- Author:
- pmr2.import <nobody@models.cellml.org>
- Date:
- 2008-07-17 02:18:21+12:00
- Desc:
- committing version02 of purvis_butera_2005
- Permanent Source URI:
- http://models.cellml.org/workspace/purvis_butera_2005/rawfile/f33430047c6f6c2f195381b678b87499bd2f3475/purvis_butera_2005.cellml
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This CellML file was generated on 17/07/2008 at 2:14:10 at p.m. using:
COR (0.9.31.979)
Copyright 2002-2008 Dr Alan Garny
http://COR.physiol.ox.ac.uk/ - COR@physiol.ox.ac.uk
CellML 1.0 was used to generate this model
http://www.CellML.org/
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<title>Ionic Current Model of Hypoglossal Motoneuron</title>
<author>
<firstname>Geoffrey</firstname>
<surname>Nunns</surname>
<affiliation>
<shortaffil>Bioengineering Institute, University of Auckland</shortaffil>
</affiliation>
</author>
</articleinfo>
<section id="sec_status">
<title>Model Status</title>
<para>
This CellML model is known to run in both PCEnv and COR and the units have been checked and are consistent. However the CellML model is curently unable to perfectly recreate the published results (we are several orders of magnitude out with certain variables). Further curation is required and we are endevouring to seek the help of the original model authors.
</para>
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<sect1 id="sec_structure">
<title>Model Structure</title>
<para>
Abstract: The hypoglossal motoneuron (HM) innervates the tongue and along with its roles in mastication and swallowing, the tongue plays an important role in breathing (Bartlett <emphasis>et al.</emphasis> 1990; Lowe 1980; reviewed in Sawczuk and Mosier 2001). The muscle of the tongue contributes to effective breathing by maintaining a patent upper airway and problems with the control of HMs can cause the breathing disorder obstructive sleep apnea (Fogel <emphasis>et al.</emphasis> 2000; Remmers <emphasis>et al.</emphasis> 1978). For this reason, extensive experimental work has been done on these motoneurons. In this communication, Butera and Smith present a computational model of a HM that provides a framework for analyzing the role of ionic currents in these neurons.
</para>
<para>
In this paper, Liston Purvis and Robert Butera have developed a single-compartment, electrophysiological, hypoglossal motoneuron (HM) model based primarily on experimental data from neonatal rat HMs. The model is able to reproduce the fine features of the HM action potential: the fast afterhyperpolarization, the afterdepolarization, and the medium-duration afterhyperpolarization (mAHP). The model also reproduces the repetitive firing properties seen in neonatal HMs and replicates the neuron's response to pharmacological experiments. The model was used to study the role of specific ionic currents in HM firing and how variations in the densities of these currents may account for age-dependent changes in excitability seen in HMs. By varying the density of a fast inactivating calcium current, the model alternates between accelerating and adapting firing patterns. Modeling the age-dependent increase in H current density accounts for the decrease in mAHP duration observed experimentally, but does not fully account for the decrease in input resistance. An increase in the density of the voltage-dependent potassium currents and the H current is required to account for the decrease in input resistance. These changes also account for the age-dependent decrease in action potential duration.</para>
<para>
The complete original paper reference is cited below:
</para>
<para>
<ulink url="http://jn.physiology.org/cgi/content/abstract/93/2/723">Ionic Current Model of a Hypoglossal Motoneuron</ulink>, Robert J. Butera, Jr., Liston K. Purvis, 2005, <ulink url="http://jn.physiology.org/">
<emphasis>Journal of Neurophysiology</emphasis>
</ulink>, 93, 723-733. (<ulink url="http://intl-jn.physiology.org/cgi/content/full/93/2/723">Full text</ulink> and <ulink url="http://jn.physiology.org/cgi/reprint/93/2/723">PDF</ulink> versions of the article are available for Journal Members on the JN website.) <ulink url="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&list_uids=15653786&dopt=Abstract">PubMed ID: 15653786</ulink>
