- Author:
- Geoff Nunns <gnunns1@jhu.edu>
- Date:
- 2010-04-12 17:08:09+12:00
- Desc:
- Added model status, updated model structure to include only abstract. Fixed circular equations so that the model can run in COR, corrected equations where different from paper. Corrected units inconsistencies and guessed initial conditions then ran model to steady state to get steady state initial conditions.
- Permanent Source URI:
- https://models.cellml.org/workspace/wolf_sohn_heinrich_kuriyama_2001/rawfile/4dbf870828c8250f44de26435b1b2be0393c7fbc/wolf_sohn_heinrich_kuriyama_2001.cellml
<?xml version="1.0" encoding="utf-8"?>
<!--
This CellML file was generated on 12/04/2010 at 4:54:42 at p.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/
-->
<model xmlns:dc="http://purl.org/dc/elements/1.1/" xmlns:rdf="http://www.w3.org/1999/02/22-rdf-syntax-ns#" xmlns:bqs="http://www.cellml.org/bqs/1.0#" xmlns:dcterms="http://purl.org/dc/terms/" xmlns:pathway_editor="http://www.physiome.com/pathway_editor/1.0#" xmlns:vCard="http://www.w3.org/2001/vcard-rdf/3.0#" name="wolf_sohn_heinrich_kuriyama_2001_version01" cmeta:id="wolf_sohn_heinrich_kuriyama_2001_version01" xmlns="http://www.cellml.org/cellml/1.0#" xmlns:cellml="http://www.cellml.org/cellml/1.0#" xmlns:cmeta="http://www.cellml.org/metadata/1.0#">
<documentation xmlns="http://cellml.org/tmp-documentation">
<article>
<articleinfo>
<title>Metabolic Oscillations in Saccharomyces cerevisiae</title>
<author>
<firstname>Catherine</firstname>
<surname>Lloyd</surname>
<affiliation>
<shortaffil>Bioengineering Institute, University of Auckland</shortaffil>
</affiliation>
</author>
</articleinfo>
<section id="sec_status">
<title>Model Status</title>
<para>
This model runs to reproduce published results in COR, OpenCell and PCEnv. The units are consistent throughout. This model currently reproduces Figure 2B, changing KH (to infinity) and f2 (to 1) will allow the model to reproduce Figure 2A. The initial conditions were estimated and the model run to steady state to yield precise values, changing any parameters will necessitate repeating this procedure.
</para>
</section>
<sect1 id="sec_structure">
<title>Model Structure</title>
<para>
ABSTRACT: Autonomous metabolic oscillations were observed in aerobic continuous culture of Saccharomyces cerevisiae. Experimental investigation of the underlying mechanism revealed that several pathways and regulatory couplings are involved. Here a hypothetical mechanism including the sulfate assimilation pathway, ethanol degradation and respiration is transformed into a mathematical model. Simulations confirm the ability of the model to produce limit cycle oscillations which reproduce most of the characteristic features of the system
</para>
<para>
The complete original paper reference is cited below:
</para>
<para>
Mathematical analysis of a mechanism for autonomous metabolic oscillations in continuous culture of <emphasis>Saccharomyces cerevisiae</emphasis>, Jana Wolf, Ho-Yong Sohn, Reinhart Heinrich and Hiroshi Kuriyama, 2001, <emphasis>FEBS Letters</emphasis>, 499, 230-234. <ulink url="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&list_uids=11423122&dopt=Abstract">PubMed ID: 11423122</ulink>
</para>
<informalfigure float="0" id="fig_cell_diagram">
<mediaobject>
<imageobject>
<objectinfo>
<title>schematic diagram</title>
</objectinfo>
<imagedata fileref="wolf_2001.png"/>
</imageobject>
</mediaobject>
<caption>Reaction scheme of the model. The following abbreviations are introduced for the metabolites: sul: sulphate ions, aps: adenylyl sulfate, pap: 3-phosphoadenylyl sulfate, hyd: hydrogen sulphide, cys: cysteine, eth: ethanol, aco: acetyl-CoA, S1 and S2: intermediates of the citric acid cycle, oxy: oxygen, C1 and C2: protein complexes involved in oxidative phosphorylation, A3: ATP, A2: ADP, N1: NAD(P)<superscript>+</superscript>, N2: NAD(P)H, oah: O-acetylhomoserine. The cytosolic and mitochondrial compartments are characterized by the superscripts c and m, respectively.</caption>
</informalfigure>
</sect1>
</article>
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total intermediates of the citrate acid cycle
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<dcterms:W3CDTF>2001-06-22</dcterms:W3CDTF>
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<vCard:Given>Jana</vCard:Given>
<vCard:Family>Wolf</vCard:Family>
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<dcterms:alternative>O-acetylhomoserine</dcterms:alternative>
<dc:title>oah</dc:title>
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<dcterms:alternative>cytosolic ATP</dcterms:alternative>
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<dcterms:alternative>mitochondrial ATP</dcterms:alternative>
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<rdf:Description rdf:about="rdf:#8f8d038c-f838-4e9b-9e50-12b75c865ad7">
<vCard:FN>Catherine Lloyd</vCard:FN>
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<rdf:value>c.lloyd@auckland.ac.nz</rdf:value>
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<dc:title>A dynamic model of metabolic oscillations in yeast</dc:title>
