- Author:
- pmr2.import <nobody@models.cellml.org>
- Date:
- 2007-03-23 04:22:31+12:00
- Desc:
- committing version02 of earm_noble_1990
- Permanent Source URI:
- https://models.physiomeproject.org/workspace/earm_noble_1990/rawfile/c45152decc437f689e2fe1947eb28a14529e03a0/earm_noble_1990.cellml
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<documentation xmlns="http://cellml.org/tmp-documentation">
<article>
<section id="sec_status">
<title>Model Status</title>
<para>
Version 02 was created from Version 01 by James Lawson (03/07) to allow the model to run in PCEnv.
</para>
</section>
<sect1 id="sec_structure">
<title>Model Structure</title>
<para>
This model is based on the paper:
A model of the single atrial cell - relation between calcium current and calcium release"
by Earm Y, Noble D
Proceedings of the Royal Society of London. Series B, Biological Sciences, Vol. 240, No. 1297. (May 22, 1990), pp. 83-96.
ABSTRACT:
"The hypothesis that calcium release from the sarcoplasmic reticulum in cardiac muscle is induced by rises in free cytosolic calcium (Fabiato 1983, Am. J. Physiol 245) allows the possibility that the release could be at least partly regenerative. There would then be a non-linear relation between calcium current and calcium release. We have investigated this possibility in a single-cell version of the rabbit-atrial model developed by Hilgemann & Noble (1987, Proc. R. Soc. Lond. B 230). The model predicts different voltage ranges of activation for calcium-dependent processes (like the sodium-calcium exchange current, contraction or Fura-2 signals) and the calcium current, in agreement with the experimental results obtained by Earm et al. (1990, Proc. R. Soc. Lond. B 240) on exchange current tails, Cannell et al. (1987, Science, Wash. 238) by using Fura-2 signals, and Fedida et al. (1987, J. Physiol., Lond. 385) and Talo et al. (1988, Biology of isolated adult cardiac myocytes) by using contraction. However, when the Fura-2 concentration is sufficiently high (greater than 200 microM) the activation ranges become very similar as the buffering properties of Fura-2 are sufficient to remove the regenerative effect. It is therefore important to allow for the buffering properties of calcium indicators when investigating the correlation between calcium current and calcium release."
</para>
<informalfigure float="0" id="fig_cell_diagram">
<mediaobject>
<imageobject>
<objectinfo>
<title>schematic diagram of a human atrial myocyte</title>
</objectinfo>
<imagedata fileref="earm_1990.png"/>
</imageobject>
</mediaobject>
<caption>A schematic representation of the Earm and Noble 1990 model.</caption>
</informalfigure>
<para>
The complete original paper reference is cited below:
</para>
<para>
<ulink url="http://www.jstor.org/stable/49496?seq=1">A model of the single atrial cell: relation between calcium current and calcium release</ulink>, Y. E. Earm and D. Noble, 1990, <ulink url="http://www.jstor.org.ezproxy.auckland.ac.nz/journals/00804649.html">
<emphasis>Proceedings of the Royal Society of London. Series B, Biological Sciences</emphasis>
</ulink>, 240, 83-96. (A <ulink url="http://www.jstor.org/stable/pdfplus/49496.pdf">PDF version</ulink> of the article is available to subscribers on the journal website.) <ulink url="http://www.ncbi.nlm.nih.gov/pubmed/1972993">PubMed ID: 1972993</ulink>
</para>
</sect1>
</article>
</documentation>
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<ci>E_Ca</ci>
<apply>
<times/>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="dimensionless">0.5</cn>
<ci>R</ci>
<ci>T</ci>
</apply>
<ci>F</ci>
</apply>
<apply>
<ln/>
<apply>
<divide/>
<ci>Ca_o</ci>
<ci>Ca_i</ci>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>i_b_Ca</ci>
<apply>
<times/>
<ci>g_b_Ca</ci>
<apply>
<minus/>
<ci>V</ci>
<ci>E_Ca</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="Na_Ca_exchanger">
<variable units="nanoA" public_interface="out" name="i_NaCa"/>
<variable units="millimolar" name="k_NaCa" initial_value="0.0001"/>
<variable units="dimensionless" name="n_NaCa" initial_value="3"/>
<variable units="millimolar" name="d_NaCa" initial_value="0.0001"/>
<variable units="dimensionless" name="gamma" initial_value="0.5"/>
<variable units="second" public_interface="in" name="time"/>
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="millijoule_per_mole_kelvin" public_interface="in" name="R"/>