</para>
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<eq/>
<ci>tau_h</ci>
<apply>
<plus/>
<apply>
<divide/>
<cn cellml:units="millisecond">20</cn>
<apply>
<plus/>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_1</ci>
</apply>
<ci>sigma_1</ci>
</apply>
</apply>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_1</ci>
</apply>
</apply>
<ci>sigma_1</ci>
</apply>
</apply>
</apply>
</apply>
<cn cellml:units="millisecond">1</cn>
</apply>
</apply>
<apply>
<eq/>
<ci>h_infinity</ci>
<apply>
<divide/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_h</ci>
</apply>
<ci>sigma_h</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>h</ci>
</apply>
<apply>
<divide/>
<apply>
<minus/>
<ci>h_infinity</ci>
<ci>h</ci>
</apply>
<ci>tau_h</ci>
</apply>
</apply>
</math>
</component>
<component name="N_HVA_calcium_current">
<variable units="nanoA" public_interface="out" name="i_N"/>
<variable units="uS" name="g_N" initial_value="0.05"/>
<variable units="millivolt" public_interface="in" name="E_Ca"/>
<variable units="millivolt" public_interface="in" private_interface="out" name="V"/>
<variable units="millisecond" public_interface="in" private_interface="out" name="time"/>
<variable units="dimensionless" private_interface="in" name="m"/>
<variable units="dimensionless" private_interface="in" name="h"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_N</ci>
<apply>
<times/>
<ci>g_N</ci>
<ci>m</ci>
<ci>h</ci>
<apply>
<minus/>
<ci>V</ci>
<ci>E_Ca</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="N_HVA_calcium_current_m_gate">
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="millisecond" public_interface="in" name="time"/>
<variable units="dimensionless" public_interface="out" name="m" initial_value="0.001"/>
<variable units="millivolt" name="theta_m" initial_value="30"/>
<variable units="millivolt" name="sigma_m" initial_value="6"/>
<variable units="millisecond" name="tau_m" initial_value="5"/>
<variable units="dimensionless" name="m_infinity"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>m_infinity</ci>
<apply>
<divide/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_m</ci>
</apply>
</apply>
<ci>sigma_m</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>m</ci>
</apply>
<apply>
<divide/>
<apply>
<minus/>
<ci>m_infinity</ci>
<ci>m</ci>
</apply>
<ci>tau_m</ci>
</apply>
</apply>
</math>
</component>
<component name="N_HVA_calcium_current_h_gate">
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="millisecond" public_interface="in" name="time"/>
<variable units="dimensionless" public_interface="out" name="h" initial_value="0.649"/>
<variable units="millivolt" name="theta_h" initial_value="70"/>
<variable units="millivolt" name="sigma_h" initial_value="3"/>
<variable units="millisecond" name="tau_h" initial_value="25"/>
<variable units="dimensionless" name="h_infinity"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>h_infinity</ci>
<apply>
<divide/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_h</ci>
</apply>
<ci>sigma_h</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>h</ci>
</apply>
<apply>
<divide/>
<apply>
<minus/>
<ci>h_infinity</ci>
<ci>h</ci>
</apply>
<ci>tau_h</ci>
</apply>
</apply>
</math>
</component>
<component name="P_HVA_calcium_current">
<variable units="nanoA" public_interface="out" name="i_P"/>
<variable units="uS" name="g_P" initial_value="0.05"/>
<variable units="millivolt" public_interface="in" name="E_Ca"/>
<variable units="millivolt" public_interface="in" private_interface="out" name="V"/>
<variable units="millisecond" public_interface="in" private_interface="out" name="time"/>
<variable units="dimensionless" private_interface="in" name="m"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_P</ci>
<apply>
<times/>
<ci>g_P</ci>
<ci>m</ci>
<apply>
<minus/>
<ci>V</ci>
<ci>E_Ca</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="P_HVA_calcium_current_m_gate">