<cmeta:comment xmlns:cmeta="http://www.cellml.org/metadata/1.0#" rdf:resource="rdf:#8a726446-10a0-4fb5-a123-0754513befa2"/>
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<vCard:Orgname>The University of Auckland</vCard:Orgname>
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<dcterms:alternative>acetyl-CoA</dcterms:alternative>
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metabolic oscillations in the yeast Saccharomyces cerevisiae.</rdf:value>
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<variable name="v10" units="flux" public_interface="in"/>
<variable name="v11A" units="flux" public_interface="in"/>
<variable name="v14" units="flux" public_interface="in"/>
<variable name="time" units="minute" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>oxy</ci>
</apply>
<apply>
<minus/>
<ci>v10</ci>
<apply>
<plus/>
<ci>v11A</ci>
<ci>v14</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="A3_c">
<variable name="A3_c" units="millimolar" initial_value="1.570847601" public_interface="out"/>
<variable name="v2" units="flux" public_interface="in"/>
<variable name="v3" units="flux" public_interface="in"/>
<variable name="v12" units="flux" public_interface="in"/>
<variable name="v16" units="flux" public_interface="in"/>
<variable name="time" units="minute" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>A3_c</ci>
</apply>
<apply>
<minus/>
<ci>v16</ci>
<apply>
<plus/>
<ci>v2</ci>
<ci>v3</ci>
<ci>v12</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="A3_m">
<variable name="A3_m" units="millimolar" initial_value="1.835158493" public_interface="out"/>
<variable name="v11B" units="flux" public_interface="in"/>
<variable name="v16" units="flux" public_interface="in"/>
<variable name="time" units="minute" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>A3_m</ci>
</apply>
<apply>
<minus/>
<ci>v11B</ci>
<ci>v16</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="A_c">
<variable name="A_c" units="millimolar" initial_value="2"/>
<variable name="A2_c" units="millimolar" public_interface="out"/>
<variable name="A3_c" units="millimolar" public_interface="in"/>
<variable name="time" units="minute" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>A2_c</ci>
<apply>
<minus/>
<ci>A_c</ci>
<ci>A3_c</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="A_m">
<variable name="A_m" units="millimolar" initial_value="2"/>
<variable name="A2_m" units="millimolar" public_interface="out"/>
<variable name="A3_m" units="millimolar" public_interface="in"/>
<variable name="time" units="minute" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>A2_m</ci>
<apply>
<minus/>
<ci>A_m</ci>
<ci>A3_m</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="N2">
<variable name="N2" units="millimolar" initial_value="1.933616149" public_interface="out"/>
<variable name="v4" units="flux" public_interface="in"/>
<variable name="v7" units="flux" public_interface="in"/>
<variable name="v9" units="flux" public_interface="in"/>
<variable name="v11A" units="flux" public_interface="in"/>
<variable name="time" units="minute" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>N2</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>v7</ci>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>v9</ci>
</apply>
</apply>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">3</cn>
<ci>v4</ci>
</apply>
<ci>v11A</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="N">
<variable name="N" units="millimolar" initial_value="2"/>
<variable name="N1" units="millimolar" public_interface="out"/>
<variable name="N2" units="millimolar" public_interface="in"/>
<variable name="time" units="minute" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>N1</ci>
<apply>
<minus/>
<ci>N</ci>
<ci>N2</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="oah">
<variable name="oah" units="millimolar" initial_value="5.571685096" public_interface="out"/>
<variable name="v5" units="flux" public_interface="in"/>
<variable name="v15" units="flux" public_interface="in"/>
<variable name="v18" units="flux" public_interface="in"/>
<variable name="time" units="minute" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>oah</ci>
</apply>
<apply>
<minus/>
<ci>v15</ci>
<apply>
<plus/>
<ci>v5</ci>
<ci>v18</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v1">
<variable name="cys" units="millimolar" public_interface="in"/>
<variable name="v0" units="flux" initial_value="1.6"/>
<variable name="f1" units="dimensionless"/>
<variable name="n" units="dimensionless" initial_value="4"/>
<variable name="Kc" units="millimolar" initial_value="0.1"/>
<variable name="v1" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v1</ci>
<apply>
<times/>
<ci>v0</ci>
<ci>f1</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>f1</ci>
<apply>
<power/>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<power/>
<apply>
<divide/>
<ci>cys</ci>
<ci>Kc</ci>
</apply>
<ci>n</ci>
</apply>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v2">
<variable name="sul" units="millimolar" public_interface="in"/>
<variable name="A3_c" units="millimolar" public_interface="in"/>
<variable name="k2" units="second_order_rate_constant" initial_value="0.2"/>
