- Author:
- pmr2.import <nobody@models.cellml.org>
- Date:
- 2006-07-09 07:21:19+12:00
- Desc:
- committing version01 of ciliberto_petrus_tyson_sible_2003
- Permanent Source URI:
- https://models.physiomeproject.org/workspace/36d/rawfile/f37e3955b3d43a63b23ad906564a8849729d1e2b/ciliberto_petrus_tyson_sible_2003.cellml
<?xml version='1.0' encoding='utf-8'?>
<!-- FILE : ciliberto_model_2003.xml
CREATED : 8th December 2003
LAST MODIFIED : 8th December 2003
AUTHOR : Catherine Lloyd
Bioengineering Institute
The University of Auckland
MODEL STATUS : This model conforms to the CellML 1.0 Specification released on
10th August 2001, and the 16/1/02 CellML Metadata 1.0 Specification.
DESCRIPTION : This file contains a CellML description of Ciliberto et al.'s 2003 kinetic model of the cyclin E/Cdk2 developmental timer in Xenopus laevis embryos.
CHANGES:
--><model xmlns="http://www.cellml.org/cellml/1.0#" xmlns:cmeta="http://www.cellml.org/metadata/1.0#" 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:cellml="http://www.cellml.org/cellml/1.0#" xmlns:dcterms="http://purl.org/dc/terms/" xmlns:vCard="http://www.w3.org/2001/vcard-rdf/3.0#" cmeta:id="ciliberto_petrus_tyson_sible_2003_version01" name="ciliberto_petrus_tyson_sible_2003_version01">
<documentation xmlns="http://cellml.org/tmp-documentation">
<article>
<articleinfo>
<title>A Kinetic Model of the Cyclin E/Cdk2 Developmental Timer in Xenopus laevis embryos</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 is the original unchecked version of the model imported from the previous
CellML model repository, 24-Jan-2006.
</para>
</section>
<sect1 id="sec_structure">
<title>Model Structure</title>
<para>
The cell cycle in <emphasis>Xenopus</emphasis> embryos has been the subject of several experimental and theoretical studies. Following fertilisation, the egg and sperm nuclei undergo DNA replication, nuclear fusion and the first mitotic cellular division. The next 11 embryonic cell cycles are rapid, synchronous oscillations between DNA replication and mitosis, lacking the usual intervening gap phases, cell growth and cell cycle checkpoints. After the 12th division, the <emphasis>Xenopus</emphasis> embryo undergoes the midblastula transition (MBT), when cells become motile, embryonic transcription begins and the cell cycle acquires the gap phases and checkpoints of a typical somatic cell.
</para>
<para>
The mechanisms underlying this cell cycle remodelling are not well understood, but it is thought that they may depend in part on a maternal developmental timer driven by oscillations in the activity of cyclin E/Cdk2. Cdk2 activity oscillates twice per cell cycle, despite a constant level of its partner cyclin E. Cyclin E breaks down at a fixed time after fertilisation, usually around MBT. In an effort to better understand the mechanisms underlying these phenomena, Ciliberto <emphasis>et al.</emphasis> have developed a mathematical model of the cyclin E/Cdk2 developmental timer that describes both the oscillations in Cdk2 activity and the sudden degradation of Cyclin E when the MBT begins (see <xref linkend="fig_reaction_diagram"/> below). In this mathematical model,
<itemizedlist>
<listitem>
<para>oscillations in Cdk2 activity depend on changes in phosphorylation;</para>
</listitem>
<listitem>
<para>Cdk2 participates in a negative feedback loop with inhibitory kinase Wee1;</para>
</listitem>
<listitem>
<para>cyclin E is cooperatively removed from the oscillatory system; and</para>
</listitem>
<listitem>
<para>removed cyclin E is degraded by a pathway activated by cyclin E/Cdk2 itself.</para>
</listitem>
</itemizedlist>
</para>
<para>
The model has been described here in CellML (the raw CellML description of the Ciliberto <emphasis>et al.</emphasis> 2003 model can be downloaded in various formats as described in <xref linkend="sec_download_this_model"/>).
