- Author:
- Catherine Lloyd <c.lloyd@auckland.ac.nz>
- Date:
- 2010-05-06 09:50:53+12:00
- Desc:
- Added "place holder" for image ready for Hanne to create image.
- Permanent Source URI:
- http://models.cellml.org/workspace/ralser_wamelink_kowald_gerisch_heeren_struys_klipp_jakobs_breitenbach_lehrach_krobitsch_2007/rawfile/50e5134bd3fea1741bcb75285ca6b053e6d013fc/ralser_2007.cellml
<?xml version="1.0" encoding="utf-8"?>
<!--
This CellML file was generated on 27/04/2010 at 11:06:52 at a.m. using:
COR (0.9.31.1371)
Copyright 2002-2010 Dr Alan Garny
http://cor.physiol.ox.ac.uk/ - cor@physiol.ox.ac.uk
CellML 1.0 was used to generate this model
http://www.cellml.org/
-->
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<documentation xmlns="http://cellml.org/tmp-documentation">
<article>
<articleinfo>
<title>Dynamic rerouting of the carbohydrate flux is key to counteracting oxidative stress</title>
<author>
<firstname>Geoffrey</firstname>
<surname>Nunns</surname>
<affiliation>
<shortaffil>Bioengineering Institute, University of Auckland</shortaffil>
</affiliation>
</author>
</articleinfo>
<section id="sec_status">
<title>Model Status</title>
<para>
This CellML model runs in COR and OpenCell and the units are consistent throughout. It reproduces the published results and was converted from SBML with the help of Lukas Endler.
</para>
</section>
<sect1 id="sec_structure">
<title>Model Structure</title>
<para>
ABSTRACT: BACKGROUND: Eukaryotic cells have evolved various response mechanisms to counteract the deleterious consequences of oxidative stress. Among these processes, metabolic alterations seem to play an important role. RESULTS: We recently discovered that yeast cells with reduced activity of the key glycolytic enzyme triosephosphate isomerase exhibit an increased resistance to the thiol-oxidizing reagent diamide. Here we show that this phenotype is conserved in Caenorhabditis elegans and that the underlying mechanism is based on a redirection of the metabolic flux from glycolysis to the pentose phosphate pathway, altering the redox equilibrium of the cytoplasmic NADP(H) pool. Remarkably, another key glycolytic enzyme, glyceraldehyde-3-phosphate dehydrogenase (GAPDH), is known to be inactivated in response to various oxidant treatments, and we show that this provokes a similar redirection of the metabolic flux. CONCLUSION: The naturally occurring inactivation of GAPDH functions as a metabolic switch for rerouting the carbohydrate flux to counteract oxidative stress. As a consequence, altering the homoeostasis of cytoplasmic metabolites is a fundamental mechanism for balancing the redox state of eukaryotic cells under stress conditions.
</para>
<para>
The original paper reference is cited below:
</para>
<para>
Dynamic rerouting of the carbohydrate flux is key to counteracting oxidative stress, Markus Ralser, Mirjam M Wamelink, Axel Kowald, Birgit Gerisch, Gino Heeren, Eduard A Struys, Edda Klipp, Cornelis Jakobs, Michael Breitenbach, Hans Lehrach and Sylvia Krobitsch, 2007, <emphasis>Journal of Biology</emphasis>, volume 6, 1-18. <ulink url="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=pubmed&cmd=Retrieve&dopt=AbstractPlus&list_uids=18154684&query_hl=1&itool=pubmed_docsum">PubMed ID: 18154684</ulink>
</para>
<informalfigure float="0" id="fig_reaction_diagram">
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<title>model diagram</title>
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<imagedata fileref="ralser_2007.png"/>
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<caption>Schematic diagram of the reactions included in the mathematical model to study the effects of a diminished TPI or GAPDH activity on the flux through glycolysis and the pentose phosphate pathway. Substances with a dashed outline are assumed to have a constant concentration</caption>
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</sect1>
</article>
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<ci>KmGLKGLCi</ci>
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<ci>KmPGIF6P</ci>
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<variable name="numerator1" units="mmol_mM2_per_minute"/>
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<times/>
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<minus/>
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<times/>
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<power/>
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<plus/>
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<power/>
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</apply>
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<apply>
<root/>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<minus/>
<apply>
<power/>
<ci>SUMAXP</ci>
<cn cellml:units="dimensionless">2</cn>
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<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>SUMAXP</ci>
<ci>P</ci>