<variable name="v2" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v2</ci>
<apply>
<times/>
<ci>k2</ci>
<ci>sul</ci>
<ci>A3_c</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v3">
<variable name="aps" units="millimolar" public_interface="in"/>
<variable name="A3_c" units="millimolar" public_interface="in"/>
<variable name="k3" units="second_order_rate_constant" initial_value="0.2"/>
<variable name="v3" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v3</ci>
<apply>
<times/>
<ci>k3</ci>
<ci>aps</ci>
<ci>A3_c</ci>
</apply>
</apply>
</math>
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<component xmlns="http://www.cellml.org/cellml/1.0#" name="v4">
<variable name="pap" units="millimolar" public_interface="in"/>
<variable name="N2" units="millimolar" public_interface="in"/>
<variable name="k4" units="second_order_rate_constant" initial_value="0.2"/>
<variable name="v4" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v4</ci>
<apply>
<times/>
<ci>k4</ci>
<ci>pap</ci>
<ci>N2</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v5">
<variable name="hyd" units="millimolar" public_interface="in"/>
<variable name="oah" units="millimolar" public_interface="in"/>
<variable name="k5" units="second_order_rate_constant" initial_value="0.1"/>
<variable name="v5" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v5</ci>
<apply>
<times/>
<ci>k5</ci>
<ci>hyd</ci>
<ci>oah</ci>
</apply>
</apply>
</math>
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<component xmlns="http://www.cellml.org/cellml/1.0#" name="v6">
<variable name="cys" units="millimolar" public_interface="in"/>
<variable name="k6" units="first_order_rate_constant" initial_value="0.12"/>
<variable name="v6" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v6</ci>
<apply>
<times/>
<ci>k6</ci>
<ci>cys</ci>
</apply>
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<component xmlns="http://www.cellml.org/cellml/1.0#" name="v7">
<variable name="eth" units="millimolar" public_interface="in"/>
<variable name="N1" units="millimolar" public_interface="in"/>
<variable name="k7" units="second_order_rate_constant" initial_value="10"/>
<variable name="v7" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v7</ci>
<apply>
<times/>
<ci>k7</ci>
<ci>eth</ci>
<ci>N1</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v8">
<variable name="aco" units="millimolar" public_interface="in"/>
<variable name="S2" units="millimolar" public_interface="in"/>
<variable name="k8" units="second_order_rate_constant" initial_value="10"/>
<variable name="v8" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v8</ci>
<apply>
<times/>
<ci>k8</ci>
<ci>aco</ci>
<ci>S2</ci>
</apply>
</apply>
</math>
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<component xmlns="http://www.cellml.org/cellml/1.0#" name="v9">
<variable name="S1" units="millimolar" public_interface="in"/>
<variable name="N1" units="millimolar" public_interface="in"/>
<variable name="k9" units="second_order_rate_constant" initial_value="10"/>
<variable name="v9" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v9</ci>
<apply>
<times/>
<ci>k9</ci>
<ci>S1</ci>
<ci>N1</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v10">
<variable name="v10" units="flux" initial_value="80" public_interface="out"/>
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<component xmlns="http://www.cellml.org/cellml/1.0#" name="v11A">
<variable name="oxy" units="millimolar" public_interface="in"/>
<variable name="hyd" units="millimolar" public_interface="in"/>
<variable name="N2" units="millimolar" public_interface="in"/>
<variable name="alpha" units="dimensionless" initial_value="0.1"/>
<variable name="f2" units="dimensionless"/>
<variable name="m" units="dimensionless" initial_value="4"/>
<variable name="KH" units="millimolar" initial_value="0.5"/>
<variable name="k11" units="first_order_rate_constant" initial_value="10"/>
<variable name="v11A" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v11A</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>k11</ci>
<ci>N2</ci>
<ci>oxy</ci>
<ci>f2</ci>
</apply>
<apply>
<plus/>
<apply>
<times/>
<ci>alpha</ci>
<ci>N2</ci>
</apply>
<ci>oxy</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>f2</ci>
<apply>
<power/>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<power/>
<apply>
<divide/>
<ci>hyd</ci>
<ci>KH</ci>
</apply>
<ci>m</ci>
</apply>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
</apply>
</apply>
</apply>
</math>
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<component xmlns="http://www.cellml.org/cellml/1.0#" name="v11B">
<variable name="A2_m" units="millimolar" public_interface="in"/>
<variable name="A3_m" units="millimolar" public_interface="in"/>
<variable name="v11A" units="flux" public_interface="in"/>
<variable name="KA" units="millimolar" initial_value="1"/>
<variable name="v11B" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v11B</ci>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="dimensionless">3</cn>
<ci>v11A</ci>
<ci>A2_m</ci>
</apply>
<apply>
<plus/>
<ci>KA</ci>
<ci>A2_m</ci>
</apply>
</apply>
</apply>
</math>