</para>
<para>
The complete original paper reference is cited below:
</para>
<para>
<ulink url="http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6TFB-48V891M-3&_user=140507&_coverDate=07%2F01%2F2003&_alid=133391549&_rdoc=1&_fmt=summary&_orig=search&_qd=1&_cdi=5222&_sort=d&_docanchor=&view=c&_acct=C000011498&_version=1&_urlVersion=0&_userid=140507&md5=ac3ac20d5deb272fb6e0b0156660bbc7">A kinetic model of the cyclin E/Cdk2 developmental timer in <emphasis>Xenopus laevis</emphasis> embryos</ulink>, Andrea Ciliberto, Matthew J. Petrus, John J. Tyson, and Jill C. Sible, 2003, <ulink url="http://www.sciencedirect.com/science?_ob=JournalURL&_cdi=5222&_auth=y&_acct=C000011498&_version=1&_urlVersion=0&_userid=140507&md5=7d06a1ef3b4293bc0633b8269aa54192">
<emphasis>Biophysical Chemistry</emphasis>
</ulink>, 104, 573-589. (<ulink url="http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6TFB-48V891M-3&_coverDate=07%2F01%2F2003&_alid=133391549&_rdoc=1&_fmt=&_orig=search&_qd=1&_cdi=5222&_sort=d&view=c&_acct=C000011498&_version=1&_urlVersion=0&_userid=140507&md5=8b3236650c4054e9e644ebbf4cd680e0">Full text (HTML)</ulink> and PDF versions of the article are available on the <emphasis>Biophysical Chemistry</emphasis> website.) <ulink url="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&list_uids=12914904&dopt=Abstract">PubMed ID: 12914904</ulink>
</para>
<informalfigure float="0" id="fig_reaction_diagram">
<mediaobject>
<imageobject>
<objectinfo>
<title>reaction diagram</title>
</objectinfo>
<imagedata fileref="ciliberto_2003.png"/>
</imageobject>
</mediaobject>
<caption>A theoretical molecular mechanism of the cyclin E/Cdk2 developmental timer was used to construct the mathematical model. The dashed box represents a mechanism for removing cyclin E/Cdk2 from the oscillatory subsystem, which is activated by the "removed" form of cyclin E/Cdk2. The removal step is cooperative in that the more cyclin E/Cdk2 there is bound to the dashed box, the faster is the association reaction. A kinase (Kin) is introduced between cyclin E/Cdk2 and Wee1 to create a time lag in the negative feedback loop.</caption>
</informalfigure>
</sect1>
</article>
</documentation>
<units name="minute">
<unit units="second" multiplier="60.0"/>
</units>
<units name="first_order_rate_constant">
<unit units="minute" exponent="-1"/>
</units>
<component name="environment">
<variable units="minute" public_interface="out" name="time"/>
</component>
<component name="Cdk2_CycE">
<variable units="dimensionless" public_interface="out" name="Cdk2_CycE"/>
<variable units="first_order_rate_constant" public_interface="in" name="kwee"/>
<variable units="first_order_rate_constant" public_interface="in" name="kon"/>
<variable units="first_order_rate_constant" public_interface="in" name="k25A"/>
<variable units="first_order_rate_constant" public_interface="in" name="koff"/>
<variable units="first_order_rate_constant" public_interface="in" name="kassoc"/>
<variable units="first_order_rate_constant" public_interface="in" name="kdissoc"/>
<variable units="dimensionless" public_interface="in" name="phi"/>
<variable units="dimensionless" public_interface="in" name="Wee1_"/>
<variable units="dimensionless" public_interface="in" name="PCdk2_CycE"/>
<variable units="dimensionless" public_interface="in" name="Xic"/>
<variable units="dimensionless" public_interface="in" name="Cdk2_CycErem"/>
<variable units="dimensionless" public_interface="in" name="Xic_Cdk2_CycE"/>
<variable units="minute" public_interface="in" name="time"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Cdk2_CycE</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
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<ci> k25A </ci>
<ci> PCdk2_CycE </ci>
</apply>
<apply>
<times/>
<ci> koff </ci>
<ci> Cdk2_CycErem </ci>
</apply>
<apply>
<times/>
<ci> kdissoc </ci>
<ci> Xic_Cdk2_CycE </ci>
</apply>
</apply>
<apply>
<plus/>
<apply>
<times/>
<ci> kwee </ci>
<ci> Wee1_ </ci>
<ci> Cdk2_CycE </ci>
</apply>