<variable units="kelvin" public_interface="in" name="T"/>
<variable units="coulomb_per_mole" public_interface="in" name="F"/>
<variable units="millimolar" public_interface="in" name="Ca_o"/>
<variable units="millimolar" public_interface="in" name="Ca_i"/>
<variable units="millimolar" public_interface="in" name="Na_i"/>
<variable units="millimolar" public_interface="in" name="Na_o"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_NaCa</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>k_NaCa</ci>
<apply>
<minus/>
<apply>
<times/>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<ci>gamma</ci>
<apply>
<minus/>
<ci>n_NaCa</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
<ci>V</ci>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
<apply>
<power/>
<ci>Na_i</ci>
<ci>n_NaCa</ci>
</apply>
<ci>Ca_o</ci>
</apply>
<apply>
<times/>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<minus/>
<ci>gamma</ci>
<cn cellml:units="dimensionless">1</cn>
</apply>
<apply>
<minus/>
<ci>n_NaCa</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
<ci>V</ci>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
<apply>
<power/>
<ci>Na_o</ci>
<ci>n_NaCa</ci>
</apply>
<ci>Ca_i</ci>
</apply>
</apply>
</apply>
<apply>
<times/>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<times/>
<ci>d_NaCa</ci>
<apply>
<plus/>
<apply>
<times/>
<ci>Ca_i</ci>
<apply>
<power/>
<ci>Na_o</ci>
<ci>n_NaCa</ci>
</apply>
</apply>
<apply>
<times/>
<ci>Ca_o</ci>
<apply>
<power/>
<ci>Na_i</ci>
<ci>n_NaCa</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<divide/>
<ci>Ca_i</ci>
<cn cellml:units="millimolar">0.0069</cn>
</apply>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="potassium_background_current">
<variable units="millivolt" public_interface="in" name="E_K"/>
<variable units="nanoA" public_interface="out" name="i_b_K"/>
<variable units="milliS_per_cm2" name="g_b_K" initial_value="0.0017"/>
<variable units="second" public_interface="in" name="time"/>
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="millijoule_per_mole_kelvin" public_interface="in" name="R"/>
<variable units="kelvin" public_interface="in" name="T"/>
<variable units="coulomb_per_mole" public_interface="in" name="F"/>
<variable units="millimolar" public_interface="in" name="K_i"/>
<variable units="millimolar" public_interface="in" name="K_c"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_b_K</ci>
<apply>
<times/>
<ci>g_b_K</ci>
<apply>
<minus/>
<ci>V</ci>
<ci>E_K</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="time_independent_potassium_current">
<variable units="nanoA" public_interface="out" name="i_K1"/>
<variable units="millivolt" public_interface="in" name="E_K"/>
<variable units="milliS_per_cm2" name="g_K1" initial_value="0.017"/>
<variable units="millimolar" name="K_m_K1" initial_value="10"/>
<variable units="second" public_interface="in" name="time"/>
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="millijoule_per_mole_kelvin" public_interface="in" name="R"/>
<variable units="kelvin" public_interface="in" name="T"/>
<variable units="coulomb_per_mole" public_interface="in" name="F"/>
<variable units="millimolar" public_interface="in" name="K_c"/>
<variable units="millimolar" public_interface="in" name="K_i"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_K1</ci>
<apply>
<divide/>
<apply>
<times/>
<apply>
<divide/>
<apply>
<times/>
<ci>g_K1</ci>
<ci>K_c</ci>
</apply>
<apply>
<plus/>
<ci>K_c</ci>
<ci>K_m_K1</ci>
</apply>
</apply>
<apply>
<minus/>
<ci>V</ci>
<ci>E_K</ci>
</apply>
</apply>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<minus/>
<apply>
<minus/>
<ci>V</ci>
<ci>E_K</ci>
</apply>
<cn cellml:units="millivolt">10</cn>
</apply>
<ci>F</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="L_type_calcium_current">
<variable units="nanoA" public_interface="out" name="i_Ca_L"/>
<variable units="nanoA" public_interface="out" name="i_Ca_L_Ca"/>
<variable units="nanoA" public_interface="out" name="i_Ca_L_K"/>
<variable units="nanoA" public_interface="out" name="i_Ca_L_Na"/>
<variable units="dimensionless" name="P_Ca_L" initial_value="0.05"/>
<variable units="second" public_interface="in" private_interface="out" name="time"/>
<variable units="millivolt" public_interface="in" private_interface="out" name="V"/>
<variable units="millimolar" public_interface="in" private_interface="out" name="Ca_i"/>
