<?xml version="1.0" encoding="utf-8"?>
<!--
This CellML file was generated on 22/04/2010 at 2:44:01 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/
-->
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<title>Mechanism of Protection of Peroxidase Activity by Oscillatory Dynamics</title>
<author>
<firstname>Catherine</firstname>
<surname>Lloyd</surname>
<affiliation>
<shortaffil>Auckland Bioengineering Institute, The University of Auckland</shortaffil>
</affiliation>
</author>
</articleinfo>
<section id="sec_status">
<title>Model Status</title>
<para>
This model runs in COR and OpenCell and the units are consistent throughout. At present it does not reproduce the published results, although the equations are a faithful match to the paper. Oscillatory behavior is seen in the published figures but cannot be reproduced in CellML, and is probably a result of an additional equation describing oxygen fluctuations. However, when run to steady state the state variables are close to published steady state values.
</para>
</section>
<sect1 id="sec_structure">
<title>Model Structure</title>
<para>
ABSTRACT: The peroxidase-oxidase reaction is known to involve reactive oxygen species as intermediates. These intermediates inactivate many types of biomolecules, including peroxidase itself. Previously, we have shown that oscillatory dynamics in the peroxidase-oxidase reaction seem to protect the enzyme from inactivation. It was suggested that this is due to a lower average concentration of reactive oxygen species in the oscillatory state compared to the steady state. Here, we studied the peroxidase-oxidase reaction with either 4-hydroxybenzoic acid or melatonin as cofactors. We show that the protective effect of oscillatory dynamics is present in both cases. We also found that the enzyme degradation depends on the concentration of the cofactor and on the pH of the reaction mixture. We simulated the oscillatory behaviour, including the oscillation/steady state bistability observed experimentally, using a detailed reaction scheme. The computational results confirm the hypothesis that protection is due to lower average concentrations of superoxide radical during oscillations. They also show that the shape of the oscillations changes with increasing cofactor concentration resulting in a further decrease in the average concentration of radicals. We therefore hypothesize that the protective effect of oscillatory dynamics is a general effect in this system.
</para>
<para>
The original paper reference is cited below:
</para>
<para>
Mechanism of protection of peroxidase activity by oscillatory dynamics, Lars F. Olsen, Marcus J. B. Hauser and Ursula Kummer, 2003, <emphasis>European Journal of Biochemistry</emphasis>, 270, 2796-2804. <ulink url="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&list_uids=12823550&dopt=Abstract">PubMed ID: 12823550</ulink>
</para>
<informalfigure float="0" id="fig_reaction_diagram">
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<title>reaction diagram</title>
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<caption>Schematic diagram of the peroxidase-oxidase reaction. Per<superscript>3+</superscript> and Per<superscript>2+</superscript> indicate iron(III) and iron(II) peroxidase respectively. Enzyme intermediates compound I, II and III are represented as coI, coII, and coIII. ArH and Ar' indicate the aromatic compound 4-hydroxybenzoic acid, or melatonin, and its free radical respectively.</caption>
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<dcterms:W3CDTF>2003-07</dcterms:W3CDTF>
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<dcterms:W3CDTF>2004-01-22</dcterms:W3CDTF>
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The University of Auckland, Auckland Bioengineering Institute
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Olsen et al.'s 2003 mathematical model of the peroxidase-oxidase reaction.