<apply>
<times/>
<ci> kon </ci>
<ci> phi </ci>
<ci> Cdk2_CycE </ci>
</apply>
<apply>
<times/>
<ci> kassoc </ci>
<ci> Xic </ci>
<ci> Cdk2_CycE </ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="PCdk2_CycE">
<variable units="dimensionless" public_interface="out" name="PCdk2_CycE"/>
<variable units="first_order_rate_constant" public_interface="in" name="kwee"/>
<variable units="first_order_rate_constant" public_interface="in" name="kon"/>
<variable units="first_order_rate_constant" public_interface="in" name="k25A"/>
<variable units="first_order_rate_constant" public_interface="in" name="koff"/>
<variable units="first_order_rate_constant" public_interface="in" name="kassoc"/>
<variable units="first_order_rate_constant" public_interface="in" name="kdissoc"/>
<variable units="dimensionless" public_interface="in" name="phi"/>
<variable units="dimensionless" public_interface="in" name="Wee1_"/>
<variable units="dimensionless" public_interface="in" name="Cdk2_CycE"/>
<variable units="dimensionless" public_interface="in" name="Xic"/>
<variable units="dimensionless" public_interface="in" name="PCdk2_CycErem"/>
<variable units="dimensionless" public_interface="in" name="Xic_PCdk2_CycE"/>
<variable units="minute" public_interface="in" name="time"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
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<eq/>
<apply>
<diff/>
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<ci>time</ci>
</bvar>
<ci>PCdk2_CycE</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
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<ci> kwee </ci>
<ci> Wee1_ </ci>
<ci> PCdk2_CycE </ci>
</apply>
<apply>
<times/>
<ci> koff </ci>
<ci> PCdk2_CycErem </ci>
</apply>
<apply>
<times/>
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<ci> Xic_PCdk2_CycE </ci>
</apply>
</apply>
<apply>
<plus/>
<apply>
<times/>
<ci> k25A </ci>
<ci> PCdk2_CycE </ci>
</apply>
<apply>
<times/>
<ci> kon </ci>
<ci> phi </ci>
<ci> PCdk2_CycE </ci>
</apply>
<apply>
<times/>
<ci> kassoc </ci>
<ci> Xic </ci>
<ci> PCdk2_CycE </ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="Wee1_">
<variable units="dimensionless" public_interface="out" name="Wee1_"/>
<variable units="dimensionless" name="Jwact" initial_value="0.01"/>
<variable units="dimensionless" name="Jwinact" initial_value="0.01"/>
<variable units="dimensionless" name="Wee1_total" initial_value="8.0"/>
<variable units="first_order_rate_constant" name="kwact" initial_value="0.75"/>
<variable units="first_order_rate_constant" name="kwinact" initial_value="1.5"/>
<variable units="dimensionless" public_interface="in" name="Kin_"/>
<variable units="minute" public_interface="in" name="time"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
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<eq/>
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<diff/>
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<ci>time</ci>
</bvar>
<ci>Wee1_</ci>
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<minus/>
<apply>
<divide/>
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<ci> kwact </ci>
<apply>
<minus/>
<ci> Wee1_total </ci>
<ci> Wee1_ </ci>
</apply>
</apply>
<apply>
<minus/>
<apply>
<plus/>
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<ci> Wee1_total </ci>
</apply>
<ci> Wee1_ </ci>
</apply>
</apply>
<apply>
<divide/>
<apply>
<times/>
<ci> kwinact </ci>
<ci> Kin_ </ci>
<ci> Wee1_ </ci>
</apply>
<apply>
<plus/>
<ci> Jwinact </ci>
<ci> Wee1_ </ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="Kin_">
<variable units="dimensionless" public_interface="out" name="Kin_"/>
<variable units="dimensionless" name="Jiact" initial_value="0.01"/>
<variable units="dimensionless" name="Jiinact" initial_value="0.01"/>
<variable units="first_order_rate_constant" name="kiact" initial_value="0.15"/>
<variable units="first_order_rate_constant" name="kiinact" initial_value="0.6"/>
<variable units="dimensionless" public_interface="in" name="Cdk2_CycE"/>