<variable units="millijoule_per_mole_kelvin" public_interface="in" name="R"/>
<variable units="kelvin" public_interface="in" name="T"/>
<variable units="coulomb_per_mole" public_interface="in" name="F"/>
<variable units="millimolar" public_interface="in" name="Na_i"/>
<variable units="millimolar" public_interface="in" name="Na_o"/>
<variable units="millimolar" public_interface="in" name="Ca_o"/>
<variable units="millimolar" public_interface="in" name="K_i"/>
<variable units="millimolar" public_interface="in" name="K_c"/>
<variable units="dimensionless" private_interface="in" name="d"/>
<variable units="dimensionless" private_interface="in" name="f_Ca"/>
<variable units="dimensionless" private_interface="in" name="CaChon"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_Ca_L</ci>
<apply>
<plus/>
<ci>i_Ca_L_Ca</ci>
<ci>i_Ca_L_K</ci>
<ci>i_Ca_L_Na</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>i_Ca_L_Ca</ci>
<apply>
<times/>
<apply>
<divide/>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>P_Ca_L</ci>
<ci>d</ci>
<ci>CaChon</ci>
<apply>
<minus/>
<ci>V</ci>
<cn cellml:units="millivolt">50</cn>
</apply>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<minus/>
<apply>
<minus/>
<ci>V</ci>
<cn cellml:units="millivolt">50</cn>
</apply>
</apply>
<ci>F</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>Ca_i</ci>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="dimensionless">100</cn>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
</apply>
<apply>
<times/>
<ci>Ca_o</ci>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<minus/>
<apply>
<minus/>
<ci>V</ci>
<cn cellml:units="millivolt">50</cn>
</apply>
</apply>
<ci>F</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>i_Ca_L_K</ci>
<apply>
<times/>
<apply>
<divide/>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="dimensionless">0.002</cn>
<ci>P_Ca_L</ci>
<ci>d</ci>
<ci>CaChon</ci>
<apply>
<minus/>
<ci>V</ci>
<cn cellml:units="millivolt">50</cn>
</apply>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<minus/>
<apply>
<minus/>
<ci>V</ci>
<cn cellml:units="millivolt">50</cn>
</apply>
</apply>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>K_i</ci>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="dimensionless">50</cn>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
</apply>
<apply>
<times/>
<ci>K_c</ci>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<minus/>
<apply>
<minus/>
<ci>V</ci>
<cn cellml:units="millivolt">50</cn>
</apply>
</apply>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>i_Ca_L_Na</ci>
<apply>
<times/>
<apply>
<divide/>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="dimensionless">0.01</cn>
<ci>P_Ca_L</ci>
<ci>d</ci>
<ci>CaChon</ci>
<apply>
<minus/>
<ci>V</ci>
<cn cellml:units="millivolt">50</cn>
</apply>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<minus/>
<apply>
<minus/>
<ci>V</ci>
<cn cellml:units="millivolt">50</cn>
</apply>
</apply>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>Na_i</ci>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="dimensionless">50</cn>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
</apply>
<apply>
<times/>
<ci>Na_o</ci>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<minus/>
<apply>
<minus/>
<ci>V</ci>
<cn cellml:units="millivolt">50</cn>
</apply>
</apply>
<ci>F</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="L_type_calcium_current_d_gate">
<variable units="dimensionless" public_interface="out" name="d" initial_value="0.0011"/>
<variable units="per_second" name="alpha_d"/>
<variable units="per_second" name="beta_d"/>
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="second" public_interface="in" name="time"/>
<variable units="millivolt" name="E0_d"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>E0_d</ci>
<apply>
<plus/>
<ci>V</ci>
<cn cellml:units="millivolt">19</cn>
</apply>
</apply>
<apply>
<eq/>
<ci>alpha_d</ci>
<piecewise>
<piece>
<cn cellml:units="per_second">120</cn>
<apply>
<lt/>
<apply>
<abs/>
<ci>E0_d</ci>
</apply>
<cn cellml:units="millivolt">0.0001</cn>
</apply>
</piece>
<otherwise>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="per_millivolt_second">30</cn>
<ci>E0_d</ci>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<ci>E0_d</ci>
</apply>
<cn cellml:units="millivolt">4</cn>
</apply>
</apply>
</apply>
</apply>
</otherwise>
</piecewise>
</apply>
<apply>
<eq/>