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<apply>
<times/>
<cn cellml:units="dimensionless">8</cn>
<ci>KeqAK</ci>
<ci>SUMAXP</ci>
<ci>P</ci>
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<power/>
<ci>P</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
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<apply>
<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>KeqAK</ci>
<apply>
<power/>
<ci>P</ci>
<cn cellml:units="dimensionless">2</cn>
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</apply>
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<apply>
<times/>
<apply>
<minus/>
<cn cellml:units="dimensionless">2</cn>
<apply>
<times/>
<cn cellml:units="dimensionless">8</cn>
<ci>KeqAK</ci>
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</apply>
<ci>KmPGKATP</ci>
</apply>
</apply>
</apply>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>BPG</ci>
<ci>KmPGKBPG</ci>
</apply>
<apply>
<divide/>
<ci>P3G</ci>
<ci>KmPGKP3G</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="vPGM">
<variable name="vPGM" units="mmol_per_min" public_interface="out"/>
<variable name="VmPGM" units="mM_per_min" initial_value="2525.81"/>
<variable name="KmPGMP3G" units="mM" initial_value="1.2"/>
<variable name="KeqPGM" units="dimensionless" initial_value="0.19"/>
<variable name="KmPGMP2G" units="mM" initial_value="0.08"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="P2G" units="mM" public_interface="in"/>
<variable name="P3G" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>vPGM</ci>
<apply>
<divide/>
<apply>
<times/>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
<ci>VmPGM</ci>
</apply>
<ci>KmPGMP3G</ci>
</apply>
<apply>
<minus/>
<ci>P3G</ci>
<apply>
<divide/>
<ci>P2G</ci>
<ci>KeqPGM</ci>
</apply>
</apply>
</apply>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>P3G</ci>
<ci>KmPGMP3G</ci>
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<apply>
<divide/>
<ci>P2G</ci>
<ci>KmPGMP2G</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="vENO">
<variable name="vENO" units="mmol_per_min" public_interface="out"/>
<variable name="VmENO" units="mM_per_min" initial_value="365.806"/>
<variable name="KmENOP2G" units="mM" initial_value="0.04"/>
<variable name="KeqENO" units="dimensionless" initial_value="6.7"/>
<variable name="KmENOPEP" units="mM" initial_value="0.5"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="P2G" units="mM" public_interface="in"/>
<variable name="PEP" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>vENO</ci>
<apply>
<divide/>
<apply>
<times/>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
<ci>VmENO</ci>
</apply>
<ci>KmENOP2G</ci>
</apply>
<apply>
<minus/>
<ci>P2G</ci>
<apply>
<divide/>
<ci>PEP</ci>
<ci>KeqENO</ci>
</apply>
</apply>
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<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>P2G</ci>
<ci>KmENOP2G</ci>
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<apply>
<divide/>
<ci>PEP</ci>
<ci>KmENOPEP</ci>
</apply>
</apply>
</apply>
</apply>
</math>
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<component xmlns="http://www.cellml.org/cellml/1.0#" name="vPYK">
<variable name="vPYK" units="mmol_per_min" public_interface="out"/>
<variable name="VmPYK" units="mM_per_min" initial_value="1088.71"/>
<variable name="KmPYKPEP" units="mM" initial_value="0.14"/>
<variable name="KmPYKADP" units="mM" initial_value="0.53"/>
<variable name="KeqAK" units="dimensionless" initial_value="0.45"/>
<variable name="KeqPYK" units="dimensionless" initial_value="6500"/>
<variable name="KmPYKPYR" units="mM" initial_value="21"/>
<variable name="KmPYKATP" units="mM" initial_value="1.5"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="PEP" units="mM" public_interface="in"/>
<variable name="P" units="mM" public_interface="in"/>
<variable name="PYR" units="mM" public_interface="in"/>
<variable name="SUMAXP" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>vPYK</ci>
<apply>
<divide/>
<apply>
<times/>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
<ci>VmPYK</ci>
</apply>
<apply>
<times/>
<ci>KmPYKPEP</ci>
<ci>KmPYKADP</ci>
</apply>
</apply>
<apply>
<minus/>
<apply>
<divide/>
<apply>
<times/>
<ci>PEP</ci>
<apply>
<minus/>
<ci>SUMAXP</ci>
<apply>
<root/>
<apply>
<plus/>
<apply>
<minus/>
<apply>
<minus/>
<apply>
<power/>
<ci>P</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>KeqAK</ci>
<apply>
<power/>
<ci>P</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>P</ci>
<ci>SUMAXP</ci>
</apply>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">8</cn>
<ci>KeqAK</ci>
<ci>P</ci>
<ci>SUMAXP</ci>
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<apply>
<power/>
<ci>SUMAXP</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
</apply>
</apply>
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<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>KeqAK</ci>
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</apply>
</apply>
<apply>
<divide/>
<apply>
<divide/>
<apply>
<times/>