<variable units="minute" public_interface="in" name="time"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
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<divide/>
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<minus/>
<cn cellml:units="dimensionless"> 1.0 </cn>
<ci> Kin_ </ci>
</apply>
</apply>
<apply>
<minus/>
<apply>
<plus/>
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<cn cellml:units="dimensionless"> 1.0 </cn>
</apply>
<ci> Kin_ </ci>
</apply>
</apply>
<apply>
<divide/>
<apply>
<times/>
<ci> kiinact </ci>
<ci> Cdk2_CycE </ci>
<ci> Kin_ </ci>
</apply>
<apply>
<plus/>
<ci> Jiinact </ci>
<ci> Kin_ </ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="Cdk2_CycErem">
<variable units="dimensionless" public_interface="out" name="Cdk2_CycErem"/>
<variable units="first_order_rate_constant" public_interface="in" name="kedeg"/>
<variable units="first_order_rate_constant" public_interface="in" name="kon"/>
<variable units="first_order_rate_constant" public_interface="in" name="kxdeg"/>
<variable units="first_order_rate_constant" public_interface="in" name="koff"/>
<variable units="first_order_rate_constant" public_interface="in" name="kassoc"/>
<variable units="first_order_rate_constant" public_interface="in" name="kdissoc"/>
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<variable units="dimensionless" public_interface="in" name="Cdk2_CycE"/>
<variable units="dimensionless" public_interface="in" name="Xic"/>
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</bvar>
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<ci> Xic_Cdk2_CycE </ci>
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<ci> kwee </ci>
<ci> Wee1_ </ci>
<ci> Xic_Cdk2_CycE </ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="Xic_PCdk2_CycE">
<variable units="dimensionless" public_interface="out" name="Xic_PCdk2_CycE"/>
<variable units="first_order_rate_constant" public_interface="in" name="kwee"/>
<variable units="first_order_rate_constant" public_interface="in" name="kon"/>
<variable units="first_order_rate_constant" public_interface="in" name="k25A"/>
<variable units="first_order_rate_constant" public_interface="in" name="koff"/>
<variable units="first_order_rate_constant" public_interface="in" name="kassoc"/>
<variable units="first_order_rate_constant" public_interface="in" name="kdissoc"/>
<variable units="dimensionless" public_interface="in" name="phi"/>
<variable units="dimensionless" public_interface="in" name="Xic"/>
<variable units="dimensionless" public_interface="in" name="PCdk2_CycE"/>
<variable units="dimensionless" public_interface="in" name="Xic_PCdk2_CycErem"/>
<variable units="dimensionless" public_interface="in" name="Xic_Cdk2_CycE"/>
<variable units="dimensionless" public_interface="in" name="Wee1_"/>
<variable units="minute" public_interface="in" name="time"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Xic_PCdk2_CycE</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<ci> kassoc </ci>
<ci> Xic </ci>
<ci> PCdk2_CycE </ci>
</apply>
<apply>
<times/>
<ci> koff </ci>
<ci> Xic_PCdk2_CycErem </ci>
</apply>
<apply>
<times/>
<ci> kwee </ci>
<ci> Wee1_ </ci>
<ci> Xic_Cdk2_CycE </ci>
</apply>
</apply>
<apply>
<plus/>
<apply>
<times/>
<ci> kdissoc </ci>
<ci> Xic_PCdk2_CycE </ci>
</apply>
<apply>
<times/>
<ci> kon </ci>
<ci> phi </ci>
<ci> Xic_PCdk2_CycE </ci>
</apply>
<apply>
<times/>
<ci> k25A </ci>
<ci> Xic_PCdk2_CycE </ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="Xic_Cdk2_CycErem">
<variable units="dimensionless" public_interface="out" name="Xic_Cdk2_CycErem"/>
<variable units="first_order_rate_constant" public_interface="in" name="kedeg"/>
<variable units="first_order_rate_constant" public_interface="in" name="kon"/>
<variable units="first_order_rate_constant" public_interface="in" name="kxdeg"/>
<variable units="first_order_rate_constant" public_interface="in" name="koff"/>
<variable units="first_order_rate_constant" public_interface="in" name="kassoc"/>
