- Author:
- Tessa Paris <tpar054@aucklanduni.ac.nz>
- Date:
- 2010-02-19 14:26:51+13:00
- Desc:
- Changed the model abstract
- Permanent Source URI:
- https://staging.physiomeproject.org/workspace/bertram_arnot_zamponi_2002/rawfile/a95d0b851e656802444c4ccd442403c6356ea2e3/bertram_arnot_zamponi_2002.cellml
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COR (0.9.31.1333)
Copyright 2002-2009 Dr Alan Garny
http://cor.physiol.ox.ac.uk/ - cor@physiol.ox.ac.uk
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http://www.cellml.org/
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<title>G-Protein Specificity In Synaptic Signalling</title>
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<section id="sec_status">
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This model was created by James Lawson on 21/05/07 and has been modified by Tessa Paris on 15/12/09.
<para>At present, this model is not able to reproduce the results in the publication. The the presynaptic and postsynaptic cell models have been created but the model still does not run correctly.ValidateCellML verifies this model as valid CellML, although a warning is given as there are equations where the dimensions are equivalent but the units are not.
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<sect1 id="sec_structure">
<title>Model Structure</title>
<para>
<para>ABSTRACT:
</para>
Computational modeling is used to investigate the functional impact of G protein-mediated presynaptic autoinhibition on synaptic filtering properties. It is demonstrated that this form of autoinhibition, which is relieved by depolarization, acts as a high-pass filter. This contrasts with vesicle depletion, which acts as a low-pass filter. Model parameters are adjusted to reproduce kinetic slowing data from different Gbetagamma dimeric isoforms, which produce different degrees of slowing. With these sets of parameter values, we demonstrate that the range of frequencies filtered out by the autoinhibition varies greatly depending on the Gbetagamma isoform activated by the autoreceptors. It is shown that G protein autoinhibition can enhance the spatial contrast between a spatially distributed high-frequency signal and surrounding low-frequency noise, providing an alternate mechanism to lateral inhibition. It is also shown that autoinhibition can increase the fidelity of coincidence detection by increasing the signal-to-noise ratio in the postsynaptic cell. The filter cut, the input frequency below which signals are filtered, depends on several biophysical parameters in addition to those related to Gbetagamma binding and unbinding. By varying one such parameter, the rate at which transmitter unbinds from autoreceptors, we show that the filter cut can be adjusted up or down for several of the Gbetagamma isoforms. This allows for great synapse-to-synapse variability in the distinction between signal and noise. This mathematical model has been translated into a CellML description which can be downloaded in various formats as described in <xref linkend="sec_download_this_model"/>.
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<para>
The complete original paper reference is cited below:
</para>
<para>
Role for G Protein G-Beta-Gamma Isoform Specificity in Synaptic Signal Processing: A Computational Study, Richard Bertram, Michelle I. Arnot, and Gerald W. Zamponi, 2002,
<emphasis>Journal of Neurophysiology</emphasis>
, 87, 2612-2623. <ulink url="http://www.ncbi.nlm.nih.gov/pubmed/11976397?dopt=Abstract">PubMed ID: 11976397</ulink>
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<para>
G-protein autoinhibitory feedback on the presynaptic terminal acts like a high-pass filter, allowing only high-frequency signals to pass through the to the postsynaptic cell. Low-frequency signals are effectively filtered out. Model simulations in this study show how different G-beta-gamma isoforms have different filtering properties. They also emphasise that the different filtering characteristics associated with a specific G-beta-gamma subunit depend on many biophysical parameters, such as the unbinding rate of a transmitter molecule from the presynaptic autoreceptor. For example faster unbinding lowers the filter cut while slower unbinding raises it. This allows for great synapse-tot-synapse variability in the distinction between signal and background noise.