<ci>beta_d</ci>
<piecewise>
<piece>
<cn cellml:units="per_second">120</cn>
<apply>
<lt/>
<apply>
<abs/>
<ci>E0_d</ci>
</apply>
<cn cellml:units="millivolt">0.0001</cn>
</apply>
</piece>
<otherwise>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="per_millivolt_second">12</cn>
<ci>E0_d</ci>
</apply>
<apply>
<minus/>
<apply>
<exp/>
<apply>
<divide/>
<ci>E0_d</ci>
<cn cellml:units="millivolt">10</cn>
</apply>
</apply>
<cn cellml:units="dimensionless">1</cn>
</apply>
</apply>
</otherwise>
</piecewise>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>d</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>alpha_d</ci>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>d</ci>
</apply>
</apply>
<apply>
<times/>
<ci>beta_d</ci>
<ci>d</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="L_type_calcium_current_f_Ca_gate">
<variable units="dimensionless" public_interface="out" name="f_Ca" initial_value="0.785"/>
<variable units="per_second" name="alpha_f_Ca"/>
<variable units="per_second" name="beta_f_Ca"/>
<variable units="dimensionless" name="CaChoff"/>
<variable units="dimensionless" public_interface="out" name="CaChon"/>
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="second" public_interface="in" name="time"/>
<variable units="millimolar" public_interface="in" name="Ca_i"/>
<variable units="millivolt" name="E0_f"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>E0_f</ci>
<apply>
<plus/>
<ci>V</ci>
<cn cellml:units="millivolt">34</cn>
</apply>
</apply>
<apply>
<eq/>
<ci>alpha_f_Ca</ci>
<piecewise>
<piece>
<cn cellml:units="per_second">25</cn>
<apply>
<lt/>
<apply>
<abs/>
<ci>E0_f</ci>
</apply>
<cn cellml:units="millivolt">0.0001</cn>
</apply>
</piece>
<otherwise>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="per_millivolt_second">6.25</cn>
<ci>E0_f</ci>
</apply>
<apply>
<minus/>
<apply>
<exp/>
<apply>
<divide/>
<ci>E0_f</ci>
<cn cellml:units="millivolt">4</cn>
</apply>
</apply>
<cn cellml:units="dimensionless">1</cn>
</apply>
</apply>
</otherwise>
</piecewise>
</apply>
<apply>
<eq/>
<ci>beta_f_Ca</ci>
<apply>
<divide/>
<cn cellml:units="per_second">12</cn>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<ci>E0_f</ci>
</apply>
<cn cellml:units="millivolt">4</cn>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>f_Ca</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="per_second">120</cn>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>f_Ca</ci>
</apply>
<ci>CaChoff</ci>
</apply>
<apply>
<times/>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>f_Ca</ci>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>CaChoff</ci>
</apply>
</apply>
</apply>
<ci>beta_f_Ca</ci>
</apply>
<apply>
<times/>
<ci>alpha_f_Ca</ci>
<ci>f_Ca</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>CaChoff</ci>
<apply>
<divide/>
<ci>Ca_i</ci>
<apply>
<plus/>
<cn cellml:units="dimensionless">0.001</cn>
<ci>Ca_i</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>CaChon</ci>
<apply>
<times/>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>f_Ca</ci>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>CaChoff</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="sarcoplasmic_reticulum_calcium_pump">
<variable units="millimolar_per_second" public_interface="out" name="i_up"/>
<variable units="per_second" name="K_1"/>
<variable units="per_second" name="K_2"/>
<variable units="millimolar" name="K_cyca" initial_value="0.0003"/>
<variable units="millimolar" name="K_xcs" initial_value="0.4"/>
<variable units="dimensionless" name="K_srca" initial_value="0.5"/>
<variable units="per_second" name="alpha_up" initial_value="3"/>
<variable units="per_second" name="beta_up" initial_value="0.23"/>
<variable units="millimolar" public_interface="in" name="Ca_up"/>
<variable units="second" public_interface="in" name="time"/>
<variable units="millimolar" public_interface="in" name="Ca_i"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>K_1</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>K_cyca</ci>
<ci>K_xcs</ci>
</apply>
<ci>K_srca</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>K_2</ci>
<apply>
<plus/>
<ci>Ca_i</ci>
<apply>
<times/>
<ci>Ca_up</ci>
<ci>K_1</ci>
</apply>
<apply>
<times/>
<ci>K_cyca</ci>
<ci>K_xcs</ci>
</apply>
<ci>K_cyca</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>i_up</ci>
<apply>
<minus/>
<apply>
<times/>
<apply>
<divide/>
<ci>Ca_i</ci>
<ci>K_2</ci>