<variable units="first_order_rate_constant" public_interface="in" name="kdissoc"/>
<variable units="dimensionless" public_interface="in" name="phi"/>
<variable units="dimensionless" public_interface="in" name="Deg"/>
<variable units="dimensionless" public_interface="in" name="Xic"/>
<variable units="dimensionless" public_interface="in" name="Cdk2_CycErem"/>
<variable units="dimensionless" public_interface="in" name="Xic_Cdk2_CycE"/>
<variable units="minute" public_interface="in" name="time"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Xic_Cdk2_CycErem</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<ci> kassoc </ci>
<ci> Xic </ci>
<ci> Cdk2_CycErem </ci>
</apply>
<apply>
<times/>
<ci> kon </ci>
<ci> phi </ci>
<ci> Xic_Cdk2_CycE </ci>
</apply>
</apply>
<apply>
<plus/>
<apply>
<times/>
<ci> kdissoc </ci>
<ci> Xic_Cdk2_CycErem </ci>
</apply>
<apply>
<times/>
<ci> koff </ci>
<ci> Xic_Cdk2_CycErem </ci>
</apply>
<apply>
<times/>
<ci> kedeg </ci>
<ci> Deg </ci>
<ci> Xic_Cdk2_CycErem </ci>
</apply>
<apply>
<times/>
<ci> kxdeg </ci>
<ci> Xic_Cdk2_CycErem </ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="Xic_PCdk2_CycErem">
<variable units="dimensionless" public_interface="out" name="Xic_PCdk2_CycErem"/>
<variable units="first_order_rate_constant" public_interface="in" name="kedeg"/>
<variable units="first_order_rate_constant" public_interface="in" name="kon"/>
<variable units="first_order_rate_constant" public_interface="in" name="kxdeg"/>
<variable units="first_order_rate_constant" public_interface="in" name="koff"/>
<variable units="first_order_rate_constant" public_interface="in" name="kassoc"/>
<variable units="first_order_rate_constant" public_interface="in" name="kdissoc"/>
<variable units="dimensionless" public_interface="in" name="phi"/>
<variable units="dimensionless" public_interface="in" name="Deg"/>
<variable units="dimensionless" public_interface="in" name="Xic"/>
<variable units="dimensionless" public_interface="in" name="PCdk2_CycErem"/>
<variable units="dimensionless" public_interface="in" name="Xic_PCdk2_CycE"/>
<variable units="minute" public_interface="in" name="time"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Xic_PCdk2_CycErem</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<ci> kassoc </ci>
<ci> Xic </ci>
<ci> PCdk2_CycErem </ci>
</apply>
<apply>
<times/>
<ci> kon </ci>
<ci> phi </ci>
<ci> Xic_PCdk2_CycE </ci>
</apply>
</apply>
<apply>
<plus/>
<apply>
<times/>
<ci> kdissoc </ci>
<ci> Xic_PCdk2_CycErem </ci>
</apply>
<apply>
<times/>
<ci> koff </ci>
<ci> Xic_PCdk2_CycErem </ci>
</apply>
<apply>
<times/>
<ci> kedeg </ci>
<ci> Deg </ci>
<ci> Xic_PCdk2_CycErem </ci>
</apply>
<apply>
<times/>
<ci> kxdeg </ci>
<ci> Xic_PCdk2_CycErem </ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="Xicrem">
<variable units="dimensionless" public_interface="out" name="Xicrem" initial_value="0.0"/>
<variable units="first_order_rate_constant" public_interface="in" name="kedeg"/>
<variable units="first_order_rate_constant" public_interface="in" name="kxdeg"/>
<variable units="dimensionless" public_interface="in" name="Deg"/>
<variable units="dimensionless" public_interface="in" name="Xic_Cdk2_CycErem"/>
<variable units="dimensionless" public_interface="in" name="Xic_PCdk2_CycErem"/>
<variable units="minute" public_interface="in" name="time"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Xicrem</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<ci> kedeg </ci>
<ci> Deg </ci>
<ci> Xic_Cdk2_CycErem </ci>
</apply>
<apply>
<times/>
<ci> kedeg </ci>
<ci> Deg </ci>
<ci> Xic_PCdk2_CycErem </ci>
</apply>
</apply>
<apply>
<times/>
<ci> kxdeg </ci>
<ci> Xicrem </ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="kinetic_parameters">
<variable units="first_order_rate_constant" public_interface="out" name="kwee" initial_value="1.5"/>
<variable units="first_order_rate_constant" public_interface="out" name="k25A" initial_value="0.1"/>
<variable units="first_order_rate_constant" public_interface="out" name="kon" initial_value="0.02"/>