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<variable name="Cm" units="uF_per_cm2" initial_value="1" public_interface="out"/>
<variable name="i_app" units="uA_per_cm2" public_interface="in"/>
<variable name="i_Na" units="uA_per_cm2" public_interface="in"/>
<variable name="i_K" units="uA_per_cm2" public_interface="in"/>
<variable name="i_leak" units="uA_per_cm2" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>V</ci>
</apply>
<apply>
<divide/>
<apply>
<minus/>
<apply>
<minus/>
<apply>
<plus/>
<ci>i_Na</ci>
<ci>i_K</ci>
<ci>i_leak</ci>
</apply>
<ci>i_app</ci>
</apply>
</apply>
<ci>Cm</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="stimulus_protocol">
<variable name="Istim" units="uA_per_cm2" public_interface="out"/>
<variable name="IstimStart" units="ms" initial_value="10"/>
<variable name="IstimEnd" units="ms" initial_value="50000"/>
<variable name="IstimAmplitude" units="uA_per_cm2" initial_value="40"/>
<variable name="IstimPeriod" units="ms" initial_value="300"/>
<variable name="IstimPulseDuration" units="ms" initial_value="1"/>
<variable name="time" units="s" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>Istim</ci>
<piecewise>
<piece>
<ci>IstimAmplitude</ci>
<apply>
<and/>
<apply>
<geq/>
<ci>time</ci>
<ci>IstimStart</ci>
</apply>
<apply>
<leq/>
<ci>time</ci>
<ci>IstimEnd</ci>
</apply>
<apply>
<leq/>
<apply>
<minus/>
<apply>
<minus/>
<ci>time</ci>
<ci>IstimStart</ci>
</apply>
<apply>
<times/>
<apply>
<floor/>
<apply>
<divide/>
<apply>
<minus/>
<ci>time</ci>
<ci>IstimStart</ci>
</apply>
<ci>IstimPeriod</ci>
</apply>
</apply>
<ci>IstimPeriod</ci>
</apply>
</apply>
<ci>IstimPulseDuration</ci>
</apply>
</apply>
</piece>
<otherwise>
<cn cellml:units="uA_per_cm2">0</cn>
</otherwise>
</piecewise>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="sodium_current">
<variable name="i_Na" units="uA_per_cm2" public_interface="out"/>
<variable name="x_infinity" units="dimensionless" public_interface="out"/>
<variable name="time" units="s" public_interface="in"/>
<variable name="V" units="mV" public_interface="in"/>
<variable name="n" units="dimensionless" public_interface="in"/>
<variable name="alpha_x" units="dimensionless" public_interface="out"/>
<variable name="beta_x" units="dimensionless" public_interface="out"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_Na</ci>
<apply>
<times/>
<cn cellml:units="pS_per_cm2">120</cn>
<apply>
<power/>
<ci>x_infinity</ci>
<cn cellml:units="dimensionless">3</cn>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>n</ci>
</apply>
<apply>
<minus/>
<ci>V</ci>
<cn cellml:units="mV">120</cn>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>x_infinity</ci>
<apply>
<divide/>
<ci>alpha_x</ci>
<apply>
<plus/>
<ci>alpha_x</ci>
<ci>beta_x</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>alpha_x</ci>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="dimensionless">0.2</cn>
<apply>
<plus/>
<ci>V</ci>
<cn cellml:units="mV">40</cn>
</apply>
</apply>
<apply>
<minus/>
<cn cellml:units="mV">1</cn>
<apply>
<times/>
<cn cellml:units="mV">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<apply>
<plus/>
<ci>V</ci>
<cn cellml:units="mV">40</cn>
</apply>
</apply>
<cn cellml:units="mV">10</cn>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>beta_x</ci>
<apply>
<times/>
<cn cellml:units="dimensionless">8</cn>
<apply>
<exp/>
<apply>
<divide/>
<cn cellml:units="mV">1</cn>
<apply>
<minus/>
<apply>
<plus/>
<ci>V</ci>
<apply>
<divide/>
<cn cellml:units="mV">65</cn>
<cn cellml:units="dimensionless">18</cn>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="potassium_current">
<variable name="i_K" units="uA_per_cm2" public_interface="out"/>
<variable name="time" units="s" public_interface="in"/>
<variable name="V" units="mV" public_interface="in"/>
<variable name="n" units="dimensionless" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_K</ci>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="pS">36</cn>
<apply>
<power/>
<ci>n</ci>
<cn cellml:units="dimensionless">4</cn>
</apply>
<apply>
<plus/>
<ci>V</ci>
<cn cellml:units="mV">77</cn>
</apply>
</apply>
<cn cellml:units="cm2">1</cn>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="potassium_current_n_gate">
<variable name="n" units="dimensionless" initial_value="0" public_interface="out"/>
<variable name="alpha_n" units="per_ms" public_interface="out"/>
<variable name="beta_n" units="per_ms" public_interface="out"/>
<variable name="V" units="mV" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>n</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>alpha_n</ci>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>n</ci>
</apply>
</apply>
<apply>
<times/>
<ci>beta_n</ci>