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Mechanism of protection of peroxidase activity by oscillatory dynamics
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<variable name="Per2" units="molar" initial_value="1e-10" public_interface="out"/>
<variable name="v10" units="flux" public_interface="in"/>
<variable name="v11" units="flux" public_interface="in"/>
<variable name="time" units="second" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Per2</ci>
</apply>
<apply>
<plus/>
<apply>
<minus/>
<ci>v11</ci>
</apply>
<ci>v10</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="coI">
<variable name="coI" units="molar" initial_value="8.9e-10" public_interface="out"/>
<variable name="v2" units="flux" public_interface="in"/>
<variable name="v3" units="flux" public_interface="in"/>
<variable name="v8" units="flux" public_interface="in"/>
<variable name="time" units="second" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>coI</ci>
</apply>
<apply>
<plus/>
<apply>
<minus/>
<ci>v2</ci>
<ci>v3</ci>
</apply>
<ci>v8</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="coII">
<variable name="coII" units="molar" initial_value="0.026e-6" public_interface="out"/>
<variable name="v3" units="flux" public_interface="in"/>
<variable name="v4" units="flux" public_interface="in"/>
<variable name="time" units="second" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>coII</ci>
</apply>
<apply>
<minus/>
<ci>v3</ci>
<ci>v4</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="coIII">
<variable name="coIII" units="molar" initial_value="1.31e-6" public_interface="out"/>
<variable name="v6" units="flux" public_interface="in"/>
<variable name="v8" units="flux" public_interface="in"/>
<variable name="v11" units="flux" public_interface="in"/>
<variable name="time" units="second" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>coIII</ci>
</apply>
<apply>
<plus/>
<apply>
<minus/>
<ci>v6</ci>
<ci>v8</ci>
</apply>
<ci>v11</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="Ar_radical">
<variable name="Ar_radical" units="molar" initial_value="0.12e-6" public_interface="out"/>
<variable name="v3" units="flux" public_interface="in"/>
<variable name="v4" units="flux" public_interface="in"/>
<variable name="v14" units="flux" public_interface="in"/>
<variable name="time" units="second" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Ar_radical</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<ci>v3</ci>
<ci>v4</ci>
</apply>
<ci>v14</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="ArH">
<variable name="ArH" units="molar" initial_value="299.88e-6" public_interface="out"/>
<variable name="v3" units="flux" public_interface="in"/>
<variable name="v4" units="flux" public_interface="in"/>
<variable name="v14" units="flux" public_interface="in"/>
<variable name="time" units="second" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>ArH</ci>
</apply>
<apply>
<plus/>
<apply>
<minus/>
<apply>
<minus/>
<ci>v3</ci>
</apply>
<ci>v4</ci>
</apply>
<ci>v14</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v1">
<variable name="v1" units="flux" public_interface="out"/>
<variable name="NADH" units="molar" public_interface="in"/>
<variable name="O2" units="molar" public_interface="in"/>
<variable name="k1" units="second_order_rate_constant" initial_value="3"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v1</ci>
<apply>
<times/>
<ci>k1</ci>
<ci>NADH</ci>
<ci>O2</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v2">
<variable name="v2" units="flux" public_interface="out"/>
<variable name="H2O2" units="molar" public_interface="in"/>
<variable name="Per3" units="molar" public_interface="in"/>
<variable name="k2" units="second_order_rate_constant" initial_value="1.8e7"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v2</ci>
<apply>
<times/>
<ci>k2</ci>
<ci>H2O2</ci>
<ci>Per3</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v3">
<variable name="v3" units="flux" public_interface="out"/>
<variable name="coI" units="molar" public_interface="in"/>
<variable name="ArH" units="molar" public_interface="in"/>
<variable name="k3" units="second_order_rate_constant" initial_value="1.5e5"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v3</ci>
<apply>
<times/>
<ci>k3</ci>
<ci>coI</ci>
<ci>ArH</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v4">
<variable name="v4" units="flux" public_interface="out"/>
<variable name="coII" units="molar" public_interface="in"/>
<variable name="ArH" units="molar" public_interface="in"/>
<variable name="k4" units="second_order_rate_constant" initial_value="5.2e3"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v4</ci>
<apply>
<times/>
<ci>k4</ci>
<ci>coII</ci>
<ci>ArH</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v5">
<variable name="v5" units="flux" public_interface="out"/>
<variable name="NAD_radical" units="molar" public_interface="in"/>
<variable name="O2" units="molar" public_interface="in"/>
<variable name="k5" units="second_order_rate_constant" initial_value="2e7"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v5</ci>
<apply>
<times/>
<ci>k5</ci>
<ci>NAD_radical</ci>
<ci>O2</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v6">
<variable name="v6" units="flux" public_interface="out"/>
<variable name="O2_radical" units="molar" public_interface="in"/>
<variable name="Per3" units="molar" public_interface="in"/>