<variable units="first_order_rate_constant" public_interface="out" name="koff" initial_value="0.0001"/>
<variable units="first_order_rate_constant" public_interface="out" name="kassoc" initial_value="0.1"/>
<variable units="first_order_rate_constant" public_interface="out" name="kdissoc" initial_value="0.001"/>
<variable units="first_order_rate_constant" public_interface="out" name="kedeg" initial_value="0.017"/>
<variable units="first_order_rate_constant" public_interface="out" name="kxdeg" initial_value="0.01"/>
<variable units="dimensionless" public_interface="out" name="phi"/>
<variable units="dimensionless" public_interface="out" name="Deg" initial_value="0.0"/>
<variable units="dimensionless" name="epsilon" initial_value="0.001"/>
<variable units="dimensionless" name="pool"/>
<variable units="dimensionless" name="n" initial_value="4.0"/>
<variable units="dimensionless" name="L" initial_value="0.4"/>
<variable units="dimensionless" public_interface="in" name="Cdk2_CycErem"/>
<variable units="dimensionless" public_interface="in" name="PCdk2_CycErem"/>
<variable units="dimensionless" public_interface="in" name="Xic_Cdk2_CycErem"/>
<variable units="dimensionless" public_interface="in" name="Xic_PCdk2_CycErem"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>phi</ci>
<apply>
<divide/>
<apply>
<plus/>
<ci> epsilon </ci>
<apply>
<power/>
<ci> pool </ci>
<ci> n </ci>
</apply>
</apply>
<apply>
<plus/>
<apply>
<power/>
<ci> L </ci>
<ci> n </ci>
</apply>
<apply>
<power/>
<ci> pool </ci>
<ci> n </ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<connection>
<map_components component_2="environment" component_1="Cdk2_CycE"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="PCdk2_CycE"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="Wee1_"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="Kin_"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="Cdk2_CycErem"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="PCdk2_CycErem"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="Deg_"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="Xic"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="Xic_Cdk2_CycE"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="Xic_PCdk2_CycE"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="Xic_Cdk2_CycErem"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="Xic_PCdk2_CycErem"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="Xicrem"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="Wee1_" component_1="Cdk2_CycE"/>
<map_variables variable_2="Wee1_" variable_1="Wee1_"/>
</connection>
<connection>
<map_components component_2="PCdk2_CycE" component_1="Cdk2_CycE"/>
<map_variables variable_2="PCdk2_CycE" variable_1="PCdk2_CycE"/>
<map_variables variable_2="Cdk2_CycE" variable_1="Cdk2_CycE"/>
</connection>
<connection>
<map_components component_2="Cdk2_CycErem" component_1="Cdk2_CycE"/>
<map_variables variable_2="Cdk2_CycErem" variable_1="Cdk2_CycErem"/>
<map_variables variable_2="Cdk2_CycE" variable_1="Cdk2_CycE"/>
</connection>
<connection>
<map_components component_2="Xic" component_1="Cdk2_CycE"/>
<map_variables variable_2="Xic" variable_1="Xic"/>
<map_variables variable_2="Cdk2_CycE" variable_1="Cdk2_CycE"/>
</connection>
<connection>
<map_components component_2="Xic_Cdk2_CycE" component_1="Cdk2_CycE"/>
<map_variables variable_2="Xic_Cdk2_CycE" variable_1="Xic_Cdk2_CycE"/>
<map_variables variable_2="Cdk2_CycE" variable_1="Cdk2_CycE"/>
</connection>
<connection>
<map_components component_2="Wee1_" component_1="PCdk2_CycE"/>
<map_variables variable_2="Wee1_" variable_1="Wee1_"/>
</connection>
<connection>
<map_components component_2="PCdk2_CycErem" component_1="PCdk2_CycE"/>
<map_variables variable_2="PCdk2_CycErem" variable_1="PCdk2_CycErem"/>
<map_variables variable_2="PCdk2_CycE" variable_1="PCdk2_CycE"/>
</connection>
<connection>
<map_components component_2="Xic" component_1="PCdk2_CycE"/>
<map_variables variable_2="Xic" variable_1="Xic"/>