</apply>
<ci>alpha_up</ci>
</apply>
<apply>
<times/>
<apply>
<divide/>
<apply>
<times/>
<ci>Ca_up</ci>
<ci>K_1</ci>
</apply>
<ci>K_2</ci>
</apply>
<ci>beta_up</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="calcium_release">
<variable units="millimolar_per_second" public_interface="out" name="i_rel"/>
<variable units="millivolt" name="VoltDep"/>
<variable units="dimensionless" name="RegBindSite"/>
<variable units="per_second" name="ActRate"/>
<variable units="per_second" name="InactRate"/>
<variable units="per_second" name="K_leak_rate" initial_value="0"/>
<variable units="per_second" name="K_m_rel" initial_value="250"/>
<variable units="dimensionless" name="PrecFrac"/>
<variable units="dimensionless" name="ActFrac" initial_value="0"/>
<variable units="dimensionless" name="ProdFrac" initial_value="0"/>
<variable units="second" public_interface="in" name="time"/>
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="millimolar" public_interface="in" name="Ca_i"/>
<variable units="millimolar" public_interface="in" name="Ca_rel"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>PrecFrac</ci>
<apply>
<minus/>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>ActFrac</ci>
</apply>
<ci>ProdFrac</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>VoltDep</ci>
<apply>
<exp/>
<apply>
<times/>
<cn cellml:units="dimensionless">0.08</cn>
<apply>
<minus/>
<ci>V</ci>
<cn cellml:units="millivolt">40</cn>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>RegBindSite</ci>
<apply>
<power/>
<apply>
<divide/>
<ci>Ca_i</ci>
<apply>
<plus/>
<ci>Ca_i</ci>
<cn cellml:units="millimolar">0.0005</cn>
</apply>
</apply>
<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
<apply>
<eq/>
<ci>ActRate</ci>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="per_millivolt_second">600</cn>
<ci>VoltDep</ci>
</apply>
<apply>
<times/>
<cn cellml:units="per_second">500</cn>
<ci>RegBindSite</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>InactRate</ci>
<apply>
<plus/>
<cn cellml:units="per_second">60</cn>
<apply>
<times/>
<cn cellml:units="per_second">500</cn>
<ci>RegBindSite</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>ActFrac</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>PrecFrac</ci>
<ci>ActRate</ci>
</apply>
<apply>
<times/>
<ci>ActFrac</ci>
<ci>InactRate</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>ProdFrac</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>ActFrac</ci>
<ci>InactRate</ci>
</apply>
<apply>
<times/>
<cn cellml:units="per_second">0.6</cn>
<ci>ProdFrac</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>i_rel</ci>
<apply>
<times/>
<apply>
<plus/>
<apply>
<times/>
<apply>
<power/>
<apply>
<divide/>
<ci>ActFrac</ci>
<apply>
<plus/>
<ci>ActFrac</ci>
<cn cellml:units="dimensionless">0.25</cn>
</apply>
</apply>
<cn cellml:units="dimensionless">2</cn>
</apply>
<ci>K_m_rel</ci>
</apply>
<ci>K_leak_rate</ci>
</apply>
<ci>Ca_rel</ci>
</apply>
</apply>
</math>
</component>
<component name="calcium_translocation">
<variable units="millimolar_per_second" public_interface="out" name="i_trans"/>
<variable units="millimolar" public_interface="in" name="Ca_rel"/>
<variable units="millimolar" public_interface="in" name="Ca_up"/>
<variable units="per_second" name="alpha_tr" initial_value="50"/>
<variable units="second" public_interface="in" name="time"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_trans</ci>
<apply>
<times/>
<apply>
<minus/>
<ci>Ca_up</ci>
<ci>Ca_rel</ci>
</apply>
<ci>alpha_tr</ci>
</apply>
</apply>
</math>
</component>
<component name="extracellular_sodium_concentration">
<variable units="millimolar" public_interface="out" name="Na_o" initial_value="140"/>
</component>
<component name="intracellular_sodium_concentration">
<variable units="millimolar" public_interface="out" name="Na_i" initial_value="6.48"/>
<variable units="microlitre" public_interface="in" name="V_i"/>
<variable units="second" public_interface="in" name="time"/>
<variable units="coulomb_per_mole" public_interface="in" name="F"/>
<variable units="nanoA" public_interface="in" name="i_Na"/>
<variable units="nanoA" public_interface="in" name="i_b_Na"/>
<variable units="nanoA" public_interface="in" name="i_Ca_L_Na"/>
<variable units="nanoA" public_interface="in" name="i_NaK"/>
<variable units="nanoA" public_interface="in" name="i_NaCa"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Na_i</ci>
