasis:entry colname="col10">0.004</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">1000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">0.80</oasis:entry>  
         <oasis:entry colname="col5">0.58</oasis:entry>  
         <oasis:entry colname="col6">1.09</oasis:entry>  
         <oasis:entry colname="col7">1.41</oasis:entry>  
         <oasis:entry colname="col8">0.02</oasis:entry>  
         <oasis:entry colname="col9"><italic>4.62</italic></oasis:entry>  
         <oasis:entry colname="col10">0.002</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">3.49</oasis:entry>  
         <oasis:entry colname="col5">1.59</oasis:entry>  
         <oasis:entry colname="col6">0.64</oasis:entry>  
         <oasis:entry colname="col7">0.85</oasis:entry>  
         <oasis:entry colname="col8">0.02</oasis:entry>  
         <oasis:entry colname="col9"><italic>4.75</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">5.00</oasis:entry>  
         <oasis:entry colname="col5">2.17</oasis:entry>  
         <oasis:entry colname="col6">2.78</oasis:entry>  
         <oasis:entry colname="col7">8.56</oasis:entry>  
         <oasis:entry colname="col8">0.04</oasis:entry>  
         <oasis:entry colname="col9"><italic>4.78</italic></oasis:entry>  
         <oasis:entry colname="col10">0.004</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.004</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">2000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">1.84</oasis:entry>  
         <oasis:entry colname="col5">0.42</oasis:entry>  
         <oasis:entry colname="col6">0.47</oasis:entry>  
         <oasis:entry colname="col7">7.94</oasis:entry>  
         <oasis:entry colname="col8">0.02</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.59</italic></oasis:entry>  
         <oasis:entry colname="col10">0.002</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">3.49</oasis:entry>  
         <oasis:entry colname="col5">2.05</oasis:entry>  
         <oasis:entry colname="col6">0.71</oasis:entry>  
         <oasis:entry colname="col7">3.11</oasis:entry>  
         <oasis:entry colname="col8">0.04</oasis:entry>  
         <oasis:entry colname="col9"><italic>3.29</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">6.84</oasis:entry>  
         <oasis:entry colname="col5">1.26</oasis:entry>  
         <oasis:entry colname="col6">2.60</oasis:entry>  
         <oasis:entry colname="col7">4.26</oasis:entry>  
         <oasis:entry colname="col8">0.05</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.79</italic></oasis:entry>  
         <oasis:entry colname="col10">0.004</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">4000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">1.41</oasis:entry>  
         <oasis:entry colname="col5">0.67</oasis:entry>  
         <oasis:entry colname="col6">0.56</oasis:entry>  
         <oasis:entry colname="col7">0.36</oasis:entry>  
         <oasis:entry colname="col8">0.02</oasis:entry>  
         <oasis:entry colname="col9"><italic>2.94</italic></oasis:entry>  
         <oasis:entry colname="col10">0.002</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">3.55</oasis:entry>  
         <oasis:entry colname="col5">1.20</oasis:entry>  
         <oasis:entry colname="col6">0.63</oasis:entry>  
         <oasis:entry colname="col7">3.02</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9"><italic>2.80</italic></oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">6.74</oasis:entry>  
         <oasis:entry colname="col5">1.17</oasis:entry>  
         <oasis:entry colname="col6">1.66</oasis:entry>  
         <oasis:entry colname="col7">6.94</oasis:entry>  
         <oasis:entry colname="col8">0.05</oasis:entry>  
         <oasis:entry colname="col9">2.15</oasis:entry>  
         <oasis:entry colname="col10">0.004</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">8000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">1.23</oasis:entry>  
         <oasis:entry colname="col5">0.37</oasis:entry>  
         <oasis:entry colname="col6">0.61</oasis:entry>  
         <oasis:entry colname="col7">0.71</oasis:entry>  
         <oasis:entry colname="col8">0.02</oasis:entry>  
         <oasis:entry colname="col9"><italic>3.20</italic></oasis:entry>  
         <oasis:entry colname="col10">0.002</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"><bold>Moderate</bold></oasis:entry>  
         <oasis:entry colname="col4"><bold>2.88</bold></oasis:entry>  
         <oasis:entry colname="col5"><bold>1.10</bold></oasis:entry>  
         <oasis:entry colname="col6"><bold>0.73</bold></oasis:entry>  
         <oasis:entry colname="col7"><bold>7.83</bold></oasis:entry>  
         <oasis:entry colname="col8"><bold>0.03</bold></oasis:entry>  
         <oasis:entry colname="col9"><bold>3.72</bold></oasis:entry>  
         <oasis:entry colname="col10"><bold>0.003</bold></oasis:entry>  
         <oasis:entry colname="col11"><bold>0.001</bold></oasis:entry>  
         <oasis:entry colname="col12"><bold>0.003</bold></oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">7.78</oasis:entry>  
         <oasis:entry colname="col5">1.56</oasis:entry>  
         <oasis:entry colname="col6">1.26</oasis:entry>  
         <oasis:entry colname="col7">7.46</oasis:entry>  
         <oasis:entry colname="col8">0.05</oasis:entry>  
         <oasis:entry colname="col9">3.44</oasis:entry>  
         <oasis:entry colname="col10">0.004</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">16 000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">1.46</oasis:entry>  
         <oasis:entry colname="col5">0.45</oasis:entry>  
         <oasis:entry colname="col6">0.77</oasis:entry>  
         <oasis:entry colname="col7">1.70</oasis:entry>  
         <oasis:entry colname="col8">0.02</oasis:entry>  
         <oasis:entry colname="col9"><italic>3.15</italic></oasis:entry>  
         <oasis:entry colname="col10">0.002</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.002</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">3.22</oasis:entry>  
         <oasis:entry colname="col5">1.13</oasis:entry>  
         <oasis:entry colname="col6">0.65</oasis:entry>  
         <oasis:entry colname="col7">2.16</oasis:entry>  
         <oasis:entry colname="col8">0.03</oasis:entry>  
         <oasis:entry colname="col9">3.08</oasis:entry>  
         <oasis:entry colname="col10">0.003</oasis:entry>  
         <oasis:entry colname="col11">0.001</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">5.67</oasis:entry>  
         <oasis:entry colname="col5">1.71</oasis:entry>  
         <oasis:entry colname="col6">1.50</oasis:entry>  
         <oasis:entry colname="col7">6.07</oasis:entry>  
         <oasis:entry colname="col8">0.05</oasis:entry>  