<map_variables variable_2="PCdk2_CycE" variable_1="PCdk2_CycE"/>
</connection>
<connection>
<map_components component_2="Xic_PCdk2_CycE" component_1="PCdk2_CycE"/>
<map_variables variable_2="Xic_PCdk2_CycE" variable_1="Xic_PCdk2_CycE"/>
<map_variables variable_2="PCdk2_CycE" variable_1="PCdk2_CycE"/>
</connection>
<connection>
<map_components component_2="Cdk2_CycE" component_1="Kin_"/>
<map_variables variable_2="Cdk2_CycE" variable_1="Cdk2_CycE"/>
</connection>
<connection>
<map_components component_2="Xic" component_1="Cdk2_CycErem"/>
<map_variables variable_2="Xic" variable_1="Xic"/>
<map_variables variable_2="Cdk2_CycErem" variable_1="Cdk2_CycErem"/>
</connection>
<connection>
<map_components component_2="Xic_Cdk2_CycErem" component_1="Cdk2_CycErem"/>
<map_variables variable_2="Xic_Cdk2_CycErem" variable_1="Xic_Cdk2_CycErem"/>
<map_variables variable_2="Cdk2_CycErem" variable_1="Cdk2_CycErem"/>
</connection>
<connection>
<map_components component_2="Xic" component_1="PCdk2_CycErem"/>
<map_variables variable_2="Xic" variable_1="Xic"/>
<map_variables variable_2="PCdk2_CycErem" variable_1="PCdk2_CycErem"/>
</connection>
<connection>
<map_components component_2="Xic_PCdk2_CycErem" component_1="PCdk2_CycErem"/>
<map_variables variable_2="Xic_PCdk2_CycErem" variable_1="Xic_PCdk2_CycErem"/>
<map_variables variable_2="PCdk2_CycErem" variable_1="PCdk2_CycErem"/>
</connection>
<connection>
<map_components component_2="Kin_" component_1="Wee1_"/>
<map_variables variable_2="Kin_" variable_1="Kin_"/>
</connection>
<connection>
<map_components component_2="Cdk2_CycErem" component_1="Deg_"/>
<map_variables variable_2="Cdk2_CycErem" variable_1="Cdk2_CycErem"/>
</connection>
<connection>
<map_components component_2="Xic_Cdk2_CycE" component_1="Xic"/>
<map_variables variable_2="Xic_Cdk2_CycE" variable_1="Xic_Cdk2_CycE"/>
<map_variables variable_2="Xic" variable_1="Xic"/>
</connection>
<connection>
<map_components component_2="Xic_PCdk2_CycE" component_1="Xic"/>
<map_variables variable_2="Xic_PCdk2_CycE" variable_1="Xic_PCdk2_CycE"/>
<map_variables variable_2="Xic" variable_1="Xic"/>
</connection>
<connection>
<map_components component_2="Xic_Cdk2_CycErem" component_1="Xic"/>
<map_variables variable_2="Xic_Cdk2_CycErem" variable_1="Xic_Cdk2_CycErem"/>
<map_variables variable_2="Xic" variable_1="Xic"/>
</connection>
<connection>
<map_components component_2="Xic_PCdk2_CycErem" component_1="Xic"/>
<map_variables variable_2="Xic_PCdk2_CycErem" variable_1="Xic_PCdk2_CycErem"/>
<map_variables variable_2="Xic" variable_1="Xic"/>
</connection>
<connection>
<map_components component_2="Xic_Cdk2_CycErem" component_1="Xic_Cdk2_CycE"/>
<map_variables variable_2="Xic_Cdk2_CycErem" variable_1="Xic_Cdk2_CycErem"/>
<map_variables variable_2="Xic_Cdk2_CycE" variable_1="Xic_Cdk2_CycE"/>
</connection>
<connection>
<map_components component_2="Xic_PCdk2_CycE" component_1="Xic_Cdk2_CycE"/>
<map_variables variable_2="Xic_PCdk2_CycE" variable_1="Xic_PCdk2_CycE"/>
<map_variables variable_2="Xic_Cdk2_CycE" variable_1="Xic_Cdk2_CycE"/>
</connection>
<connection>
<map_components component_2="Wee1_" component_1="Xic_Cdk2_CycE"/>
<map_variables variable_2="Wee1_" variable_1="Wee1_"/>
</connection>
<connection>
<map_components component_2="Xic_PCdk2_CycErem" component_1="Xic_PCdk2_CycE"/>
<map_variables variable_2="Xic_PCdk2_CycErem" variable_1="Xic_PCdk2_CycErem"/>
<map_variables variable_2="Xic_PCdk2_CycE" variable_1="Xic_PCdk2_CycE"/>
</connection>
<connection>
<map_components component_2="Wee1_" component_1="Xic_PCdk2_CycE"/>
<map_variables variable_2="Wee1_" variable_1="Wee1_"/>
</connection>
<connection>
<map_components component_2="Xic_Cdk2_CycErem" component_1="Xicrem"/>
<map_variables variable_2="Xic_Cdk2_CycErem" variable_1="Xic_Cdk2_CycErem"/>
</connection>
<connection>
<map_components component_2="Xic_PCdk2_CycErem" component_1="Xicrem"/>
<map_variables variable_2="Xic_PCdk2_CycErem" variable_1="Xic_PCdk2_CycErem"/>
</connection>
<connection>