</apply>
<apply>
<times/>
<apply>
<divide/>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
</apply>
<apply>
<times/>
<ci>V_i</ci>
<ci>F</ci>
</apply>
</apply>
<apply>
<plus/>
<ci>i_Na</ci>
<ci>i_b_Na</ci>
<apply>
<times/>
<cn cellml:units="dimensionless">3</cn>
<ci>i_NaK</ci>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">3</cn>
<ci>i_NaCa</ci>
</apply>
<ci>i_Ca_L_Na</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="extracellular_calcium_concentration">
<variable units="millimolar" public_interface="out" name="Ca_o" initial_value="2"/>
</component>
<component name="extracellular_potassium_concentration">
<variable units="millimolar" public_interface="out" name="K_c" initial_value="4"/>
</component>
<component name="intracellular_potassium_concentration">
<variable units="millimolar" public_interface="out" name="K_i" initial_value="140"/>
<variable units="microlitre" public_interface="in" name="V_i"/>
<variable units="nanoA" public_interface="in" name="i_K1"/>
<variable units="nanoA" public_interface="in" name="i_Ca_L_K"/>
<variable units="nanoA" public_interface="in" name="i_to"/>
<variable units="nanoA" public_interface="in" name="i_b_K"/>
<variable units="nanoA" public_interface="in" name="i_NaK"/>
<variable units="second" public_interface="in" name="time"/>
<variable units="coulomb_per_mole" public_interface="in" name="F"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>K_i</ci>
</apply>
<apply>
<times/>
<apply>
<divide/>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
</apply>
<apply>
<times/>
<ci>V_i</ci>
<ci>F</ci>
</apply>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<ci>i_K1</ci>
<ci>i_Ca_L_K</ci>
<ci>i_to</ci>
<ci>i_b_K</ci>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>i_NaK</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component name="intracellular_calcium_concentration">
<variable units="millimolar" public_interface="out" name="Ca_i" initial_value="1e-5"/>
<variable units="millimolar" public_interface="out" name="Ca_up" initial_value="0.3"/>
<variable units="millimolar" public_interface="out" name="Ca_rel" initial_value="0.3"/>
<variable units="millimolar" name="Ca_Calmod" initial_value="0.0005"/>
<variable units="millimolar" name="Ca_Trop" initial_value="0.0015"/>
<variable units="millimolar" name="Calmod" initial_value="0.02"/>
<variable units="millimolar" name="Trop" initial_value="0.15"/>
<variable units="per_second" name="alpha_Calmod" initial_value="100000"/>
<variable units="per_second" name="beta_Calmod" initial_value="50"/>
<variable units="per_second" name="alpha_Trop" initial_value="100000"/>
<variable units="per_second" name="beta_Trop" initial_value="200"/>
<variable units="micrometre" name="radius" initial_value="10"/>
<variable units="micrometre" name="length" initial_value="80"/>
<variable units="microlitre" public_interface="out" name="V_i"/>
<variable units="microlitre" name="V_Cell"/>
<variable units="dimensionless" name="V_i_ratio"/>
<variable units="dimensionless" name="V_rel_ratio" initial_value="0.1"/>
<variable units="dimensionless" name="V_e_ratio" initial_value="0.4"/>
<variable units="dimensionless" name="V_up_ratio" initial_value="0.01"/>
<variable units="millimolar_per_second" public_interface="in" name="i_up"/>
<variable units="millimolar_per_second" public_interface="in" name="i_trans"/>
<variable units="millimolar_per_second" public_interface="in" name="i_rel"/>
<variable units="nanoA" public_interface="in" name="i_NaCa"/>
<variable units="nanoA" public_interface="in" name="i_Ca_L_Ca"/>
<variable units="nanoA" public_interface="in" name="i_b_Ca"/>
<variable units="coulomb_per_mole" public_interface="in" name="F"/>
<variable units="second" public_interface="in" name="time"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>V_Cell</ci>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="dimensionless">3.141592654</cn>
<apply>
<power/>
<apply>
<divide/>
<ci>radius</ci>
<cn cellml:units="dimensionless">1000</cn>
</apply>
<cn cellml:units="dimensionless">2</cn>
</apply>
<ci>length</ci>
</apply>
<cn cellml:units="dimensionless">1000</cn>
</apply>
</apply>
<apply>
<eq/>
<ci>V_i_ratio</ci>
<apply>
<minus/>
<apply>
<minus/>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>V_e_ratio</ci>
</apply>
<ci>V_up_ratio</ci>
</apply>
<ci>V_rel_ratio</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>V_i</ci>