         <oasis:entry colname="col9">1.36</oasis:entry>  
         <oasis:entry colname="col10">0.004</oasis:entry>  
         <oasis:entry colname="col11">0.002</oasis:entry>  
         <oasis:entry colname="col12">0.003</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"><bold>(b)</bold></oasis:entry>  
         <oasis:entry colname="col2">Particle</oasis:entry>  
         <oasis:entry colname="col3">Resampling</oasis:entry>  
         <oasis:entry colname="col4">Pmax for</oasis:entry>  
         <oasis:entry colname="col5">Pmax for</oasis:entry>  
         <oasis:entry colname="col6">Dor for</oasis:entry>  
         <oasis:entry colname="col7">Dor for</oasis:entry>  
         <oasis:entry colname="col8">LAI</oasis:entry>  
         <oasis:entry colname="col9">Biomass</oasis:entry>  
         <oasis:entry colname="col10">GPP</oasis:entry>  
         <oasis:entry colname="col11">RE</oasis:entry>  
         <oasis:entry colname="col12">NEE</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">sizes</oasis:entry>  
         <oasis:entry colname="col3">perturbation sizes</oasis:entry>  
         <oasis:entry colname="col4">tree</oasis:entry>  
         <oasis:entry colname="col5">grass</oasis:entry>  
         <oasis:entry colname="col6">tree</oasis:entry>  
         <oasis:entry colname="col7">grass</oasis:entry>  
         <oasis:entry colname="col8">(m<inline-formula><mml:math id="M98" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:math></inline-formula> m<inline-formula><mml:math id="M99" 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>)</oasis:entry>  
         <oasis:entry colname="col9">(MgC ha<inline-formula><mml:math id="M100" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>  
         <oasis:entry colname="col10">(MgC ha<inline-formula><mml:math id="M101" 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> day<inline-formula><mml:math id="M102" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>  
         <oasis:entry colname="col11">(MgC ha<inline-formula><mml:math id="M103" 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> day<inline-formula><mml:math id="M104" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>  
         <oasis:entry colname="col12">(MgC ha<inline-formula><mml:math id="M105" 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> day<inline-formula><mml:math id="M106" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">500</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">12.41</oasis:entry>  
         <oasis:entry colname="col5">3.98</oasis:entry>  
         <oasis:entry colname="col6">11.29</oasis:entry>  
         <oasis:entry colname="col7"><italic>18.21</italic></oasis:entry>  
         <oasis:entry colname="col8"><italic>0.13</italic></oasis:entry>  
         <oasis:entry colname="col9"><italic>0.37</italic></oasis:entry>  
         <oasis:entry colname="col10">0.012</oasis:entry>  
         <oasis:entry colname="col11">0.004</oasis:entry>  
         <oasis:entry colname="col12">0.010</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">25.82</oasis:entry>  
         <oasis:entry colname="col5">7.47</oasis:entry>  
         <oasis:entry colname="col6">23.68</oasis:entry>  
         <oasis:entry colname="col7">31.83</oasis:entry>  
         <oasis:entry colname="col8">0.18</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.44</italic></oasis:entry>  
         <oasis:entry colname="col10">0.015</oasis:entry>  
         <oasis:entry colname="col11">0.006</oasis:entry>  
         <oasis:entry colname="col12">0.013</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">47.51</oasis:entry>  
         <oasis:entry colname="col5">11.38</oasis:entry>  
         <oasis:entry colname="col6">52.32</oasis:entry>  
         <oasis:entry colname="col7">40.18</oasis:entry>  
         <oasis:entry colname="col8">0.24</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.70</italic></oasis:entry>  
         <oasis:entry colname="col10">0.022</oasis:entry>  
         <oasis:entry colname="col11">0.008</oasis:entry>  
         <oasis:entry colname="col12">0.018</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">1000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">14.31</oasis:entry>  
         <oasis:entry colname="col5">4.17</oasis:entry>  
         <oasis:entry colname="col6">13.85</oasis:entry>  
         <oasis:entry colname="col7">16.57</oasis:entry>  
         <oasis:entry colname="col8">0.14</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.46</italic></oasis:entry>  
         <oasis:entry colname="col10">0.010</oasis:entry>  
         <oasis:entry colname="col11">0.004</oasis:entry>  
         <oasis:entry colname="col12">0.009</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">24.18</oasis:entry>  
         <oasis:entry colname="col5">8.02</oasis:entry>  
         <oasis:entry colname="col6">23.60</oasis:entry>  
         <oasis:entry colname="col7">26.80</oasis:entry>  
         <oasis:entry colname="col8">0.18</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.42</italic></oasis:entry>  
         <oasis:entry colname="col10">0.017</oasis:entry>  
         <oasis:entry colname="col11">0.006</oasis:entry>  
         <oasis:entry colname="col12">0.014</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">44.68</oasis:entry>  
         <oasis:entry colname="col5">11.57</oasis:entry>  
         <oasis:entry colname="col6">53.50</oasis:entry>  
         <oasis:entry colname="col7">45.94</oasis:entry>  
         <oasis:entry colname="col8">0.24</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.60</italic></oasis:entry>  
         <oasis:entry colname="col10">0.022</oasis:entry>  
         <oasis:entry colname="col11">0.009</oasis:entry>  
         <oasis:entry colname="col12">0.018</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">2000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">14.10</oasis:entry>  
         <oasis:entry colname="col5">4.40</oasis:entry>  
         <oasis:entry colname="col6">12.69</oasis:entry>  
         <oasis:entry colname="col7">31.61</oasis:entry>  
         <oasis:entry colname="col8">0.16</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.46</italic></oasis:entry>  
         <oasis:entry colname="col10">0.010</oasis:entry>  
         <oasis:entry colname="col11">0.004</oasis:entry>  
         <oasis:entry colname="col12">0.009</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">24.08</oasis:entry>  
         <oasis:entry colname="col5">9.05</oasis:entry>  
         <oasis:entry colname="col6">26.24</oasis:entry>  
         <oasis:entry colname="col7">29.50</oasis:entry>  
         <oasis:entry colname="col8">0.20</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.63</italic></oasis:entry>  
         <oasis:entry colname="col10">0.018</oasis:entry>  