<ci>n</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>alpha_n</ci>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="per_mV_per_ms">0.02</cn>
<apply>
<plus/>
<ci>V</ci>
<cn cellml:units="mV">55</cn>
</apply>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<apply>
<plus/>
<ci>V</ci>
<cn cellml:units="mV">55</cn>
</apply>
</apply>
<cn cellml:units="mV">10</cn>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>beta_n</ci>
<apply>
<times/>
<cn cellml:units="per_ms">0.25</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<apply>
<plus/>
<ci>V</ci>
<cn cellml:units="mV">65</cn>
</apply>
</apply>
<cn cellml:units="mV">80</cn>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="leak_current">
<variable name="i_leak" units="uA_per_cm2" public_interface="out"/>
<variable name="time" units="s" public_interface="in"/>
<variable name="V" units="mV" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_leak</ci>
<apply>
<times/>
<cn cellml:units="pS_per_cm2">0.3</cn>
<apply>
<plus/>
<ci>V</ci>
<cn cellml:units="mV">54</cn>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="transmitter_release">
<variable name="R" units="dimensionless" initial_value="0" public_interface="out"/>
<variable name="kr_plus" units="per_mM_per_ms" initial_value="0.00015"/>
<variable name="kr_minus" units="per_ms" initial_value="2.5"/>
<variable name="Ca" units="uM" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<variable name="T" units="mM" public_interface="out"/>
<variable name="T_bar" units="mM" initial_value="4"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>R</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>kr_plus</ci>
<ci>Ca</ci>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>R</ci>
</apply>
</apply>
<apply>
<times/>
<ci>kr_minus</ci>
<ci>R</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>T</ci>
<apply>
<times/>
<ci>T_bar</ci>
<ci>R</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="calcium_concentration">
<variable name="Ca" units="uM" public_interface="out"/>
<variable name="Ca_ex" units="mM" initial_value="2"/>
<variable name="Ca_open" units="uM"/>
<variable name="Dc" units="um2_per_second" initial_value="220"/>
<variable name="r" units="nm" initial_value="10"/>
<variable name="sigma" units="uM_per_ms"/>
<variable name="i_V" units="uA"/>
<variable name="g_Ca" units="pS" initial_value="1.2"/>
<variable name="P" units="mV_per_mM" initial_value="6"/>
<variable name="R_gas_const" units="millijoule_per_mole_kelvin" public_interface="in"/>
<variable name="F" units="coulomb_per_mole" public_interface="in"/>
<variable name="Temp" units="kelvin" public_interface="in"/>
<variable name="V" units="mV" public_interface="in"/>
<variable name="O" units="dimensionless" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>Ca_open</ci>
<apply>
<divide/>
<ci>sigma</ci>
<apply>
<times/>
<cn cellml:units="per_um3">2</cn>
<ci>Dc</ci>
<ci>r</ci>
<pi/>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>sigma</ci>
<apply>
<times/>
<apply>
<minus/>
<cn cellml:units="uM_per_coulomb">5.182</cn>
</apply>
<ci>i_V</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>i_V</ci>
<apply>
<divide/>
<apply>
<times/>
<apply>
<divide/>
<apply>
<times/>
<ci>g_Ca</ci>
<ci>P</ci>
<cn cellml:units="dimensionless">2</cn>
<ci>F</ci>
<ci>V</ci>
</apply>
<apply>
<times/>
<ci>R_gas_const</ci>
<ci>Temp</ci>
</apply>
</apply>
<ci>Ca_ex</ci>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>F</ci>
<ci>V</ci>
</apply>
<apply>
<times/>
<ci>R_gas_const</ci>
<ci>Temp</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>Ca</ci>
<apply>
<plus/>
<apply>
<times/>
<ci>O</ci>
<ci>Ca_open</ci>
</apply>
<cn cellml:units="uM">0.1</cn>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="rate_constants">
<variable name="alpha" units="per_ms" public_interface="out"/>
<variable name="alpha_" units="per_ms" public_interface="out"/>
<variable name="beta" units="per_ms" public_interface="out"/>
<variable name="beta_" units="per_ms" public_interface="out"/>
<variable name="kG_plus" units="per_ms" public_interface="out"/>
<variable name="a" units="dimensionless" initial_value="0"/>
<variable name="ka_plus" units="per_mM_per_ms" initial_value="0.2"/>
<variable name="ka_minus" units="per_ms" initial_value="0.0015"/>
<variable name="V" units="mV" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<variable name="kG_minus" units="per_ms" initial_value="0.00025" public_interface="out"/>
<variable name="kG2_minus" units="per_ms" initial_value="0.016" public_interface="out"/>
<variable name="kG3_minus" units="per_ms" initial_value="1.024" public_interface="out"/>
<variable name="T" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>alpha</ci>
<apply>
<times/>