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="millisecond" public_interface="in" name="time"/>
<variable units="dimensionless" public_interface="out" name="m" initial_value="0"/>
<variable units="millivolt" name="theta_m" initial_value="17"/>
<variable units="millivolt" name="sigma_m" initial_value="3"/>
<variable units="millisecond" name="tau_m" initial_value="10"/>
<variable units="dimensionless" name="m_infinity"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>m_infinity</ci>
<apply>
<divide/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_m</ci>
</apply>
</apply>
<ci>sigma_m</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>m</ci>
</apply>
<apply>
<divide/>
<apply>
<minus/>
<ci>m_infinity</ci>
<ci>m</ci>
</apply>
<ci>tau_m</ci>
</apply>
</apply>
</math>
</component>
<component name="calcium_dependent_potassium_current">
<variable units="nanoA" public_interface="out" name="i_SK"/>
<variable units="uS" name="g_SK" initial_value="0.3"/>
<variable units="millivolt" public_interface="in" name="E_K"/>
<variable units="millivolt" public_interface="in" private_interface="out" name="V"/>
<variable units="millisecond" public_interface="in" private_interface="out" name="time"/>
<variable units="nanoA" public_interface="in" private_interface="out" name="i_T"/>
<variable units="nanoA" public_interface="in" private_interface="out" name="i_N"/>
<variable units="nanoA" public_interface="in" private_interface="out" name="i_P"/>
<variable units="dimensionless" private_interface="in" name="z"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_SK</ci>
<apply>
<times/>
<ci>g_SK</ci>
<apply>
<power/>
<ci>z</ci>
<cn cellml:units="dimensionless">2</cn>
</apply>
<apply>
<minus/>
<ci>V</ci>
<ci>E_K</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="calcium_dependent_potassium_current_z_gate">
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="millisecond" public_interface="in" name="time"/>
<variable units="nanoA" public_interface="in" name="i_T"/>
<variable units="nanoA" public_interface="in" name="i_N"/>
<variable units="nanoA" public_interface="in" name="i_P"/>
<variable units="dimensionless" public_interface="out" name="z" initial_value="0"/>
<variable units="uM_per_nanocoulomb" name="K1" initial_value="-500"/>
<variable units="per_ms" name="K2" initial_value="0.04"/>
<variable units="uM" name="Ca_conc" initial_value="0.0604"/>
<variable units="millisecond" name="tau_z" initial_value="1"/>
<variable units="dimensionless" name="z_infinity"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>Ca_conc</ci>
</apply>
<apply>
<minus/>
<apply>
<times/>
<apply>
<divide/>
<cn cellml:units="second">1</cn>
<cn cellml:units="millisecond">1000</cn>
</apply>
<ci>K1</ci>
<apply>
<plus/>
<ci>i_T</ci>
<ci>i_N</ci>
<ci>i_P</ci>
</apply>
</apply>
<apply>
<times/>
<ci>K2</ci>
<ci>Ca_conc</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>z_infinity</ci>
<apply>
<divide/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<power/>
<apply>
<divide/>
<cn cellml:units="uM">0.003</cn>
<ci>Ca_conc</ci>
</apply>
<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>z</ci>
</apply>
<apply>
<divide/>
<apply>
<minus/>
<ci>z_infinity</ci>
<ci>z</ci>
</apply>
<ci>tau_z</ci>
</apply>
</apply>
</math>
</component>
<component name="fast_transient_potassium_current">
<variable units="nanoA" public_interface="out" name="i_A"/>
<variable units="uS" name="g_A" initial_value="1"/>
<variable units="millivolt" public_interface="in" name="E_K"/>
<variable units="millivolt" public_interface="in" private_interface="out" name="V"/>
<variable units="millisecond" public_interface="in" private_interface="out" name="time"/>
<variable units="dimensionless" private_interface="in" name="m"/>
<variable units="dimensionless" private_interface="in" name="h"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_A</ci>
<apply>
<times/>
<ci>g_A</ci>
<ci>m</ci>
<ci>h</ci>
<apply>
<minus/>
<ci>V</ci>
<ci>E_K</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="fast_transient_potassium_current_m_gate">