         <oasis:entry colname="col11">0.007</oasis:entry>  
         <oasis:entry colname="col12">0.015</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">46.88</oasis:entry>  
         <oasis:entry colname="col5">12.23</oasis:entry>  
         <oasis:entry colname="col6">49.80</oasis:entry>  
         <oasis:entry colname="col7">46.06</oasis:entry>  
         <oasis:entry colname="col8">0.25</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.82</italic></oasis:entry>  
         <oasis:entry colname="col10">0.023</oasis:entry>  
         <oasis:entry colname="col11">0.009</oasis:entry>  
         <oasis:entry colname="col12">0.018</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">4000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">14.27</oasis:entry>  
         <oasis:entry colname="col5">5.03</oasis:entry>  
         <oasis:entry colname="col6">13.62</oasis:entry>  
         <oasis:entry colname="col7">15.53</oasis:entry>  
         <oasis:entry colname="col8">0.14</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.51</italic></oasis:entry>  
         <oasis:entry colname="col10">0.011</oasis:entry>  
         <oasis:entry colname="col11">0.004</oasis:entry>  
         <oasis:entry colname="col12">0.009</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">27.11</oasis:entry>  
         <oasis:entry colname="col5">8.62</oasis:entry>  
         <oasis:entry colname="col6">27.73</oasis:entry>  
         <oasis:entry colname="col7">28.68</oasis:entry>  
         <oasis:entry colname="col8">0.20</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.70</italic></oasis:entry>  
         <oasis:entry colname="col10">0.018</oasis:entry>  
         <oasis:entry colname="col11">0.007</oasis:entry>  
         <oasis:entry colname="col12">0.015</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">43.99</oasis:entry>  
         <oasis:entry colname="col5">12.93</oasis:entry>  
         <oasis:entry colname="col6">50.11</oasis:entry>  
         <oasis:entry colname="col7">45.44</oasis:entry>  
         <oasis:entry colname="col8">0.25</oasis:entry>  
         <oasis:entry colname="col9">6.28</oasis:entry>  
         <oasis:entry colname="col10">0.023</oasis:entry>  
         <oasis:entry colname="col11">0.010</oasis:entry>  
         <oasis:entry colname="col12">0.019</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">8000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">15.30</oasis:entry>  
         <oasis:entry colname="col5">5.07</oasis:entry>  
         <oasis:entry colname="col6">13.78</oasis:entry>  
         <oasis:entry colname="col7">16.72</oasis:entry>  
         <oasis:entry colname="col8">0.15</oasis:entry>  
         <oasis:entry colname="col9"><italic>0.67</italic></oasis:entry>  
         <oasis:entry colname="col10">0.011</oasis:entry>  
         <oasis:entry colname="col11">0.004</oasis:entry>  
         <oasis:entry colname="col12">0.009</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"><bold>Moderate</bold></oasis:entry>  
         <oasis:entry colname="col4"><bold>28.32</bold></oasis:entry>  
         <oasis:entry colname="col5"><bold>9.29</bold></oasis:entry>  
         <oasis:entry colname="col6"><bold>26.23</bold></oasis:entry>  
         <oasis:entry colname="col7"><bold>33.99</bold></oasis:entry>  
         <oasis:entry colname="col8"><bold>0.20</bold></oasis:entry>  
         <oasis:entry colname="col9"><bold>11.4</bold></oasis:entry>  
         <oasis:entry colname="col10"><bold>0.017</bold></oasis:entry>  
         <oasis:entry colname="col11"><bold>0.008</bold></oasis:entry>  
         <oasis:entry colname="col12"><bold>0.015</bold></oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">45.92</oasis:entry>  
         <oasis:entry colname="col5">12.70</oasis:entry>  
         <oasis:entry colname="col6">53.92</oasis:entry>  
         <oasis:entry colname="col7">44.86</oasis:entry>  
         <oasis:entry colname="col8">0.25</oasis:entry>  
         <oasis:entry colname="col9">8.39</oasis:entry>  
         <oasis:entry colname="col10">0.024</oasis:entry>  
         <oasis:entry colname="col11">0.010</oasis:entry>  
         <oasis:entry colname="col12">0.020</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">16 000</oasis:entry>  
         <oasis:entry colname="col3">Small</oasis:entry>  
         <oasis:entry colname="col4">15.36</oasis:entry>  
         <oasis:entry colname="col5">5.33</oasis:entry>  
         <oasis:entry colname="col6">13.70</oasis:entry>  
         <oasis:entry colname="col7">23.18</oasis:entry>  
         <oasis:entry colname="col8">0.15</oasis:entry>  
         <oasis:entry colname="col9"><italic>2.13</italic></oasis:entry>  
         <oasis:entry colname="col10">0.011</oasis:entry>  
         <oasis:entry colname="col11">0.004</oasis:entry>  
         <oasis:entry colname="col12">0.010</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Moderate</oasis:entry>  
         <oasis:entry colname="col4">28.47</oasis:entry>  
         <oasis:entry colname="col5">8.85</oasis:entry>  
         <oasis:entry colname="col6">27.86</oasis:entry>  
         <oasis:entry colname="col7">31.91</oasis:entry>  
         <oasis:entry colname="col8">0.21</oasis:entry>  
         <oasis:entry colname="col9">6.53</oasis:entry>  
         <oasis:entry colname="col10">0.017</oasis:entry>  
         <oasis:entry colname="col11">0.008</oasis:entry>  
         <oasis:entry colname="col12">0.015</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">Large</oasis:entry>  
         <oasis:entry colname="col4">44.05</oasis:entry>  
         <oasis:entry colname="col5">12.27</oasis:entry>  
         <oasis:entry colname="col6">52.21</oasis:entry>  
         <oasis:entry colname="col7">46.10</oasis:entry>  
         <oasis:entry colname="col8">0.25</oasis:entry>  
         <oasis:entry colname="col9">7.76</oasis:entry>  
         <oasis:entry colname="col10">0.024</oasis:entry>  
         <oasis:entry colname="col11">0.010</oasis:entry>  
         <oasis:entry colname="col12">0.019</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup><?xmltex \end{scaleboxenv}?></oasis:table></table-wrap>

      <p>Table 5 shows similar tables as Table 3 but for the sensitivity to the
resampling perturbation sizes. We use the similar notation of ( ; ) denoting
(particle size; resampling perturbation setting). The results show that the
filter diverges for biomass in 13 out of 18 experiments. The five experiments
that do not diverge are (4000; large), (8000; moderate) <inline-formula><mml:math id="M107" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> TEST, (8000;
large), (16 000; moderate), and (16 000; large). The (500; small) experiment is the most
unstable, with more variables and parameters showing filter divergence.