<cn cellml:units="per_ms">0.45</cn>
<apply>
<exp/>
<apply>
<divide/>
<ci>V</ci>
<cn cellml:units="mV">22</cn>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>alpha_</ci>
<apply>
<divide/>
<ci>alpha</ci>
<cn cellml:units="dimensionless">8</cn>
</apply>
</apply>
<apply>
<eq/>
<ci>beta</ci>
<apply>
<times/>
<cn cellml:units="per_ms">0.015</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<ci>V</ci>
</apply>
<cn cellml:units="mV">14</cn>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>beta_</ci>
<apply>
<times/>
<ci>beta</ci>
<cn cellml:units="dimensionless">8</cn>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>a</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>ka_plus</ci>
<ci>T</ci>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>a</ci>
</apply>
</apply>
<apply>
<times/>
<ci>ka_minus</ci>
<ci>a</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>kG_plus</ci>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="per_ms">3</cn>
<ci>a</ci>
</apply>
<apply>
<plus/>
<cn cellml:units="dimensionless">680</cn>
<apply>
<times/>
<cn cellml:units="dimensionless">320</cn>
<ci>a</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="C1">
<variable name="alpha" units="per_ms" public_interface="in"/>
<variable name="beta" units="per_ms" public_interface="in"/>
<variable name="C1" units="dimensionless" initial_value="1" public_interface="out"/>
<variable name="C2" units="dimensionless" public_interface="in"/>
<variable name="C_G1" units="dimensionless" public_interface="in"/>
<variable name="kG_minus" units="per_ms" public_interface="in"/>
<variable name="kG_plus" units="per_ms" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>C1</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<ci>beta</ci>
<ci>C2</ci>
</apply>
<apply>
<times/>
<ci>kG_minus</ci>
<ci>C_G1</ci>
</apply>
</apply>
<apply>
<times/>
<ci>C1</ci>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>alpha</ci>
</apply>
<ci>kG_plus</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="C2">
<variable name="alpha" units="per_ms" public_interface="in"/>
<variable name="beta" units="per_ms" public_interface="in"/>
<variable name="C1" units="dimensionless" public_interface="in"/>
<variable name="C2" units="dimensionless" initial_value="0" public_interface="out"/>
<variable name="C3" units="dimensionless" public_interface="in"/>
<variable name="C_G2" units="dimensionless" public_interface="in"/>
<variable name="kG_plus" units="per_ms" public_interface="in"/>
<variable name="kG2_minus" units="per_ms" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>C2</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>alpha</ci>
<ci>C1</ci>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>beta</ci>
<ci>C3</ci>
</apply>
<apply>
<times/>
<ci>kG2_minus</ci>
<ci>C_G2</ci>
</apply>
</apply>
<apply>
<times/>
<ci>C2</ci>
<apply>
<plus/>
<ci>beta</ci>
<apply>
<times/>
<cn cellml:units="dimensionless">3</cn>
<ci>alpha</ci>
</apply>
<ci>kG_plus</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="C3">
<variable name="alpha" units="per_ms" public_interface="in"/>
<variable name="beta" units="per_ms" public_interface="in"/>
<variable name="C2" units="dimensionless" public_interface="in"/>
<variable name="C3" units="dimensionless" initial_value="0" public_interface="out"/>
<variable name="C4" units="dimensionless" public_interface="in"/>
<variable name="C_G3" units="dimensionless" public_interface="in"/>
<variable name="kG_plus" units="per_ms" public_interface="in"/>
<variable name="kG3_minus" units="per_ms" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>C3</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">3</cn>
<ci>alpha</ci>
<ci>C2</ci>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">3</cn>
<ci>beta</ci>
<ci>C4</ci>
</apply>
<apply>
<times/>
<ci>kG3_minus</ci>
<ci>C_G3</ci>
</apply>
</apply>
<apply>
<times/>
<ci>C3</ci>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>beta</ci>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>alpha</ci>
</apply>
<ci>kG_plus</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="C4">
<variable name="alpha" units="per_ms" public_interface="in"/>
<variable name="beta" units="per_ms" public_interface="in"/>
<variable name="C3" units="dimensionless" public_interface="in"/>
<variable name="C4" units="dimensionless" initial_value="0" public_interface="out"/>
<variable name="O" units="dimensionless" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>C4</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>alpha</ci>
<ci>C3</ci>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>beta</ci>
<ci>O</ci>
</apply>
</apply>
<apply>
<times/>
<ci>C4</ci>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">3</cn>