<variable name="k6" units="second_order_rate_constant" initial_value="1.7e7"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v6</ci>
<apply>
<times/>
<ci>k6</ci>
<ci>O2_radical</ci>
<ci>Per3</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v7">
<variable name="v7" units="flux" public_interface="out"/>
<variable name="O2_radical" units="molar" public_interface="in"/>
<variable name="k7" units="second_order_rate_constant" initial_value="2e7"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v7</ci>
<apply>
<times/>
<ci>k7</ci>
<apply>
<power/>
<ci>O2_radical</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v8">
<variable name="v8" units="flux" public_interface="out"/>
<variable name="coIII" units="molar" public_interface="in"/>
<variable name="NAD_radical" units="molar" public_interface="in"/>
<variable name="k8" units="second_order_rate_constant" initial_value="4e7"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v8</ci>
<apply>
<times/>
<ci>k8</ci>
<ci>coIII</ci>
<ci>NAD_radical</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v9">
<variable name="v9" units="flux" public_interface="out"/>
<variable name="NAD_radical" units="molar" public_interface="in"/>
<variable name="k9" units="second_order_rate_constant" initial_value="6e7"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v9</ci>
<apply>
<times/>
<ci>k9</ci>
<apply>
<power/>
<ci>NAD_radical</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v10">
<variable name="v10" units="flux" public_interface="out"/>
<variable name="NAD_radical" units="molar" public_interface="in"/>
<variable name="Per3" units="molar" public_interface="in"/>
<variable name="k10" units="second_order_rate_constant" initial_value="1.8e6"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v10</ci>
<apply>
<times/>
<ci>k10</ci>
<ci>Per3</ci>
<ci>NAD_radical</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v11">
<variable name="v11" units="flux" public_interface="out"/>
<variable name="Per2" units="molar" public_interface="in"/>
<variable name="O2" units="molar" public_interface="in"/>
<variable name="k11" units="second_order_rate_constant" initial_value="1e5"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v11</ci>
<apply>
<times/>
<ci>k11</ci>
<ci>Per2</ci>
<ci>O2</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v12">
<variable name="v12" units="flux" public_interface="out"/>
<variable name="k12" units="flux" initial_value="0.08e-6"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v12</ci>
<ci>k12</ci>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v13">
<variable name="v13" units="flux" public_interface="out"/>
<variable name="O2eq" units="molar" initial_value="1.2e-5"/>
<variable name="k13" units="first_order_rate_constant" initial_value="6e-3"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v13</ci>
<apply>
<times/>
<ci>k13</ci>
<ci>O2eq</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v13_back">
<variable name="v13_back" units="flux" public_interface="out"/>
<variable name="O2" units="molar" public_interface="in"/>
<variable name="k13_" units="first_order_rate_constant" initial_value="6e-3"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v13_back</ci>
<apply>
<times/>
<ci>k13_</ci>
<ci>O2</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="v14">
<variable name="v14" units="flux" public_interface="out"/>
<variable name="Ar_radical" units="molar" public_interface="in"/>
<variable name="NADH" units="molar" public_interface="in"/>
<variable name="k14" units="second_order_rate_constant" initial_value="7e5"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>v14</ci>
<apply>
<times/>
<ci>k14</ci>
<ci>Ar_radical</ci>
<ci>NADH</ci>
</apply>
</apply>
</math>
</component>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NADH" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NAD" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NAD2" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NAD_radical" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="O2" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="O2_radical" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="Per3" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="Per2" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="coI" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="coII" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="coIII" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="H2O2" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="ArH" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="Ar_radical" component_2="environment"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NADH" component_2="v1"/>
<map_variables variable_1="v1" variable_2="v1"/>
<map_variables variable_1="NADH" variable_2="NADH"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="O2" component_2="v1"/>
<map_variables variable_1="v1" variable_2="v1"/>
<map_variables variable_1="O2" variable_2="O2"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NAD" component_2="v1"/>
<map_variables variable_1="v1" variable_2="v1"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="H2O2" component_2="v1"/>
<map_variables variable_1="v1" variable_2="v1"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="H2O2" component_2="v2"/>
<map_variables variable_1="v2" variable_2="v2"/>
<map_variables variable_1="H2O2" variable_2="H2O2"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="Per3" component_2="v2"/>