Resampling perturbations act as variance inflation in the ensemble filters
(e.g., Anderson and Anderson, 1999). It is known that variance inflation
generally stabilizes the filter, and the results obtained here seem to be
consistent. With 4000 particles or more, the parameters and state variables
except for biomass were estimated accurately, although the filter collapsed
for biomass with smaller perturbations even with large particle sizes.</p>
</sec>
</sec>
<sec id="Ch1.S5">
  <title>Real-world experiment</title>
<sec id="Ch1.S5.SS1">
  <title>Experimental settings</title>
      <p>Here, the OSSE is extended to the real world by replacing the simulated
observations with the real observations. The sensitivity results in the
previous section showed that the settings used for the TEST experiment
provided stable filter performance; therefore, we follow the TEST experiment
here with the moderate initial and resampling perturbation sizes and with
8000 particles.</p>
      <p>Since the OSSE used the actual climate forcing in 2004 to 2007, we used the
quality-controlled MODIS LAI product of MCD15A3 for those years with flagged
as “good quality”, “Terra or Aqua”, “detectors apparently fine for up to
50 %”, “significant clouds not present”, and “main method used with
or without saturation”. We took the median of the LAI observations in the
10 km radius from the study site (62<inline-formula><mml:math id="M108" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>15<inline-formula><mml:math id="M109" display="inline"><mml:msup><mml:mi/><mml:mo>′</mml:mo></mml:msup></mml:math></inline-formula>18<inline-formula><mml:math id="M110" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>′</mml:mo><mml:mo>′</mml:mo></mml:mrow></mml:msup></mml:math></inline-formula> N,
129<inline-formula><mml:math id="M111" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>14<inline-formula><mml:math id="M112" display="inline"><mml:msup><mml:mi/><mml:mo>′</mml:mo></mml:msup></mml:math></inline-formula>29<inline-formula><mml:math id="M113" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>′</mml:mo><mml:mo>′</mml:mo></mml:mrow></mml:msup></mml:math></inline-formula> E). There are a number of missing data in the
quality-controlled MODIS data. Therefore, if the number of the data in the
10 km radius is less than 300, we set these data as the missing data for DA.
Since the MODIS data resolution is 1 km, the 10 km radius area contains
about 314 data. The observation error standard deviations are assigned to
each LAI datum in the original MODIS product (Knyazikhin et al., 1999). We
rely on the estimate of the observation error standard deviations and take
the median of the error standard deviations in the same way as getting the
LAI data. The observation error standard deviation is used in the particle
filter when computing the likelihood function (Eq. 1).</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F7" specific-use="star"><caption><p>Similar to Fig. 3, showing LAI for <bold>(a)</bold> tree plus grass, <bold>(b)</bold> tree, and
<bold>(c)</bold> grass but for the real-world experiment. Red dots with error bars
indicate the observations and their error standard deviations.</p></caption>
          <?xmltex \igopts{width=389.802756pt}?><graphic xlink:href="https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017-f07.pdf"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F8" specific-use="star"><caption><p>Similar to Fig. 4, showing the model parameters: <bold>(a)</bold> Pmax for tree,
<bold>(b)</bold> Pmax for grass, <bold>(c)</bold> Dor for tree, and <bold>(d)</bold> Dor for grass but for the real-world
experiment.</p></caption>
          <?xmltex \igopts{width=389.802756pt}?><graphic xlink:href="https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017-f08.pdf"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F9" specific-use="star"><caption><p>Similar to Fig. 5, showing the unobserved model variables: <bold>(a)</bold> GPP,
<bold>(b)</bold> RE, <bold>(c)</bold> NEE,  and <bold>(d)</bold> biomass but for the real-world experiment. Red lines
indicate the direct field observations made instantaneously every 30 min
at the AsiaFlux site, while the model simulates only the daily averages.</p></caption>
          <?xmltex \igopts{width=389.802756pt}?><graphic xlink:href="https://npg.copernicus.org/articles/24/553/2017/npg-24-553-2017-f09.pdf"/>

        </fig>

      <p>The model-simulated NEE was validated with the field observation data at this
AsiaFlux site (Ohta et al., 2001, 2008, 2014). The data were quality
controlled by the steady-state test as indicated by the quality flag 0.