<ci>beta</ci>
</apply>
<ci>alpha</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="O">
<variable name="C1" units="dimensionless" public_interface="in"/>
<variable name="C2" units="dimensionless" public_interface="in"/>
<variable name="C3" units="dimensionless" public_interface="in"/>
<variable name="C4" units="dimensionless" public_interface="in"/>
<variable name="O" units="dimensionless" public_interface="out"/>
<variable name="C_G1" units="dimensionless" public_interface="in"/>
<variable name="C_G2" units="dimensionless" public_interface="in"/>
<variable name="C_G3" units="dimensionless" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<variable name="C_G" units="dimensionless"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>O</ci>
<apply>
<minus/>
<apply>
<minus/>
<apply>
<minus/>
<apply>
<minus/>
<apply>
<minus/>
<apply>
<minus/>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>C1</ci>
</apply>
<ci>C2</ci>
</apply>
<ci>C3</ci>
</apply>
<ci>C4</ci>
</apply>
<ci>C_G1</ci>
</apply>
<ci>C_G2</ci>
</apply>
<ci>C_G3</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>C_G</ci>
<apply>
<plus/>
<ci>C_G1</ci>
<ci>C_G2</ci>
<ci>C_G3</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="C_G1">
<variable name="alpha_" units="per_ms" public_interface="in"/>
<variable name="beta_" units="per_ms" public_interface="in"/>
<variable name="C1" units="dimensionless" public_interface="in"/>
<variable name="C_G1" units="dimensionless" initial_value="0" public_interface="out"/>
<variable name="C_G2" units="dimensionless" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<variable name="kG_plus" units="per_ms" public_interface="in"/>
<variable name="kG_minus" units="per_ms" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>C_G1</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<ci>beta_</ci>
<ci>C_G2</ci>
</apply>
<apply>
<times/>
<ci>kG_plus</ci>
<ci>C1</ci>
</apply>
</apply>
<apply>
<times/>
<ci>C_G1</ci>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>alpha_</ci>
</apply>
<ci>kG_minus</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="C_G2">
<variable name="alpha_" units="per_ms" public_interface="in"/>
<variable name="beta_" units="per_ms" public_interface="in"/>
<variable name="C2" units="dimensionless" public_interface="in"/>
<variable name="C_G1" units="dimensionless" public_interface="in"/>
<variable name="C_G2" units="dimensionless" initial_value="0" public_interface="out"/>
<variable name="C_G3" units="dimensionless" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<variable name="kG_plus" units="per_ms" public_interface="in"/>
<variable name="kG2_minus" units="per_ms" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>C_G2</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">4</cn>
<ci>alpha_</ci>
<ci>C_G1</ci>
</apply>
<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>beta_</ci>
<ci>C_G3</ci>
</apply>
<apply>
<times/>
<ci>kG_plus</ci>
<ci>C2</ci>
</apply>
</apply>
<apply>
<times/>
<ci>C_G2</ci>
<apply>
<plus/>
<ci>beta_</ci>
<apply>
<times/>
<cn cellml:units="dimensionless">3</cn>
<ci>alpha_</ci>
</apply>
<ci>kG2_minus</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="C_G3">
<variable name="alpha_" units="per_ms" public_interface="in"/>
<variable name="beta_" units="per_ms" public_interface="in"/>
<variable name="C3" units="dimensionless" public_interface="in"/>
<variable name="C_G2" units="dimensionless" public_interface="in"/>
<variable name="C_G3" units="dimensionless" initial_value="0" public_interface="out"/>
<variable name="time" units="s" public_interface="in"/>
<variable name="kG_plus" units="per_ms" public_interface="in"/>
<variable name="kG3_minus" units="per_ms" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>C_G3</ci>
</apply>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">3</cn>
<ci>alpha_</ci>
<ci>C_G2</ci>
</apply>
<apply>
<times/>
<ci>kG_plus</ci>
<ci>C3</ci>
</apply>
</apply>
<apply>
<times/>
<ci>C_G3</ci>
<apply>
<plus/>
<apply>
<times/>
<cn cellml:units="dimensionless">2</cn>
<ci>beta_</ci>
</apply>
<ci>kG3_minus</ci>
</apply>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="membrane_post">
<variable name="V_post" units="mV" initial_value="-65" public_interface="out"/>
<variable name="Cm" units="uF_per_cm2" public_interface="in"/>
<variable name="i_syn" units="uA_per_cm2" public_interface="in"/>
<variable name="i_Na_post" units="uA_per_cm2" public_interface="in"/>
<variable name="i_K_post" units="uA_per_cm2" public_interface="in"/>
<variable name="i_leak_post" units="uA_per_cm2" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>V_post</ci>
</apply>
<apply>
<divide/>
<apply>
<minus/>
<apply>
<plus/>
<ci>i_Na_post</ci>