<ci>PYR</ci>
<apply>
<plus/>
<apply>
<minus/>
<apply>
<minus/>
<ci>P</ci>
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<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>KeqAK</ci>
<ci>P</ci>
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<ci>SUMAXP</ci>
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<apply>
<root/>
<apply>
<plus/>
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<minus/>
<apply>
<minus/>
<apply>
<power/>
<ci>P</ci>
<cn cellml:units="dimensionless">2</cn>
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<apply>
<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>KeqAK</ci>
<apply>
<power/>
<ci>P</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
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<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>P</ci>
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<apply>
<times/>
<cn cellml:units="dimensionless">8</cn>
<ci>KeqAK</ci>
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<power/>
<ci>SUMAXP</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
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<apply>
<minus/>
<cn cellml:units="dimensionless">2</cn>
<apply>
<times/>
<cn cellml:units="dimensionless">8</cn>
<ci>KeqAK</ci>
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<ci>KeqPYK</ci>
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<apply>
<times/>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>PEP</ci>
<ci>KmPYKPEP</ci>
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<apply>
<divide/>
<ci>PYR</ci>
<ci>KmPYKPYR</ci>
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<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
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<divide/>
<apply>
<plus/>
<apply>
<minus/>
<apply>
<minus/>
<ci>P</ci>
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<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>KeqAK</ci>
<ci>P</ci>
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<ci>SUMAXP</ci>
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<apply>
<root/>
<apply>
<plus/>
<apply>
<minus/>
<apply>
<minus/>
<apply>
<power/>
<ci>P</ci>
<cn cellml:units="dimensionless">2</cn>
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<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>KeqAK</ci>
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<power/>
<ci>P</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
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<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>P</ci>
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<times/>
<cn cellml:units="dimensionless">8</cn>
<ci>KeqAK</ci>
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<power/>
<ci>SUMAXP</ci>
<cn cellml:units="dimensionless">2</cn>
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<apply>
<minus/>
<cn cellml:units="dimensionless">2</cn>
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<times/>
<cn cellml:units="dimensionless">8</cn>
<ci>KeqAK</ci>
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<ci>KmPYKATP</ci>
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<divide/>
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<divide/>
<apply>
<minus/>
<ci>SUMAXP</ci>
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<root/>
<apply>
<plus/>
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<minus/>
<apply>
<minus/>
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<power/>
<ci>P</ci>
<cn cellml:units="dimensionless">2</cn>
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<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>KeqAK</ci>
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<power/>
<ci>P</ci>
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<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>P</ci>
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<times/>
<cn cellml:units="dimensionless">8</cn>
<ci>KeqAK</ci>
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<power/>
<ci>SUMAXP</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
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<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
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<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>KeqAK</ci>
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<ci>KmPYKADP</ci>
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</apply>
</apply>
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<variable name="vPDC" units="mmol_per_min" public_interface="out"/>
<variable name="VmPDC" units="mM_per_min" initial_value="174.194"/>
<variable name="nPDC" units="dimensionless" initial_value="1.9"/>
<variable name="KmPDCPYR" units="mM" initial_value="4.33"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="PYR" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
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<eq/>
<ci>vPDC</ci>
<apply>