Although the model simulates daily-average NEE, the field observation data
represent instantaneous NEE every 30 min. The observation data are missing
frequently, and it is not trivial to derive daily averages. Therefore, the
raw data are compared with the DA results directly. This allows only a rough
verification about whether or not the simulated NEE is in a reasonable range,
but this is the only possible verification with an independent source.</p>
</sec>
<sec id="Ch1.S5.SS2">
  <title>Results</title>
      <p>Figures 7, 8, and 9 show similar time series to Figs. 3, 4, and 5,
respectively, but with the real MODIS LAI observations. Although the
particles of the NODA experiments are widely spread, DA makes the particles
much narrower (right) for all variables and parameters. With DA, the median
of LAI is very close to the observations, within the range of the observation
error standard deviations (Fig. 7a). The grass and tree LAIs are estimated
separately (Fig. 7b, c), but there is no direct observation or other
verification truth to compare with. This is similar to the model parameters
(Fig. 8) and other model variables (Fig. 9) except for NEE, for which direct
field observation data are available. As in the OSSE results, the range of
uncertainties for NEE is reduced significantly by DA (Fig. 9c). Since the
field observations are made instantaneously every 30 min, the observation
values (red) appear to have a wider range. However, the SEIB-DGVM simulates
only daily-average NEE, and it is not straightforward to compare the outputs
from SEIB-DGVM with the field observations. We still find that the median of
NEE becomes closer to the observations, particularly near the dormancy
period. The simulated NEE generally stays within the reasonable range
compared with the field observations. In general, the particle filter shows
promising results with the real MODIS LAI data.</p>
</sec>
</sec>
<sec id="Ch1.S6" sec-type="conclusions">
  <title>Conclusion</title>
      <p>We assimilated the satellite-based MODIS LAI data using a non-Gaussian
ensemble DA system with the SEIB-DGVM based on the SIR particle filter
approach. To the best of the authors' knowledge, this is the first study to
assimilate the fine timescale satellite data with an
individual-based DGVM. We found that DA performed generally well
both for the OSSE and real-world experiments. Although we assimilated only LAI as
a whole, the tree and grass LAIs were estimated separately. This suggests
that the satellite-based DA reduced the uncertainties in the initial
vegetation structure of the individual-based DGVM toward the
simulation of future vegetation change. Another notable result includes
that the model parameters of the individual-based DGVM were
estimated successfully and that the uncertainties in the unobserved model
variables relevant to carbon cycle and vegetation states were also reduced
significantly. Similarly to the previous studies with a static
vegetation model (Stöckli et al., 2011) and a
non-individual-based DGVM (Demarty et al., 2007), the results in the present
study also suggest that LAI is the key to DA for phenology and carbon
dynamics.</p>
      <p>Generally, particle filters do not work well in high-dimensional problems
(e.g., Bickel et al., 2008; Snyder et al., 2008, 2015; Snyder, 2012). The SEIB-DGVM
has several thousand state variables, but we applied random perturbations to
only four model parameters in the particle filter. The four model parameters,
i.e., Pmax and Dor for tree and grass, control the leaf season and
photosynthesis rate of the forest as a whole. Therefore, the effective
degrees of freedom of the estimation problem would be substantially lower
than the number of variables of the SEIB-DGVM. This may be why the particle
filter worked well in this study.</p>
      <p>Additional sensitivity experiments revealed general robustness but some
sensitivities to the nature run, initial and resampling perturbation sizes,
and particle size, particularly for biomass, which tends to show particle
degeneracy. When resampling, the random perturbations were applied only to
Pmax and Dor, and not to model state variables. This contributes to reduce the
variety of vegetation structures such as tree densities and tree heights due
to the frequent DA every 4 days (not shown). This tends to cause particle
degeneracy for biomass even with large particle sizes when the resampling
perturbation size is small (Table 5). When the resampling perturbation size
is relatively large, degeneracy of the vegetation structure is mitigated to
some extent. Therefore, in this study, we tuned the resampling perturbation
sizes to avoid the filter collapse for biomass, and found that the
“moderate” perturbation size with 8000 particles is a reasonable choice.
However, the moderate perturbation size may be large for variables other than
biomass, and this may be why the estimated parameters show occasional jumps.
Adding resampling perturbations to other variables in addition to Pmax and
Dor would be better. Also, since resampling perturbations affect the particle
spread strongly, the DA technique does not necessarily provide accurate
estimates of the errors. In future studies, we will explore more effective
resampling methods to avoid the filter collapse for biomass and to represent
error estimates more accurately.</p>
      <p>As a potential limitation, it is important to note that we have made strong
assumptions in OSSE. For example, the only source of model imperfections was
the model parameter uncertainties of the four parameters. It was also
assumed that the observation error statistics were perfectly known. These
conditions would have never been met in the real-world experiment.</p>
      <p>As the first step, this study focused on the methodological development of
the data assimilation system with SEIB-DGVM and estimated only four
parameters of two PFTs using LAI observations at a single location. As a
next step, more parameters and distributions of more diverse PFTs should be
considered at different locations. Local-scale experiments can be performed
in parallel for different locations since the satellite-based LAI
observations are available globally. The simulation with the initial states
and parameter sets obtained from the SEIB-DGVM-based DA system would be
expected to improve the estimates of the carbon cycle changes over the
globe.</p>
</sec>

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

      <p>All data and source codes are archived in RIKEN Advanced
Institute for Computational Science and available upon request from the
corresponding authors under the license of the original providers. The
AsiaFlux data cannot be redistributed and are available from the AsiaFlux
database (<uri>https://db.cger.nies.go.jp/asiafluxdb/</uri>). The original source
code of the SEIB-DGVM is available at <uri>http://seib-dgvm.com/</uri> from the
developer, Hisashi Sato.</p>
  </notes><?xmltex \hack{\clearpage}?><app-group>

<app id="App1.Ch1.S1">
  <title/>

<?xmltex \floatpos{h!}?><table-wrap id="App1.Ch1.T1"><?xmltex \hack{\hsize\textwidth}?><caption><p>List of modifications to SEIB-DGVM ver.2.71.</p></caption><oasis:table frame="topbot"><?xmltex \begin{scaleboxenv}{.95}[.95]?><oasis:tgroup cols="2">
     <oasis:colspec colnum="1" colname="col1" align="justify" colwidth="398.338583pt"/>
     <oasis:colspec colnum="2" colname="col2" align="justify" colwidth="96.73937pt"/>
     <oasis:thead>
       <oasis:row rowsep="1">  
         <oasis:entry namest="col1" nameend="col2" align="left">Modification to main.f90 </oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">SUBROUTINE main_loop </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">  Initialize variables: parameters (Pmax for tree and grass, Dor for tree and grass) are read here. </oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry namest="col1" nameend="col2" align="left">  Wild fire subroutines: fire function was excluded. </oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry namest="col1" nameend="col2" align="left">Modification to metabolic.f90 </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">SUBROUTINE photosynthesis_condition: </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">  ce_water (no dimension, the minimum value): limitation on photosynthesis via soil water:  </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">  <inline-formula><mml:math id="M116" display="inline"><mml:mi>x</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M117" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> min (1.0, max (0.001, stat_water(<inline-formula><mml:math id="M118" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>))) <inline-formula><mml:math id="M119" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M120" display="inline"><mml:mi>x</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M121" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> min (1.0, max (0.1, stat_water(<inline-formula><mml:math id="M122" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>))) </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">SUBROUTINE leaf_season: </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">  Days_leaf_shed (days): day length required for full leaf drop <inline-formula><mml:math id="M123" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> from 14 to 30 </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">  Days_release_larch (days): days required for full release of stock energy for larch <inline-formula><mml:math id="M124" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> from 7 to 60 </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">  Checker (foliage <inline-formula><mml:math id="M125" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> dormancy) </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">     case (1): if (<inline-formula><mml:math id="M126" display="inline"><mml:mi>x</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M127" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 7.0) flag(<inline-formula><mml:math id="M128" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M129" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> .true. <inline-formula><mml:math id="M130" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> if (DOY <inline-formula><mml:math id="M131" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M132" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> Dor_f) flag(<inline-formula><mml:math id="M133" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M134" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> .true. (DOY: day of the year, Dor_f: Dor for tree) </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">     case (5:6): if (<inline-formula><mml:math id="M135" display="inline"><mml:mi>y</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M136" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 0.01) flag(<inline-formula><mml:math id="M137" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M138" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> .false. <inline-formula><mml:math id="M139" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> if (DOY <inline-formula><mml:math id="M140" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M141" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> Dor_g) flag(<inline-formula><mml:math id="M142" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M143" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> .true. (DOY: day of the year, Dor_g: Dor for grass) </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">     If (dfl_leaf_onset(<inline-formula><mml:math id="M144" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M145" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> Days_foliation_min) flag(<inline-formula><mml:math id="M146" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M147" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> .false. <inline-formula><mml:math id="M148" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> comment out </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">  Checker (dormancy <inline-formula><mml:math id="M149" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> foliage) </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">     case (1): if (<inline-formula><mml:math id="M150" display="inline"><mml:mi>x</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M151" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M152" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 65.0) flag(<inline-formula><mml:math id="M153" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M154" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> .true. <inline-formula><mml:math id="M155" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> if (DOY <inline-formula><mml:math id="M156" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M157" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 110) flag(<inline-formula><mml:math id="M158" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M159" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> .true (DOY: day of the year) </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">     case (5:6): if (<inline-formula><mml:math id="M160" display="inline"><mml:mi>y</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M161" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M162" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 0.01) flag(<inline-formula><mml:math id="M163" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M164" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> .false. <inline-formula><mml:math id="M165" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> if (DOY <inline-formula><mml:math id="M166" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M167" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 110) flag(<inline-formula><mml:math id="M168" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M169" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> .true (DOY: day of the year) </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">  Gradual release of stock energy: (for bug fix) </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">     IF (dfl_leaf_onset(<inline-formula><mml:math id="M170" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M171" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M172" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> day_length_release) cycle <inline-formula><mml:math id="M173" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> IF (dfl_leaf_onset(<inline-formula><mml:math id="M174" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M175" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M176" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> (day_length_release-1)) cycle </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">SUBROUTINE maintenance_resp: </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">  Herbaceous PFT source 1: (for bug fix<inline-formula><mml:math id="M177" display="inline"><mml:msup><mml:mi/><mml:mtext>a</mml:mtext></mml:msup></mml:math></inline-formula>)  </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">    mass_combust <inline-formula><mml:math id="M178" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> mass_combust <inline-formula><mml:math id="M179" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> mass_required <inline-formula><mml:math id="M180" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> mass_combust <inline-formula><mml:math id="M181" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> mass_combust <inline-formula><mml:math id="M182" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> mass_required <inline-formula><mml:math id="M183" display="inline"><mml:mo>⋅</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M184" display="inline"><mml:mi>x</mml:mi></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">    npp(<inline-formula><mml:math id="M185" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M186" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> npp(<inline-formula><mml:math id="M187" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M188" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula> mass_required <inline-formula><mml:math id="M189" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> npp(<inline-formula><mml:math id="M190" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M191" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> npp(<inline-formula><mml:math id="M192" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula>) <inline-formula><mml:math id="M193" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula> mass_required <inline-formula><mml:math id="M194" display="inline"><mml:mo>⋅</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M195" display="inline"><mml:mi>x</mml:mi></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry namest="col1" nameend="col2" align="left">SUBROUTINE growth_wood: </oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry namest="col1" nameend="col2" align="left"> Delay_from_foliation (days): delay of stem growth and reproduction process after foliation <inline-formula><mml:math id="M196" display="inline"><mml:mo>→</mml:mo></mml:math></inline-formula> from 21 to 0<inline-formula><mml:math id="M197" display="inline"><mml:msup><mml:mi/><mml:mtext>a</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry namest="col1" nameend="col2" align="left">Modification to parameter.txt </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">TO_f (times yr<inline-formula><mml:math id="M198" 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>): turn over time for foliage (grass)</oasis:entry>  
         <oasis:entry colname="col2">from 0.50 to 3.19<inline-formula><mml:math id="M199" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">TO_r (times yr<inline-formula><mml:math id="M200" 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>): turn over time for root (tree). We set the same value as the other boreal tree PFTs.</oasis:entry>  