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<component xmlns="http://www.cellml.org/cellml/1.0#" name="v12">
<variable name="A3_c" units="millimolar" public_interface="in"/>
<variable name="k12" units="first_order_rate_constant" initial_value="5"/>
<variable name="v12" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v12</ci>
<apply>
<times/>
<ci>k12</ci>
<ci>A3_c</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v13">
<variable name="v13" units="flux" initial_value="4" public_interface="out"/>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v14">
<variable name="oxy" units="millimolar" public_interface="in"/>
<variable name="k14" units="first_order_rate_constant" initial_value="10"/>
<variable name="v14" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v14</ci>
<apply>
<times/>
<ci>k14</ci>
<ci>oxy</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v15">
<variable name="aco" units="millimolar" public_interface="in"/>
<variable name="k15" units="first_order_rate_constant" initial_value="5"/>
<variable name="v15" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v15</ci>
<apply>
<times/>
<ci>k15</ci>
<ci>aco</ci>
</apply>
</apply>
</math>
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<component xmlns="http://www.cellml.org/cellml/1.0#" name="v16">
<variable name="A3_m" units="millimolar" public_interface="in"/>
<variable name="A2_c" units="millimolar" public_interface="in"/>
<variable name="k16" units="second_order_rate_constant" initial_value="10"/>
<variable name="v16" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v16</ci>
<apply>
<times/>
<ci>k16</ci>
<ci>A3_m</ci>
<ci>A2_c</ci>
</apply>
</apply>
</math>
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<component xmlns="http://www.cellml.org/cellml/1.0#" name="v17">
<variable name="hyd" units="millimolar" public_interface="in"/>
<variable name="k17" units="first_order_rate_constant" initial_value="0.02"/>
<variable name="v17" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v17</ci>
<apply>
<times/>
<ci>k17</ci>
<ci>hyd</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v18">
<variable name="oah" units="millimolar" public_interface="in"/>
<variable name="k18" units="first_order_rate_constant" initial_value="1"/>
<variable name="v18" units="flux" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v18</ci>
<apply>
<times/>
<ci>k18</ci>
<ci>oah</ci>
</apply>
</apply>
</math>
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<map_components component_1="S1" component_2="environment"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="S" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="sul" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="aps" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="pap" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="hyd" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="cys" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="eth" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="aco" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="oxy" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A_c" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
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<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A_m" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A3_m" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="N" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="N2" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="oah" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="S1" component_2="v8"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="S1" component_2="v9"/>
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<map_variables variable_1="v9" variable_2="v9"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="S" component_2="v8"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="S1" component_2="S"/>
<map_variables variable_1="S1" variable_2="S1"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="sul" component_2="v1"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="sul" component_2="v2"/>
<map_variables variable_1="sul" variable_2="sul"/>
<map_variables variable_1="v2" variable_2="v2"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="aps" component_2="v2"/>
<map_variables variable_1="v2" variable_2="v2"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="aps" component_2="v3"/>
<map_variables variable_1="aps" variable_2="aps"/>
<map_variables variable_1="v3" variable_2="v3"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="pap" component_2="v3"/>
<map_variables variable_1="v3" variable_2="v3"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="pap" component_2="v4"/>
<map_variables variable_1="pap" variable_2="pap"/>
<map_variables variable_1="v4" variable_2="v4"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="hyd" component_2="v4"/>
<map_variables variable_1="v4" variable_2="v4"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="hyd" component_2="v5"/>
<map_variables variable_1="hyd" variable_2="hyd"/>
<map_variables variable_1="v5" variable_2="v5"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="hyd" component_2="v11A"/>