<divide/>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
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<power/>
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<power/>
<ci>KmPDCPYR</ci>
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<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<apply>
<power/>
<ci>PYR</ci>
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<power/>
<ci>KmPDCPYR</ci>
<ci>nPDC</ci>
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<variable name="vSUC" units="mmol_per_min" public_interface="out"/>
<variable name="KSUCC" units="per_min" initial_value="21.4"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="ACE" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>vSUC</ci>
<apply>
<times/>
<ci>cytoplasm</ci>
<ci>KSUCC</ci>
<ci>ACE</ci>
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<variable name="vADH" units="mmol_per_min" public_interface="out"/>
<variable name="VmADH" units="mM_per_min" initial_value="810"/>
<variable name="KiADHNAD" units="mM" initial_value="0.92"/>
<variable name="KmADHETOH" units="mM" initial_value="17"/>
<variable name="KeqADH" units="dimensionless" initial_value="6.9e-5"/>
<variable name="KmADHNAD" units="mM" initial_value="0.17"/>
<variable name="KmADHNADH" units="mM" initial_value="0.11"/>
<variable name="KiADHNADH" units="mM" initial_value="0.031"/>
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<variable name="KiADHACE" units="mM" initial_value="1.1"/>
<variable name="KiADHETOH" units="mM" initial_value="90"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="ACE" units="mM" public_interface="in"/>
<variable name="ETOH" units="mM" public_interface="in"/>
<variable name="NAD" units="mM" public_interface="in"/>
<variable name="NADH" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>vADH</ci>
<apply>
<divide/>
<apply>
<times/>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
<apply>
<minus/>
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<apply>
<times/>
<ci>KiADHNAD</ci>
<ci>KmADHETOH</ci>
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</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>NAD</ci>
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<apply>
<divide/>
<apply>
<times/>
<ci>NADH</ci>
<ci>ACE</ci>
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<ci>KeqADH</ci>
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</apply>
</apply>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>NAD</ci>
<ci>KiADHNAD</ci>
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<apply>
<divide/>
<apply>
<times/>
<ci>KmADHNAD</ci>
<ci>ETOH</ci>
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<apply>
<times/>
<ci>KiADHNAD</ci>
<ci>KmADHETOH</ci>
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<apply>
<divide/>
<apply>
<times/>
<ci>KmADHNADH</ci>
<ci>ACE</ci>
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<apply>
<times/>
<ci>KiADHNADH</ci>
<ci>KmADHACE</ci>
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<apply>
<divide/>
<ci>NADH</ci>
<ci>KiADHNADH</ci>
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<apply>
<divide/>
<apply>
<times/>
<ci>NAD</ci>
<ci>ETOH</ci>
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<apply>
<times/>
<ci>KiADHNAD</ci>
<ci>KmADHETOH</ci>
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<divide/>
<apply>
<times/>
<ci>KmADHNADH</ci>
<ci>NAD</ci>
<ci>ACE</ci>
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<times/>
<ci>KiADHNAD</ci>
<ci>KiADHNADH</ci>
<ci>KmADHACE</ci>
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<apply>
<divide/>
<apply>
<times/>
<ci>KmADHNAD</ci>
<ci>ETOH</ci>
<ci>NADH</ci>
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<times/>
<ci>KiADHNAD</ci>
<ci>KmADHETOH</ci>
<ci>KiADHNADH</ci>
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<apply>
<divide/>
<apply>
<times/>
<ci>NADH</ci>
<ci>ACE</ci>
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<times/>
<ci>KiADHNADH</ci>
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<apply>
<divide/>
<apply>
<times/>
<ci>NAD</ci>
<ci>ETOH</ci>
<ci>ACE</ci>
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<apply>
<times/>
<ci>KiADHNAD</ci>
<ci>KmADHETOH</ci>
<ci>KiADHACE</ci>
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<apply>
<divide/>
<apply>
<times/>
<ci>ETOH</ci>
<ci>NADH</ci>
<ci>ACE</ci>
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<apply>
<times/>
<ci>KiADHETOH</ci>
<ci>KiADHNADH</ci>
<ci>KmADHACE</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
</math>
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<component xmlns="http://www.cellml.org/cellml/1.0#" name="vATP">
<variable name="vATP" units="mmol_per_min" public_interface="out"/>
<variable name="KATPASE" units="per_min" initial_value="39.5"/>
<variable name="KeqAK" units="dimensionless" initial_value="0.45"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="P" units="mM" public_interface="in"/>
<variable name="SUMAXP" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