         <oasis:entry colname="col2">from 0.16 to 0.42</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">ALM1 (m<inline-formula><mml:math id="M201" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:math></inline-formula> m<inline-formula><mml:math id="M202" 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>): allometry index of LA vs. dbh of sapwood (tree)</oasis:entry>  
         <oasis:entry colname="col2">from 6000 to 0<inline-formula><mml:math id="M203" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">ALM3 (g dm m<inline-formula><mml:math id="M204" display="inline"><mml:mrow><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>: allometry index of trunk mass (tree)</oasis:entry>  
         <oasis:entry colname="col2">from 0 to 700 000<inline-formula><mml:math id="M205" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">FR_ratio (g dm g dm<inline-formula><mml:math id="M206" 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>): ratio of leaf mass vs. root mass (tree)</oasis:entry>  
         <oasis:entry colname="col2">from 0.17 to 0.35<inline-formula><mml:math id="M207" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">FR_ratio (g dm g dm<inline-formula><mml:math id="M208" 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>): ratio of leaf mass vs. root mass (grass)</oasis:entry>  
         <oasis:entry colname="col2">from 0.33 to 0.10<inline-formula><mml:math id="M209" display="inline"><mml:msup><mml:mi/><mml:mtext>a</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">SLA (one sided m<inline-formula><mml:math id="M210" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:math></inline-formula> g dm<inline-formula><mml:math id="M211" 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>): specific leaf area (tree)</oasis:entry>  
         <oasis:entry colname="col2">from 0.014 to 0.010<inline-formula><mml:math id="M212" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">SLA (one sided m<inline-formula><mml:math id="M213" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:math></inline-formula> g dm<inline-formula><mml:math id="M214" 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>): specific leaf area (grass)</oasis:entry>  
         <oasis:entry colname="col2">from 0.015 to 0.020<inline-formula><mml:math id="M215" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Topt0 (<inline-formula><mml:math id="M216" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C): optimum temperature (tree)</oasis:entry>  
         <oasis:entry colname="col2">from 20.0 to 21.0<inline-formula><mml:math id="M217" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Tmin (<inline-formula><mml:math id="M218" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C): minimum temperature (tree). We set the same value as the other boreal tree PFTs.</oasis:entry>  
         <oasis:entry colname="col2">from 5.0 to <inline-formula><mml:math id="M219" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>4.0</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Tmax (<inline-formula><mml:math id="M220" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C): maximum temperature (tree)</oasis:entry>  
         <oasis:entry colname="col2">from 35.0 to 38.0<inline-formula><mml:math id="M221" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">GS_b2 (no dimension): parameters of stomatal conductance (grass)</oasis:entry>  
         <oasis:entry colname="col2">from 3.0 to 5.0<inline-formula><mml:math id="M222" display="inline"><mml:msup><mml:mi/><mml:mtext>a</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">M1 (no dimension): asymptotic maximum mortality rate (tree)</oasis:entry>  
         <oasis:entry colname="col2">from 0.003 to 0.001<inline-formula><mml:math id="M223" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">TC_min (<inline-formula><mml:math id="M224" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C): minimum coldest month temperature for persisting (tree and grass)</oasis:entry>  
         <oasis:entry colname="col2">from <inline-formula><mml:math id="M225" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>1000.0 to <inline-formula><mml:math id="M226" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>45.0<inline-formula><mml:math id="M227" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">GDD_min (5 <inline-formula><mml:math id="M228" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C base): minimum degree-day sum for establishment (tree)</oasis:entry>  
         <oasis:entry colname="col2">from 350 to 250<inline-formula><mml:math id="M229" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Est_scenario: scenario for establishment for tree. Only specified woody PFT was set to establish.</oasis:entry>  
         <oasis:entry colname="col2"><inline-formula><mml:math id="M230" display="inline"><mml:msup><mml:mi/><mml:mtext>a</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Est_pft_OnOff: establish switch for tree. Only boreal deciduous needleleaf tree was set to establish.</oasis:entry>  
         <oasis:entry colname="col2"><inline-formula><mml:math id="M231" display="inline"><mml:msup><mml:mi/><mml:mtext>a</mml:mtext></mml:msup></mml:math></inline-formula></oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup><?xmltex \end{scaleboxenv}?></oasis:table><?xmltex \begin{scaleboxenv}{.95}[.95]?><table-wrap-foot><p>without footnote: modifications in this
study, <inline-formula><mml:math id="M114" display="inline"><mml:msup><mml:mi/><mml:mtext>a</mml:mtext></mml:msup></mml:math></inline-formula> H. Sato (personal communications, 2014), <inline-formula><mml:math id="M115" display="inline"><mml:msup><mml:mi/><mml:mtext>b</mml:mtext></mml:msup></mml:math></inline-formula> Sato
et al. (2016).</p></table-wrap-foot><?xmltex \end{scaleboxenv}?></table-wrap>

<?xmltex \hack{\clearpage}?>
</app>
  </app-group><notes notes-type="competinginterests">

      <p>The authors declare that they have no conflict of
interest.</p>
  </notes><ack><title>Acknowledgements</title><p>The authors thank Hisashi Sato, the main developer of SEIB-DGVM, for useful
discussions. The authors also thank reviewers Matthias Morzfeld and Malaquias Pena Mendez for their careful reviews and constructive comments that helped
improve the manuscript significantly. The source code of SEIB-DGVM and the
climate forcing data are available at <uri>http://seib-dgvm.com/</uri>. MODIS LAI
product of MCD15A3 was retrieved from the online data pool, courtesy of the
NASA Land Processes Distributed Active Archive Center (LP DAAC), USGS/Earth
Resources Observation and Science (EROS) Center, Sioux Falls, South Dakota
(<uri>https://lpdaac.usgs.gov/data_access/data_pool</uri>). Carbon flux data were
retrieved from the AsiaFlux database (<uri>https://db.cger.nies.go.jp/asiafluxdb/</uri>).<?xmltex \hack{\newline}?><?xmltex \hack{\newline}?>
Edited by: Amit Apte<?xmltex \hack{\newline}?>
Reviewed by: Matthias Morzfeld and Malaquias Pena</p></ack><ref-list>
    <title>References</title>

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  </ref-list><app-group content-type="float"><app><title/>

    </app></app-group></back>
    <!--<article-title-html>Non-Gaussian data assimilation of satellite-based leaf area index observations with an individual-based dynamic global vegetation model</article-title-html>
<abstract-html><p class="p">We developed a data assimilation system based on a particle filter
approach with the spatially explicit individual-based dynamic global
vegetation model (SEIB-DGVM). We first performed an idealized observing
system simulation experiment to evaluate the impact of assimilating the leaf
area index (LAI) data every 4 days, simulating the satellite-based LAI.
Although we assimilated only LAI as a whole, the tree and grass LAIs were
estimated separately with high accuracy. Uncertain model parameters and
other state variables were also estimated accurately. Therefore, we extended
the experiment to the real world using the real Moderate Resolution Imaging
Spectroradiometer (MODIS) LAI data and obtained promising results.</p></abstract-html>
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Kalnay, E., Kanamitsu, M., Kistler, R., Collins, W., Deaven, D., Gandin, L.,
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Ebisuzaki, W., Higgins, W., Janowiak, J., Mo, K. C., Ropelewski, C., Wang,
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Grant, J. P., Heimann, M., Hooker-Stroud, A., Houweling, S., Kato, T.,
Kattge, J., Kelley, D., Kemp, S., Koffi, E. N., Köstler, C., Mathieu,
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Sebald, C., Stacke, T., Terwisscha van Scheltinga, A., Vossbeck, M.,
Widmann, H., and Ziehn, T.: The BETHY/JSBACH Carbon Cycle Data Assimilation
System: experiences and challenges, J. Geophys. Res.-Biogeo., 118,
1414–1426, 2013.