<map_variables variable_1="hyd" variable_2="hyd"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="hyd" component_2="v17"/>
<map_variables variable_1="hyd" variable_2="hyd"/>
<map_variables variable_1="v17" variable_2="v17"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="cys" component_2="v1"/>
<map_variables variable_1="cys" variable_2="cys"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="cys" component_2="v5"/>
<map_variables variable_1="v5" variable_2="v5"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="cys" component_2="v6"/>
<map_variables variable_1="cys" variable_2="cys"/>
<map_variables variable_1="v6" variable_2="v6"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="eth" component_2="v13"/>
<map_variables variable_1="v13" variable_2="v13"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="eth" component_2="v7"/>
<map_variables variable_1="eth" variable_2="eth"/>
<map_variables variable_1="v7" variable_2="v7"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="aco" component_2="v7"/>
<map_variables variable_1="v7" variable_2="v7"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="aco" component_2="v8"/>
<map_variables variable_1="aco" variable_2="aco"/>
<map_variables variable_1="v8" variable_2="v8"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="aco" component_2="v15"/>
<map_variables variable_1="aco" variable_2="aco"/>
<map_variables variable_1="v15" variable_2="v15"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="oxy" component_2="v10"/>
<map_variables variable_1="v10" variable_2="v10"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="oxy" component_2="v11A"/>
<map_variables variable_1="oxy" variable_2="oxy"/>
<map_variables variable_1="v11A" variable_2="v11A"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="oxy" component_2="v14"/>
<map_variables variable_1="oxy" variable_2="oxy"/>
<map_variables variable_1="v14" variable_2="v14"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A3_m" component_2="v11B"/>
<map_variables variable_1="A3_m" variable_2="A3_m"/>
<map_variables variable_1="v11B" variable_2="v11B"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A3_m" component_2="v16"/>
<map_variables variable_1="A3_m" variable_2="A3_m"/>
<map_variables variable_1="v16" variable_2="v16"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="N2" component_2="v4"/>
<map_variables variable_1="N2" variable_2="N2"/>
<map_variables variable_1="v4" variable_2="v4"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="N2" component_2="v7"/>
<map_variables variable_1="v7" variable_2="v7"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A_c" component_2="v16"/>
<map_variables variable_1="A2_c" variable_2="A2_c"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="N2" component_2="v9"/>
<map_variables variable_1="v9" variable_2="v9"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="N2" component_2="v11A"/>
<map_variables variable_1="N2" variable_2="N2"/>
<map_variables variable_1="v11A" variable_2="v11A"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A3_c" component_2="v2"/>
<map_variables variable_1="A3_c" variable_2="A3_c"/>
<map_variables variable_1="v2" variable_2="v2"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A3_c" component_2="v3"/>
<map_variables variable_1="A3_c" variable_2="A3_c"/>
<map_variables variable_1="v3" variable_2="v3"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A3_c" component_2="v12"/>
<map_variables variable_1="A3_c" variable_2="A3_c"/>
<map_variables variable_1="v12" variable_2="v12"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A3_c" component_2="v16"/>
<map_variables variable_1="v16" variable_2="v16"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="oah" component_2="v5"/>
<map_variables variable_1="oah" variable_2="oah"/>
<map_variables variable_1="v5" variable_2="v5"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="oah" component_2="v15"/>
<map_variables variable_1="v15" variable_2="v15"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="oah" component_2="v18"/>
<map_variables variable_1="oah" variable_2="oah"/>
<map_variables variable_1="v18" variable_2="v18"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A_c" component_2="A3_c"/>
<map_variables variable_1="A3_c" variable_2="A3_c"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A_m" component_2="v11B"/>
<map_variables variable_1="A2_m" variable_2="A2_m"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="A_m" component_2="A3_m"/>
<map_variables variable_1="A3_m" variable_2="A3_m"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="N" component_2="v7"/>
<map_variables variable_1="N1" variable_2="N1"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="N" component_2="v9"/>
<map_variables variable_1="N1" variable_2="N1"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="N" component_2="N2"/>
<map_variables variable_1="N2" variable_2="N2"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="v11A" component_2="v11B"/>
<map_variables variable_1="v11A" variable_2="v11A"/>
</connection>
</model>