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<eq/>
<ci>vATP</ci>
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<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
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<plus/>
<apply>
<minus/>
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<minus/>
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<times/>
<cn cellml:units="dimensionless">4</cn>
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<root/>
<apply>
<plus/>
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<minus/>
<apply>
<minus/>
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<power/>
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<times/>
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<ci>KeqAK</ci>
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<power/>
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<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>P</ci>
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<apply>
<times/>
<cn cellml:units="dimensionless">8</cn>
<ci>KeqAK</ci>
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<power/>
<ci>SUMAXP</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
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<apply>
<minus/>
<cn cellml:units="dimensionless">2</cn>
<apply>
<times/>
<cn cellml:units="dimensionless">8</cn>
<ci>KeqAK</ci>
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</apply>
</math>
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<variable name="vTPI" units="mmol_per_min" public_interface="out"/>
<variable name="KmGA3P" units="mM" initial_value="1.27"/>
<variable name="KmDHAP" units="mM" initial_value="1.23"/>
<variable name="VmDHAP" units="mM_per_min" initial_value="10900"/>
<variable name="VmGA3P" units="mM_per_min" initial_value="555"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="DHAP" units="mM" public_interface="in"/>
<variable name="GA3P" units="mM" public_interface="in"/>
<variable name="k_rel_TPI" units="dimensionless" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>vTPI</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
<ci>k_rel_TPI</ci>
<apply>
<minus/>
<apply>
<divide/>
<apply>
<times/>
<ci>VmDHAP</ci>
<ci>GA3P</ci>
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<ci>KmGA3P</ci>
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<apply>
<divide/>
<apply>
<times/>
<ci>VmGA3P</ci>
<ci>DHAP</ci>
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<ci>KmDHAP</ci>
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</apply>
</apply>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
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<divide/>
<ci>GA3P</ci>
<ci>KmGA3P</ci>
</apply>
<apply>
<divide/>
<ci>DHAP</ci>
<ci>KmDHAP</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="vG6PDH">
<variable name="vG6PDH" units="mmol_per_min" public_interface="out"/>
<variable name="VmG6PDH" units="mM_per_min" initial_value="4"/>
<variable name="KmG6P" units="mM" initial_value="0.04"/>
<variable name="KmNADP" units="mM" initial_value="0.02"/>
<variable name="KiNADPH" units="mM" initial_value="0.017"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="G6P" units="mM" public_interface="in"/>
<variable name="NADP" units="mM" public_interface="in"/>
<variable name="NADPH" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>vG6PDH</ci>
<apply>
<divide/>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
<ci>VmG6PDH</ci>
<ci>G6P</ci>
<ci>NADP</ci>
</apply>
<apply>
<times/>
<ci>KmG6P</ci>
<ci>KmNADP</ci>
</apply>
</apply>
<apply>
<times/>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>G6P</ci>
<ci>KmG6P</ci>
</apply>
<apply>
<divide/>
<ci>NADPH</ci>
<ci>KiNADPH</ci>
</apply>
</apply>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>NADP</ci>
<ci>KmNADP</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v6PGL">
<variable name="v6PGL" units="mmol_per_min" public_interface="out"/>
<variable name="Vm6PGL" units="mM_per_min" initial_value="4"/>
<variable name="Km6PGL" units="mM" initial_value="0.8"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="D6PGluconoLactone" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v6PGL</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
<ci>Vm6PGL</ci>
<ci>D6PGluconoLactone</ci>
</apply>
<apply>
<plus/>
<ci>Km6PGL</ci>
<ci>D6PGluconoLactone</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="vGluDH">
<variable name="vGluDH" units="mmol_per_min" public_interface="out"/>
<variable name="VmGluDH" units="mM_per_min" initial_value="4"/>
<variable name="KmGluconate" units="mM" initial_value="0.02"/>
<variable name="KmNADP" units="mM" initial_value="0.03"/>
<variable name="KiNADPH" units="mM" initial_value="0.03"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="D6PGluconate" units="mM" public_interface="in"/>
<variable name="NADP" units="mM" public_interface="in"/>
<variable name="NADPH" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>vGluDH</ci>