<apply>
<times/>
<ci>V_Cell</ci>
<ci>V_i_ratio</ci>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Ca_i</ci>
</apply>
<apply>
<minus/>
<apply>
<minus/>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<apply>
<divide/>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
</apply>
<cn cellml:units="dimensionless">2</cn>
</apply>
<ci>V_i</ci>
<ci>F</ci>
<apply>
<minus/>
<apply>
<plus/>
<ci>i_Ca_L_Ca</ci>
<ci>i_b_Ca</ci>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>i_NaCa</ci>
</apply>
</apply>
</apply>
<apply>
<divide/>
<apply>
<times/>
<ci>i_rel</ci>
<ci>V_rel_ratio</ci>
</apply>
<ci>V_i_ratio</ci>
</apply>
</apply>
<ci>dCa_Calmod_dtime</ci>
</apply>
<ci>dCa_Trop_dtime</ci>
</apply>
<ci>i_up</ci>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Ca_up</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<apply>
<divide/>
<ci>V_i_ratio</ci>
<ci>V_up_ratio</ci>
</apply>
<ci>i_up</ci>
</apply>
<ci>i_trans</ci>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Ca_rel</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<apply>
<divide/>
<ci>V_up_ratio</ci>
<ci>V_rel_ratio</ci>
</apply>
<ci>i_trans</ci>
</apply>
<ci>i_rel</ci>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Ca_Calmod</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>alpha_Calmod</ci>
<ci>Ca_i</ci>
<apply>
<minus/>
<ci>Calmod</ci>
<ci>Ca_Calmod</ci>
</apply>
</apply>
<apply>
<times/>
<ci>beta_Calmod</ci>
<ci>Ca_Calmod</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Ca_Trop</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>alpha_Trop</ci>
<ci>Ca_i</ci>
<apply>
<minus/>
<ci>Trop</ci>
<ci>Ca_Trop</ci>
</apply>
</apply>
<apply>
<times/>
<ci>beta_Trop</ci>
<ci>Ca_Trop</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>dCa_Calmod_dtime</ci>
<apply>
<minus/>
<apply>
<times/>
<ci>alpha_Calmod</ci>
<ci>Ca_i</ci>
<apply>
<minus/>
<ci>Calmod</ci>
<ci>Ca_Calmod</ci>
</apply>
</apply>
<apply>
<times/>
<ci>beta_Calmod</ci>
<ci>Ca_Calmod</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>dCa_Trop_dtime</ci>
<apply>
<minus/>
<apply>
<times/>
<ci>alpha_Trop</ci>
<ci>Ca_i</ci>
<apply>
<minus/>
<ci>Trop</ci>
<ci>Ca_Trop</ci>
</apply>
</apply>
<apply>
<times/>
<ci>beta_Trop</ci>
<ci>Ca_Trop</ci>
</apply>
</apply>
</apply>
</math>
<variable units="millimolar_per_second" name="dCa_Calmod_dtime"/>
<variable units="millimolar_per_second" name="dCa_Trop_dtime"/>
</component>
<group>
<relationship_ref relationship="containment"/>
<component_ref component="membrane">
<component_ref component="fast_sodium_current">
<component_ref component="fast_sodium_current_m_gate"/>
<component_ref component="fast_sodium_current_h_gate"/>
</component_ref>
<component_ref component="sodium_potassium_pump"/>
<component_ref component="sodium_background_current"/>
<component_ref component="calcium_background_current"/>
<component_ref component="Na_Ca_exchanger"/>
<component_ref component="potassium_background_current"/>
<component_ref component="time_independent_potassium_current"/>
<component_ref component="transient_outward_current">
<component_ref component="transient_outward_current_s_gate"/>
<component_ref component="transient_outward_current_r_gate"/>
</component_ref>
<component_ref component="L_type_calcium_current">
<component_ref component="L_type_calcium_current_d_gate"/>
<component_ref component="L_type_calcium_current_f_Ca_gate"/>
</component_ref>
<component_ref component="sarcoplasmic_reticulum_calcium_pump"/>
<component_ref component="calcium_release"/>
<component_ref component="calcium_translocation"/>
<component_ref component="extracellular_potassium_concentration"/>
<component_ref component="intracellular_potassium_concentration"/>
<component_ref component="extracellular_sodium_concentration"/>
<component_ref component="intracellular_sodium_concentration"/>
<component_ref component="extracellular_calcium_concentration"/>
<component_ref component="intracellular_calcium_concentration"/>
</component_ref>
</group>
<group>
<relationship_ref relationship="encapsulation"/>
<component_ref component="fast_sodium_current">
<component_ref component="fast_sodium_current_m_gate"/>
<component_ref component="fast_sodium_current_h_gate"/>
</component_ref>
<component_ref component="transient_outward_current">
<component_ref component="transient_outward_current_r_gate"/>
<component_ref component="transient_outward_current_s_gate"/>