<map_variables variable_1="v2" variable_2="v2"/>
<map_variables variable_1="Per3" variable_2="Per3"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="coI" component_2="v2"/>
<map_variables variable_1="v2" variable_2="v2"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="coI" component_2="v3"/>
<map_variables variable_1="v3" variable_2="v3"/>
<map_variables variable_1="coI" variable_2="coI"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="ArH" component_2="v3"/>
<map_variables variable_1="v3" variable_2="v3"/>
<map_variables variable_1="ArH" variable_2="ArH"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="Ar_radical" 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="coII" 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="coII" component_2="v4"/>
<map_variables variable_1="v4" variable_2="v4"/>
<map_variables variable_1="coII" variable_2="coII"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="ArH" component_2="v4"/>
<map_variables variable_1="v4" variable_2="v4"/>
<map_variables variable_1="ArH" variable_2="ArH"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="Per3" 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="Ar_radical" 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="NAD_radical" component_2="v5"/>
<map_variables variable_1="v5" variable_2="v5"/>
<map_variables variable_1="NAD_radical" variable_2="NAD_radical"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="O2" component_2="v5"/>
<map_variables variable_1="v5" variable_2="v5"/>
<map_variables variable_1="O2" variable_2="O2"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NAD" 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="O2_radical" 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="O2_radical" component_2="v6"/>
<map_variables variable_1="v6" variable_2="v6"/>
<map_variables variable_1="O2_radical" variable_2="O2_radical"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="Per3" component_2="v6"/>
<map_variables variable_1="v6" variable_2="v6"/>
<map_variables variable_1="Per3" variable_2="Per3"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="coIII" component_2="v6"/>
<map_variables variable_1="v6" variable_2="v6"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="O2_radical" component_2="v7"/>
<map_variables variable_1="v7" variable_2="v7"/>
<map_variables variable_1="O2_radical" variable_2="O2_radical"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="H2O2" 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="O2" 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="coIII" component_2="v8"/>
<map_variables variable_1="v8" variable_2="v8"/>
<map_variables variable_1="coIII" variable_2="coIII"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NAD_radical" component_2="v8"/>
<map_variables variable_1="v8" variable_2="v8"/>
<map_variables variable_1="NAD_radical" variable_2="NAD_radical"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="coI" component_2="v8"/>
<map_variables variable_1="v8" variable_2="v8"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NAD" component_2="v8"/>
<map_variables variable_1="v8" variable_2="v8"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NAD_radical" component_2="v9"/>
<map_variables variable_1="v9" variable_2="v9"/>
<map_variables variable_1="NAD_radical" variable_2="NAD_radical"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NAD2" 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="Per3" component_2="v10"/>
<map_variables variable_1="v10" variable_2="v10"/>
<map_variables variable_1="Per3" variable_2="Per3"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NAD_radical" component_2="v10"/>
<map_variables variable_1="v10" variable_2="v10"/>
<map_variables variable_1="NAD_radical" variable_2="NAD_radical"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="Per2" 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="NAD" 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="Per2" component_2="v11"/>
<map_variables variable_1="v11" variable_2="v11"/>
<map_variables variable_1="Per2" variable_2="Per2"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="O2" component_2="v11"/>
<map_variables variable_1="v11" variable_2="v11"/>
<map_variables variable_1="O2" variable_2="O2"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="coIII" component_2="v11"/>
<map_variables variable_1="v11" variable_2="v11"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NADH" component_2="v12"/>
<map_variables variable_1="v12" variable_2="v12"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="O2" 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="O2" component_2="v13_back"/>
<map_variables variable_1="v13_back" variable_2="v13_back"/>
<map_variables variable_1="O2" variable_2="O2"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="Ar_radical" component_2="v14"/>
<map_variables variable_1="v14" variable_2="v14"/>
<map_variables variable_1="Ar_radical" variable_2="Ar_radical"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NADH" component_2="v14"/>
<map_variables variable_1="v14" variable_2="v14"/>
<map_variables variable_1="NADH" variable_2="NADH"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="ArH" component_2="v14"/>
<map_variables variable_1="v14" variable_2="v14"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="NAD_radical" component_2="v14"/>
<map_variables variable_1="v14" variable_2="v14"/>
</connection>
</model>