<apply>
<divide/>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
<ci>VmGluDH</ci>
<ci>D6PGluconate</ci>
<ci>NADP</ci>
</apply>
<apply>
<times/>
<ci>KmGluconate</ci>
<ci>KmNADP</ci>
</apply>
</apply>
<apply>
<times/>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>D6PGluconate</ci>
<ci>KmGluconate</ci>
</apply>
<apply>
<divide/>
<ci>NADPH</ci>
<ci>KiNADPH</ci>
</apply>
</apply>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>NADP</ci>
<ci>KmNADP</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="vPPI">
<variable name="vPPI" units="mmol_per_min" public_interface="out"/>
<variable name="VmPPIf" units="mM_per_min" initial_value="3458"/>
<variable name="VmPPIr" units="mM_per_min" initial_value="3458"/>
<variable name="KmRibu5P" units="mM" initial_value="1.6"/>
<variable name="KmRibo5P" units="mM" initial_value="1.6"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="Ribulose5P" units="mM" public_interface="in"/>
<variable name="Ribose5P" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>vPPI</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
<apply>
<minus/>
<apply>
<divide/>
<apply>
<times/>
<ci>VmPPIf</ci>
<ci>Ribulose5P</ci>
</apply>
<ci>KmRibu5P</ci>
</apply>
<apply>
<divide/>
<apply>
<times/>
<ci>VmPPIr</ci>
<ci>Ribose5P</ci>
</apply>
<ci>KmRibo5P</ci>
</apply>
</apply>
</apply>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>Ribulose5P</ci>
<ci>KmRibu5P</ci>
</apply>
<apply>
<divide/>
<ci>Ribose5P</ci>
<ci>KmRibo5P</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="vTransk1">
<variable name="vTransk1" units="mmol_per_min" public_interface="out"/>
<variable name="VmTransk1f" units="mM_per_min" initial_value="4"/>
<variable name="VmTransk1r" units="mM_per_min" initial_value="2"/>
<variable name="KmRibose5P" units="mM" initial_value="0.1"/>
<variable name="KmXyl5P" units="mM" initial_value="0.15"/>
<variable name="KmGA3P" units="mM" initial_value="0.1"/>
<variable name="KmSeduhept" units="mM" initial_value="0.15"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="GA3P" units="mM" public_interface="in"/>
<variable name="Ribose5P" units="mM" public_interface="in"/>
<variable name="Seduhept7P" units="mM" public_interface="in"/>
<variable name="Xyl5P" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>vTransk1</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
<apply>
<minus/>
<apply>
<divide/>
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<times/>
<ci>VmTransk1f</ci>
<ci>Ribose5P</ci>
<ci>Xyl5P</ci>
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<apply>
<times/>
<ci>KmRibose5P</ci>
<ci>KmXyl5P</ci>
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<apply>
<divide/>
<apply>
<times/>
<ci>VmTransk1r</ci>
<ci>GA3P</ci>
<ci>Seduhept7P</ci>
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<apply>
<times/>
<ci>KmGA3P</ci>
<ci>KmSeduhept</ci>
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</apply>
</apply>
</apply>
<apply>
<times/>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>Ribose5P</ci>
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<apply>
<divide/>
<ci>GA3P</ci>
<ci>KmGA3P</ci>
</apply>
</apply>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>Xyl5P</ci>
<ci>KmXyl5P</ci>
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<apply>
<divide/>
<ci>Seduhept7P</ci>
<ci>KmSeduhept</ci>
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</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="vR5PI">
<variable name="vR5PI" units="mmol_per_min" public_interface="out"/>
<variable name="VmR5PIf" units="mM_per_min" initial_value="1039"/>
<variable name="VmR5PIr" units="mM_per_min" initial_value="1039"/>
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<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="Ribulose5P" units="mM" public_interface="in"/>
<variable name="Xyl5P" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
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<eq/>
<ci>vR5PI</ci>
<apply>
<divide/>
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<times/>
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<minus/>
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<divide/>
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<times/>
<ci>VmR5PIf</ci>
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<divide/>
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<times/>
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<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
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<divide/>
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<divide/>
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</apply>
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<variable name="vTransald" units="mmol_per_min" public_interface="out"/>
<variable name="VmTransaldf" units="mM_per_min" initial_value="55"/>
<variable name="VmTransaldr" units="mM_per_min" initial_value="10"/>