</component_ref>
<component_ref component="L_type_calcium_current">
<component_ref component="L_type_calcium_current_d_gate"/>
<component_ref component="L_type_calcium_current_f_Ca_gate"/>
</component_ref>
</group>
<connection>
<map_components component_2="environment" component_1="membrane"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="intracellular_sodium_concentration" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="intracellular_potassium_concentration" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="intracellular_calcium_concentration" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="fast_sodium_current"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="sodium_potassium_pump"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="sodium_background_current"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="calcium_background_current"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="Na_Ca_exchanger"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="potassium_background_current"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="time_independent_potassium_current"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="transient_outward_current"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="L_type_calcium_current"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="sarcoplasmic_reticulum_calcium_pump"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="calcium_release"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="environment" component_1="calcium_translocation"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="membrane" component_1="fast_sodium_current"/>
<map_variables variable_2="i_Na" variable_1="i_Na"/>
<map_variables variable_2="V" variable_1="V"/>
<map_variables variable_2="R" variable_1="R"/>
<map_variables variable_2="F" variable_1="F"/>
<map_variables variable_2="T" variable_1="T"/>
</connection>
<connection>
<map_components component_2="membrane" component_1="transient_outward_current"/>
<map_variables variable_2="i_to" variable_1="i_to"/>
<map_variables variable_2="V" variable_1="V"/>
<map_variables variable_2="R" variable_1="R"/>
<map_variables variable_2="F" variable_1="F"/>
<map_variables variable_2="T" variable_1="T"/>
</connection>
<connection>
<map_components component_2="membrane" component_1="sodium_potassium_pump"/>
<map_variables variable_2="i_NaK" variable_1="i_NaK"/>
<map_variables variable_2="V" variable_1="V"/>
</connection>
<connection>
<map_components component_2="membrane" component_1="sodium_background_current"/>
<map_variables variable_2="i_b_Na" variable_1="i_b_Na"/>
<map_variables variable_2="V" variable_1="V"/>
<map_variables variable_2="R" variable_1="R"/>
<map_variables variable_2="F" variable_1="F"/>
<map_variables variable_2="T" variable_1="T"/>
</connection>
<connection>
<map_components component_2="membrane" component_1="calcium_background_current"/>
<map_variables variable_2="i_b_Ca" variable_1="i_b_Ca"/>
<map_variables variable_2="V" variable_1="V"/>
<map_variables variable_2="R" variable_1="R"/>
<map_variables variable_2="F" variable_1="F"/>
<map_variables variable_2="T" variable_1="T"/>
</connection>
<connection>
<map_components component_2="membrane" component_1="Na_Ca_exchanger"/>
<map_variables variable_2="V" variable_1="V"/>
<map_variables variable_2="i_NaCa" variable_1="i_NaCa"/>
<map_variables variable_2="R" variable_1="R"/>
<map_variables variable_2="F" variable_1="F"/>
<map_variables variable_2="T" variable_1="T"/>
</connection>
<connection>
<map_components component_2="membrane" component_1="potassium_background_current"/>
<map_variables variable_2="i_b_K" variable_1="i_b_K"/>
<map_variables variable_2="V" variable_1="V"/>
<map_variables variable_2="R" variable_1="R"/>
<map_variables variable_2="F" variable_1="F"/>
<map_variables variable_2="T" variable_1="T"/>
</connection>
<connection>
<map_components component_2="membrane" component_1="time_independent_potassium_current"/>
<map_variables variable_2="i_K1" variable_1="i_K1"/>
<map_variables variable_2="V" variable_1="V"/>
<map_variables variable_2="R" variable_1="R"/>
<map_variables variable_2="F" variable_1="F"/>
<map_variables variable_2="T" variable_1="T"/>
</connection>
<connection>
<map_components component_2="membrane" component_1="L_type_calcium_current"/>
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