<ci>i_K_post</ci>
<ci>i_leak_post</ci>
<ci>i_syn</ci>
</apply>
</apply>
<ci>Cm</ci>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="synaptic_current">
<variable name="i_syn" units="uA_per_cm2" public_interface="out"/>
<variable name="g_syn" units="mS_per_cm2" initial_value="0.2"/>
<variable name="V_post" units="mV" public_interface="in"/>
<variable name="V_syn" units="mV" initial_value="0"/>
<variable name="b" units="dimensionless" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_syn</ci>
<apply>
<times/>
<ci>g_syn</ci>
<ci>b</ci>
<apply>
<minus/>
<ci>V_post</ci>
<ci>V_syn</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="leak_current_post">
<variable name="i_leak_post" units="uA_per_cm2" public_interface="out"/>
<variable name="time" units="s" public_interface="in"/>
<variable name="V_post" units="mV" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_leak_post</ci>
<apply>
<times/>
<cn cellml:units="pS_per_cm2">0.3</cn>
<apply>
<plus/>
<ci>V_post</ci>
<cn cellml:units="mV">54</cn>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="sodium_current_post">
<variable name="i_Na_post" units="uA_per_cm2" public_interface="out"/>
<variable name="x_infinity" units="dimensionless" public_interface="in"/>
<variable name="V_post" units="mV" public_interface="in"/>
<variable name="n_post" units="dimensionless" public_interface="in"/>
<variable name="alpha_x" units="dimensionless" public_interface="in"/>
<variable name="beta_x" units="dimensionless" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_Na_post</ci>
<apply>
<times/>
<cn cellml:units="pS_per_cm2">120</cn>
<apply>
<power/>
<ci>x_infinity</ci>
<cn cellml:units="dimensionless">3</cn>
</apply>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>n_post</ci>
</apply>
<apply>
<minus/>
<ci>V_post</ci>
<cn cellml:units="mV">120</cn>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="potassium_current_post">
<variable name="i_K_post" units="uA_per_cm2" public_interface="out"/>
<variable name="V_post" units="mV" public_interface="in"/>
<variable name="n_post" units="dimensionless" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_K_post</ci>
<apply>
<times/>
<cn cellml:units="pS_per_cm2">36</cn>
<apply>
<power/>
<ci>n_post</ci>
<cn cellml:units="dimensionless">4</cn>
</apply>
<apply>
<plus/>
<ci>V_post</ci>
<cn cellml:units="mV">77</cn>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="potassium_current_n_gate_post">
<variable name="n_post" units="dimensionless" initial_value="0" public_interface="out"/>
<variable name="alpha_n" units="per_ms" public_interface="in"/>
<variable name="beta_n" units="per_ms" public_interface="in"/>
<variable name="time" units="s" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>n_post</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>alpha_n</ci>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>n_post</ci>
</apply>
</apply>
<apply>
<times/>
<ci>beta_n</ci>
<ci>n_post</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component xmlns="http://www.cellml.org/cellml/1.0#" name="bound_postsynaptic_receptors">
<variable name="b" units="dimensionless" initial_value="0" public_interface="out"/>
<variable name="time" units="s" public_interface="in"/>
<variable name="k_b_plus" units="per_mM_per_ms" initial_value="2"/>
<variable name="k_b_minus" units="per_ms" initial_value="1"/>
<variable name="T" units="mM" public_interface="in"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>b</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>k_b_plus</ci>
<ci>T</ci>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>b</ci>
</apply>
</apply>
<apply>
<times/>
<ci>k_b_minus</ci>
<ci>b</ci>
</apply>
</apply>
</apply>
</math>
</component>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="membrane"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="sodium_current"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="potassium_current"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="potassium_current_n_gate"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="leak_current"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="transmitter_release"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="calcium_concentration"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="rate_constants"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="C1"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="C2"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="C3"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="C4"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="O"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="C_G1"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="C_G2"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="C_G3"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="membrane_post"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="potassium_current_n_gate_post"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="bound_postsynaptic_receptors"/>