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="millisecond" public_interface="in" name="time"/>
<variable units="dimensionless" public_interface="out" name="m" initial_value="0.057"/>
<variable units="millivolt" name="theta_m" initial_value="27"/>
<variable units="millivolt" name="sigma_m" initial_value="16"/>
<variable units="millivolt" name="theta_1" initial_value="40"/>
<variable units="millivolt" name="theta_2" initial_value="74"/>
<variable units="millivolt" name="sigma_1" initial_value="5"/>
<variable units="millivolt" name="sigma_2" initial_value="7.5"/>
<variable units="millisecond" name="tau_m"/>
<variable units="dimensionless" name="m_infinity"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>tau_m</ci>
<apply>
<plus/>
<apply>
<divide/>
<cn cellml:units="millisecond">1</cn>
<apply>
<plus/>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_1</ci>
</apply>
<ci>sigma_1</ci>
</apply>
</apply>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_2</ci>
</apply>
</apply>
<ci>sigma_2</ci>
</apply>
</apply>
</apply>
</apply>
<cn cellml:units="millisecond">0.37</cn>
</apply>
</apply>
<apply>
<eq/>
<ci>m_infinity</ci>
<apply>
<divide/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_m</ci>
</apply>
</apply>
<ci>sigma_m</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>m</ci>
</apply>
<apply>
<divide/>
<apply>
<minus/>
<ci>m_infinity</ci>
<ci>m</ci>
</apply>
<ci>tau_m</ci>
</apply>
</apply>
</math>
</component>
<component name="fast_transient_potassium_current_h_gate">
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="millisecond" public_interface="in" name="time"/>
<variable units="dimensionless" public_interface="out" name="h" initial_value="0.287"/>
<variable units="millivolt" name="theta_h" initial_value="80"/>
<variable units="millivolt" name="sigma_h" initial_value="11"/>
<variable units="millisecond" name="tau_h" initial_value="20"/>
<variable units="dimensionless" name="h_infinity"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>h_infinity</ci>
<apply>
<divide/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_h</ci>
</apply>
<ci>sigma_h</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>h</ci>
</apply>
<apply>
<divide/>
<apply>
<minus/>
<ci>h_infinity</ci>
<ci>h</ci>
</apply>
<ci>tau_h</ci>
</apply>
</apply>
</math>
</component>
<component name="hyperpolarization_activated_current">
<variable units="nanoA" public_interface="out" name="i_H"/>
<variable units="uS" name="g_H" initial_value="0.005"/>
<variable units="millivolt" name="E_H" initial_value="-38.8"/>
<variable units="millivolt" public_interface="in" private_interface="out" name="V"/>
<variable units="millisecond" public_interface="in" private_interface="out" name="time"/>
<variable units="dimensionless" private_interface="in" name="m"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_H</ci>
<apply>
<times/>
<ci>g_H</ci>
<ci>m</ci>
<apply>
<minus/>
<ci>V</ci>
<ci>E_H</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="hyperpolarization_activated_current_m_gate">
<variable units="millivolt" public_interface="in" name="V"/>
<variable units="millisecond" public_interface="in" name="time"/>
<variable units="dimensionless" public_interface="out" name="m" initial_value="0.182"/>
<variable units="millivolt" name="theta_m" initial_value="79.8"/>
<variable units="millivolt" name="sigma_m" initial_value="5.3"/>
<variable units="millivolt" name="theta_1" initial_value="70"/>
<variable units="millivolt" name="sigma_1" initial_value="11"/>
<variable units="millisecond" name="tau_m"/>
<variable units="dimensionless" name="m_infinity"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>tau_m</ci>
<apply>
<plus/>
<apply>
<divide/>
<cn cellml:units="millisecond">1</cn>
<apply>
<plus/>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_1</ci>
</apply>
<ci>sigma_1</ci>
</apply>
</apply>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<minus/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_1</ci>
</apply>
</apply>
<ci>sigma_1</ci>
</apply>
</apply>
</apply>
</apply>
<cn cellml:units="millisecond">50</cn>
</apply>
</apply>
<apply>
<eq/>
<ci>m_infinity</ci>