<map_components component_2="kinetic_parameters" component_1="Cdk2_CycE"/>
<map_variables variable_2="kwee" variable_1="kwee"/>
<map_variables variable_2="k25A" variable_1="k25A"/>
<map_variables variable_2="kon" variable_1="kon"/>
<map_variables variable_2="koff" variable_1="koff"/>
<map_variables variable_2="phi" variable_1="phi"/>
<map_variables variable_2="kassoc" variable_1="kassoc"/>
<map_variables variable_2="kdissoc" variable_1="kdissoc"/>
</connection>
<connection>
<map_components component_2="kinetic_parameters" component_1="PCdk2_CycE"/>
<map_variables variable_2="kwee" variable_1="kwee"/>
<map_variables variable_2="k25A" variable_1="k25A"/>
<map_variables variable_2="kon" variable_1="kon"/>
<map_variables variable_2="koff" variable_1="koff"/>
<map_variables variable_2="phi" variable_1="phi"/>
<map_variables variable_2="kassoc" variable_1="kassoc"/>
<map_variables variable_2="kdissoc" variable_1="kdissoc"/>
</connection>
<connection>
<map_components component_2="kinetic_parameters" component_1="Cdk2_CycErem"/>
<map_variables variable_2="kedeg" variable_1="kedeg"/>
<map_variables variable_2="kxdeg" variable_1="kxdeg"/>
<map_variables variable_2="kon" variable_1="kon"/>
<map_variables variable_2="koff" variable_1="koff"/>
<map_variables variable_2="phi" variable_1="phi"/>
<map_variables variable_2="kassoc" variable_1="kassoc"/>
<map_variables variable_2="kdissoc" variable_1="kdissoc"/>
<map_variables variable_2="Deg" variable_1="Deg"/>
<map_variables variable_2="Cdk2_CycErem" variable_1="Cdk2_CycErem"/>
</connection>
<connection>
<map_components component_2="kinetic_parameters" component_1="PCdk2_CycErem"/>
<map_variables variable_2="kedeg" variable_1="kedeg"/>
<map_variables variable_2="kxdeg" variable_1="kxdeg"/>
<map_variables variable_2="kon" variable_1="kon"/>
<map_variables variable_2="koff" variable_1="koff"/>
<map_variables variable_2="phi" variable_1="phi"/>
<map_variables variable_2="kassoc" variable_1="kassoc"/>
<map_variables variable_2="kdissoc" variable_1="kdissoc"/>
<map_variables variable_2="Deg" variable_1="Deg"/>
<map_variables variable_2="PCdk2_CycErem" variable_1="PCdk2_CycErem"/>
</connection>
<connection>
<map_components component_2="kinetic_parameters" component_1="Xic"/>
<map_variables variable_2="kassoc" variable_1="kassoc"/>
<map_variables variable_2="kdissoc" variable_1="kdissoc"/>
</connection>
<connection>
<map_components component_2="kinetic_parameters" component_1="Xic_Cdk2_CycE"/>
<map_variables variable_2="kwee" variable_1="kwee"/>
<map_variables variable_2="k25A" variable_1="k25A"/>
<map_variables variable_2="kon" variable_1="kon"/>
<map_variables variable_2="koff" variable_1="koff"/>
<map_variables variable_2="phi" variable_1="phi"/>
<map_variables variable_2="kassoc" variable_1="kassoc"/>
<map_variables variable_2="kdissoc" variable_1="kdissoc"/>
</connection>
<connection>
<map_components component_2="kinetic_parameters" component_1="Xic_PCdk2_CycE"/>
<map_variables variable_2="kwee" variable_1="kwee"/>
<map_variables variable_2="k25A" variable_1="k25A"/>
<map_variables variable_2="kon" variable_1="kon"/>
<map_variables variable_2="koff" variable_1="koff"/>
<map_variables variable_2="phi" variable_1="phi"/>
<map_variables variable_2="kassoc" variable_1="kassoc"/>
<map_variables variable_2="kdissoc" variable_1="kdissoc"/>
</connection>
<connection>
<map_components component_2="kinetic_parameters" component_1="Xic_Cdk2_CycErem"/>
<map_variables variable_2="kedeg" variable_1="kedeg"/>
<map_variables variable_2="kxdeg" variable_1="kxdeg"/>
<map_variables variable_2="kon" variable_1="kon"/>
<map_variables variable_2="koff" variable_1="koff"/>
<map_variables variable_2="phi" variable_1="phi"/>
<map_variables variable_2="kassoc" variable_1="kassoc"/>
<map_variables variable_2="kdissoc" variable_1="kdissoc"/>
<map_variables variable_2="Deg" variable_1="Deg"/>
<map_variables variable_2="Xic_Cdk2_CycErem" variable_1="Xic_Cdk2_CycErem"/>
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