<map_variables variable_1="time" variable_2="time"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="leak_current_post"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane" component_2="sodium_current"/>
<map_variables variable_1="V" variable_2="V"/>
<map_variables variable_1="i_Na" variable_2="i_Na"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane" component_2="potassium_current"/>
<map_variables variable_1="V" variable_2="V"/>
<map_variables variable_1="i_K" variable_2="i_K"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane" component_2="potassium_current_n_gate"/>
<map_variables variable_1="V" variable_2="V"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="potassium_current" component_2="potassium_current_n_gate"/>
<map_variables variable_1="n" variable_2="n"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="sodium_current" component_2="potassium_current_n_gate"/>
<map_variables variable_1="n" variable_2="n"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane" component_2="leak_current"/>
<map_variables variable_1="V" variable_2="V"/>
<map_variables variable_1="i_leak" variable_2="i_leak"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane" component_2="calcium_concentration"/>
<map_variables variable_1="V" variable_2="V"/>
<map_variables variable_1="R_gas_const" variable_2="R_gas_const"/>
<map_variables variable_1="F" variable_2="F"/>
<map_variables variable_1="Temp" variable_2="Temp"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane" component_2="rate_constants"/>
<map_variables variable_1="V" variable_2="V"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="O" component_2="calcium_concentration"/>
<map_variables variable_1="O" variable_2="O"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="transmitter_release" component_2="calcium_concentration"/>
<map_variables variable_1="Ca" variable_2="Ca"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="transmitter_release" component_2="rate_constants"/>
<map_variables variable_1="T" variable_2="T"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="rate_constants" component_2="C1"/>
<map_variables variable_1="alpha" variable_2="alpha"/>
<map_variables variable_1="beta" variable_2="beta"/>
<map_variables variable_1="kG_plus" variable_2="kG_plus"/>
<map_variables variable_1="kG_minus" variable_2="kG_minus"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C2" component_2="C1"/>
<map_variables variable_1="C2" variable_2="C2"/>
<map_variables variable_1="C1" variable_2="C1"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C_G1" component_2="C1"/>
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<map_variables variable_1="C1" variable_2="C1"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="rate_constants" component_2="C2"/>
<map_variables variable_1="alpha" variable_2="alpha"/>
<map_variables variable_1="beta" variable_2="beta"/>
<map_variables variable_1="kG_plus" variable_2="kG_plus"/>
<map_variables variable_1="kG2_minus" variable_2="kG2_minus"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C3" component_2="C2"/>
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<map_variables variable_1="C2" variable_2="C2"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C_G2" component_2="C2"/>
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<map_variables variable_1="C2" variable_2="C2"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="rate_constants" component_2="C3"/>
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<map_variables variable_1="beta" variable_2="beta"/>
<map_variables variable_1="kG_plus" variable_2="kG_plus"/>
<map_variables variable_1="kG3_minus" variable_2="kG3_minus"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C4" component_2="C3"/>
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<map_variables variable_1="C3" variable_2="C3"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C_G3" component_2="C3"/>
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<map_variables variable_1="C3" variable_2="C3"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="rate_constants" component_2="C4"/>
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<map_variables variable_1="beta" variable_2="beta"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="O" component_2="C4"/>
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<map_variables variable_1="C4" variable_2="C4"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C1" component_2="O"/>