<apply>
<divide/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<plus/>
<cn cellml:units="dimensionless">1</cn>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<plus/>
<ci>V</ci>
<ci>theta_m</ci>
</apply>
<ci>sigma_m</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>m</ci>
</apply>
<apply>
<divide/>
<apply>
<minus/>
<ci>m_infinity</ci>
<ci>m</ci>
</apply>
<ci>tau_m</ci>
</apply>
</apply>
</math>
</component>
<component name="stimulus_protocol">
<variable units="nanoA" public_interface="out" name="i_app"/>
<variable units="millisecond" name="i_stimStart" initial_value="0.5"/>
<variable units="millisecond" name="i_stimEnd" initial_value="0.9"/>
<variable units="nanoA" name="i_stimAmplitude" initial_value="1"/>
<variable units="millisecond" public_interface="in" name="time"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>i_app</ci>
<piecewise>
<piece>
<ci>i_stimAmplitude</ci>
<apply>
<and/>
<apply>
<geq/>
<ci>time</ci>
<ci>i_stimStart</ci>
</apply>
<apply>
<leq/>
<ci>time</ci>
<ci>i_stimEnd</ci>
</apply>
</apply>
</piece>
<otherwise>
<cn cellml:units="nanoA">0</cn>
</otherwise>
</piecewise>
</apply>
</math>
</component>
<group>
<relationship_ref relationship="encapsulation"/>
<component_ref component="sodium_current">
<component_ref component="sodium_current_h_gate"/>
<component_ref component="sodium_current_m_gate"/>
</component_ref>
<component_ref component="persistent_sodium_current">
<component_ref component="persistent_sodium_current_h_gate"/>
<component_ref component="persistent_sodium_current_m_gate"/>
</component_ref>
<component_ref component="delayed_rectifier_current">
<component_ref component="delayed_rectifier_current_n_gate"/>
</component_ref>
<component_ref component="LVA_calcium_current">
<component_ref component="LVA_calcium_current_h_gate"/>
<component_ref component="LVA_calcium_current_m_gate"/>
</component_ref>
<component_ref component="N_HVA_calcium_current">
<component_ref component="N_HVA_calcium_current_h_gate"/>
<component_ref component="N_HVA_calcium_current_m_gate"/>
</component_ref>
<component_ref component="P_HVA_calcium_current">
<component_ref component="P_HVA_calcium_current_m_gate"/>
</component_ref>
<component_ref component="calcium_dependent_potassium_current">
<component_ref component="calcium_dependent_potassium_current_z_gate"/>
</component_ref>
<component_ref component="fast_transient_potassium_current">
<component_ref component="fast_transient_potassium_current_h_gate"/>
<component_ref component="fast_transient_potassium_current_m_gate"/>
</component_ref>
<component_ref component="hyperpolarization_activated_current">
<component_ref component="hyperpolarization_activated_current_m_gate"/>
</component_ref>
</group>
<connection>
<map_components component_2="membrane" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="sodium_current" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="persistent_sodium_current" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="delayed_rectifier_current" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="leak_current" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="LVA_calcium_current" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="N_HVA_calcium_current" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="P_HVA_calcium_current" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="calcium_dependent_potassium_current" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="fast_transient_potassium_current" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="hyperpolarization_activated_current" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="stimulus_protocol" component_1="environment"/>
<map_variables variable_2="time" variable_1="time"/>
</connection>
<connection>
<map_components component_2="stimulus_protocol" component_1="membrane"/>
<map_variables variable_2="i_app" variable_1="i_app"/>
</connection>
<connection>
<map_components component_2="sodium_current" component_1="membrane"/>
<map_variables variable_2="V" variable_1="V"/>
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