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Kato, T., Knorr, W., Scholze, M., Veenendaal, E., Kaminski, T., Kattge, J.,
and Gobron, N.: Simultaneous assimilation of satellite and eddy covariance
data for improving terrestrial water and carbon simulations at a semi-arid
woodland site in Botswana, Biogeosciences, 10, 789–802,
<a href="https://doi.org/10.5194/bg-10-789-2013" target="_blank">https://doi.org/10.5194/bg-10-789-2013</a>, 2013.
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Knyazikhin, Y., Glassy, J., Privette, J. L., Tian, Y., Lotsch, A., Zhang, Y.,
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Running, S. W.: MODIS Leaf Area Index (LAI) and Fraction of
Photosynthetically Active Radiation Absorbed by Vegetation (FPAR) product
(MOD15) Algorithm, Theoretical Basis Document, NASA Goddard Space Flight
Center, Greenbelt, MD 20771, USA, 1999.
</mixed-citation></ref-html>
<ref-html id="bib1.bib15"><label>15</label><mixed-citation>
Kolari, P., Pumpanen, J., Kulmala, L., Ilvesniemi, H., Nikinmaa, E.,
Grönholm, T., and Hari, P.: Forest floor vegetation plays an important
role in photosynthetic production of boreal forests, Forest Ecol. Manag.,
211, 241–248, 2006.
</mixed-citation></ref-html>
<ref-html id="bib1.bib16"><label>16</label><mixed-citation>
Luo, Y., Ogle, K., Tucker, C., Fei, S., Gao, C., LaDeau, S., Clark, J. S.,
and Schimel, D. S.: Ecological forecasting and data assimilation in a
data-rich era, Ecol. Appl., 21, 1429–1442, 2011.
</mixed-citation></ref-html>
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Murray-Tortarolo, G., Anav, A., Friedlingstein, P., Sitch, S., Piao, S., Zhu,
Z., Poulter, B., Zaehle, S., Ahlström, A., Lomas, M., Levis, S., Viovy,
N., and Zeng, N.: Evaluation of land surface models in reproducing
satellite-derived LAI over the high-latitude Northern Hemisphere. Part I:
Uncoupled DGVMs, Remote Sens., 5, 4819–4838, 2013.
</mixed-citation></ref-html>
<ref-html id="bib1.bib18"><label>18</label><mixed-citation>
Ohta, T., Hiyama, T., Tanaka, H., Kuwada, T., Maximov, T. C., Ohata, T., and
Fukushima, Y.: Seasonal variation in the energy and water exchanges above and
below a larch forest in eastern Siberia, Hydrol. Process., 15, 1459–1476,
2001.
</mixed-citation></ref-html>
<ref-html id="bib1.bib19"><label>19</label><mixed-citation>
Ohta, T., Maximov, T. C., Dolman, A. J., Nakai, T., van der Molen, M. K.,
Kononov, A. V., Maximov, A. P., Hiyama, T., Iijima, Y., Moors, E. J., Tanaka,
H., Toba, T., and Yabuki, H.: Interannual variation of water balance and
summer evapotranspiration in an eastern Siberian larch forest over a 7-year
period (1998–2006), Agr. Forest Meteorol., 148, 1941–1953, 2008.
</mixed-citation></ref-html>
<ref-html id="bib1.bib20"><label>20</label><mixed-citation>
Ohta, T., Kotani, A., Iijima, Y., Maximov, T. C., Ito, S., Hanamura, M.,
Kononov, A. V., and Maximov, A. P.: Effects of waterlogging on water and
carbon dioxide fluxes and environmental variables in a Siberian larch forest,
1998–2011, Agr. Forest Meteorol., 188, 64–75, 2014.
</mixed-citation></ref-html>
<ref-html id="bib1.bib21"><label>21</label><mixed-citation>
Peng, C.: From static biogeographical model to dynamic global vegetation
model: a global perspective on modelling vegetation dynamics, Ecol. Model.,
135, 33–54, 2000.
</mixed-citation></ref-html>
<ref-html id="bib1.bib22"><label>22</label><mixed-citation>
Peng, C., Guiot, J., Wu, H., Jiang, H., and Luo, Y.: Integrating models with
data in ecology and palaeoecology: advances towards a model–data fusion
approach, Ecol. Lett., 14, 522–536, 2011.
</mixed-citation></ref-html>
<ref-html id="bib1.bib23"><label>23</label><mixed-citation>
Quaife, T., Lewis, P., De Kauwe, M., Williams, M., Law, B. E., Disney, M.,
and Bowyer, P.: Assimilating canopy reflectance data into an ecosystem model
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</mixed-citation></ref-html>
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Dragoni, D., Garrity, S. R., Gough, C. M., Grant, R., Hollinger, D. Y.,
Margolis, H. A., McCaughey, H., Migliavacca, M., Monson, R. K., Munger, J.
W., Poulter, B., Raczka, B. M., Ricciuto, D. M., Sahoo, A. K., Schaefer, K.,
Tian, H., Vargas, R., Verbeeck, H., Xiao, J., and Xue, Y.: Terrestrial
biosphere models need better representation of vegetation phenology: results
from the North American Carbon Program Site Synthesis, Glob. Change Biol.,
18, 566–584, 2012.
</mixed-citation></ref-html>
<ref-html id="bib1.bib25"><label>25</label><mixed-citation>
Sato, H. and Ise, T.: Effect of plant dynamic processes on African vegetation
responses to climate change: Analysis using the spatially explicit
individual-based dynamic global vegetation model (SEIB-DGVM), J. Geophys.
Res., 117, G03017, <a href="https://doi.org/10.1029/2012JG002056" target="_blank">https://doi.org/10.1029/2012JG002056</a>, 2012.
</mixed-citation></ref-html>
<ref-html id="bib1.bib26"><label>26</label><mixed-citation>
Sato, H., Itoh, A., and Kohyama, T.: SEIB–DGVM: A new Dynamic Global
Vegetation Model using a spatially explicit individual-based approach, Ecol.
Model., 200, 279–307, 2007.
</mixed-citation></ref-html>
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