<variable name="KmGA3P" units="mM" initial_value="0.22"/>
<variable name="KmSeduhept" units="mM" initial_value="0.18"/>
<variable name="KmF6P" units="mM" initial_value="0.32"/>
<variable name="KmEry4P" units="mM" initial_value="0.018"/>
<variable name="cytoplasm" units="liter" public_interface="in"/>
<variable name="Erythrose4P" units="mM" public_interface="in"/>
<variable name="F6P" units="mM" public_interface="in"/>
<variable name="Seduhept7P" units="mM" public_interface="in"/>
<variable name="GA3P" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
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<eq/>
<ci>vTransald</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>cytoplasm</ci>
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<minus/>
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<divide/>
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<times/>
<ci>VmTransaldf</ci>
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<times/>
<ci>KmGA3P</ci>
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<apply>
<divide/>
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<times/>
<ci>VmTransaldr</ci>
<ci>F6P</ci>
<ci>Erythrose4P</ci>
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<times/>
<ci>KmF6P</ci>
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<apply>
<times/>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
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<divide/>
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<divide/>
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<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
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<divide/>
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<variable name="VmTransk2r" units="mM_per_min" initial_value="43"/>
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<variable name="KmGA3P" units="mM" initial_value="2.1"/>
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<variable name="Erythrose4P" units="mM" public_interface="in"/>
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<variable name="Xyl5P" units="mM" public_interface="in"/>
<variable name="GA3P" units="mM" public_interface="in"/>
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<eq/>
<ci>vTransk2</ci>
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<divide/>
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<times/>
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<minus/>
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<divide/>
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<times/>
<ci>VmTransk2r</ci>
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<apply>
<times/>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
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<divide/>
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<divide/>
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<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
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<divide/>
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<divide/>
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<variable name="kNADPH" units="per_min" initial_value="2"/>
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<variable name="NADPH" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
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<eq/>
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<times/>
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<variable name="GLCo" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
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<minus/>
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<divide/>
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<plus/>
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<divide/>
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<divide/>
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<times/>
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<times/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
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<map_variables variable_1="GA3P" variable_2="GA3P"/>
<map_variables variable_1="DHAP" variable_2="DHAP"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
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<map_variables variable_1="cytoplasm" variable_2="cytoplasm"/>
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<map_variables variable_1="NADH" variable_2="NADH"/>
<map_variables variable_1="DHAP" variable_2="DHAP"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="vG3PDH" component_2="extracellular"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="vGAPDH" component_2="cytoplasm"/>
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<map_variables variable_1="cytoplasm" variable_2="cytoplasm"/>
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<map_variables variable_1="NAD" variable_2="NAD"/>
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