<map_variables variable_1="C1" variable_2="C1"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C2" component_2="O"/>
<map_variables variable_1="C2" variable_2="C2"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C3" component_2="O"/>
<map_variables variable_1="C3" variable_2="C3"/>
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<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C_G1" component_2="O"/>
<map_variables variable_1="C_G1" variable_2="C_G1"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C_G2" component_2="O"/>
<map_variables variable_1="C_G2" variable_2="C_G2"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C_G3" component_2="O"/>
<map_variables variable_1="C_G3" variable_2="C_G3"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="rate_constants" component_2="C_G1"/>
<map_variables variable_1="alpha_" variable_2="alpha_"/>
<map_variables variable_1="beta_" variable_2="beta_"/>
<map_variables variable_1="kG_plus" variable_2="kG_plus"/>
<map_variables variable_1="kG_minus" variable_2="kG_minus"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C_G2" component_2="C_G1"/>
<map_variables variable_1="C_G2" variable_2="C_G2"/>
<map_variables variable_1="C_G1" variable_2="C_G1"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="rate_constants" component_2="C_G2"/>
<map_variables variable_1="alpha_" variable_2="alpha_"/>
<map_variables variable_1="beta_" variable_2="beta_"/>
<map_variables variable_1="kG_plus" variable_2="kG_plus"/>
<map_variables variable_1="kG2_minus" variable_2="kG2_minus"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="C_G3" component_2="C_G2"/>
<map_variables variable_1="C_G3" variable_2="C_G3"/>
<map_variables variable_1="C_G2" variable_2="C_G2"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="rate_constants" component_2="C_G3"/>
<map_variables variable_1="alpha_" variable_2="alpha_"/>
<map_variables variable_1="beta_" variable_2="beta_"/>
<map_variables variable_1="kG_plus" variable_2="kG_plus"/>
<map_variables variable_1="kG3_minus" variable_2="kG3_minus"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="environment" component_2="stimulus_protocol"/>
<map_variables variable_1="time" variable_2="time"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane" component_2="stimulus_protocol"/>
<map_variables variable_1="i_app" variable_2="Istim"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane" component_2="membrane_post"/>
<map_variables variable_1="Cm" variable_2="Cm"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane_post" component_2="synaptic_current"/>
<map_variables variable_1="i_syn" variable_2="i_syn"/>
<map_variables variable_1="V_post" variable_2="V_post"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane_post" component_2="sodium_current_post"/>
<map_variables variable_1="V_post" variable_2="V_post"/>
<map_variables variable_1="i_Na_post" variable_2="i_Na_post"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane_post" component_2="potassium_current_post"/>
<map_variables variable_1="i_K_post" variable_2="i_K_post"/>
<map_variables variable_1="V_post" variable_2="V_post"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="membrane_post" component_2="leak_current_post"/>
<map_variables variable_1="i_leak_post" variable_2="i_leak_post"/>
<map_variables variable_1="V_post" variable_2="V_post"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="sodium_current_post" component_2="sodium_current"/>
<map_variables variable_1="x_infinity" variable_2="x_infinity"/>
<map_variables variable_1="alpha_x" variable_2="alpha_x"/>
<map_variables variable_1="beta_x" variable_2="beta_x"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="potassium_current_n_gate" component_2="potassium_current_n_gate_post"/>
<map_variables variable_1="alpha_n" variable_2="alpha_n"/>
<map_variables variable_1="beta_n" variable_2="beta_n"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="potassium_current_n_gate_post" component_2="sodium_current_post"/>
<map_variables variable_1="n_post" variable_2="n_post"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="potassium_current_n_gate_post" component_2="potassium_current_post"/>
<map_variables variable_1="n_post" variable_2="n_post"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="bound_postsynaptic_receptors" component_2="synaptic_current"/>
<map_variables variable_1="b" variable_2="b"/>
</connection>
<connection xmlns="http://www.cellml.org/cellml/1.0#">
<map_components component_1="bound_postsynaptic_receptors" component_2="transmitter_release"/>
<map_variables variable_1="T" variable_2="T"/>
</connection>
</model>
