- Author:
- Dewan Sarwar <dsar941@UoA.auckland.ac.nz>
- Date:
- 2020-02-04 11:59:04+13:00
- Desc:
- replaced https with https
- Permanent Source URI:
- https://staging.physiomeproject.org/workspace/267/rawfile/86daa2d11994f6ee68559ac81fbea58f42bc3ced/SEDML/mackenzie_1996_1A.cellml
<?xml version='1.0' encoding='UTF-8'?>
<model cmeta:id="mackenzie_1996" name="mackenzie_1996" xmlns="http://www.cellml.org/cellml/1.1#" xmlns:cellml="http://www.cellml.org/cellml/1.1#" xmlns:cmeta="http://www.cellml.org/metadata/1.0#">
<units name="per_second">
<unit exponent="-1.0" units="second"/>
</units>
<units name="per_M2_per_second5">
<unit exponent="-2.0" units="M"/>
<unit exponent="-5.0" units="second"/>
</units>
<units name="M3">
<unit exponent="3.0" units="M"/>
</units>
<units name="per_M2_per_second">
<unit exponent="-2.0" units="M"/>
<unit exponent="-1.0" units="second"/>
</units>
<units name="uA_per_um2">
<unit exponent="-1.0" units="um2"/>
<unit units="uA"/>
</units>
<units name="pA_per_cm2">
<unit prefix="pico" units="ampere"/>
<unit exponent="-2.0" prefix="centi" units="metre"/>
</units>
<units name="M2">
<unit exponent="2.0" units="M"/>
</units>
<units name="umol">
<unit prefix="micro" units="mole"/>
</units>
<units name="per_second5">
<unit exponent="-5.0" units="second"/>
</units>
<units name="mol_per_um2">
<unit exponent="-1.0" units="um2"/>
<unit units="mole"/>
</units>
<units name="mV">
<unit prefix="milli" units="volt"/>
</units>
<units name="per_mol">
<unit exponent="-1.0" units="mole"/>
</units>
<units name="volt_per_second">
<unit units="volt"/>
<unit exponent="-1.0" units="second"/>
</units>
<units name="umol_per_second">
<unit units="umol"/>
<unit exponent="-1.0" units="second"/>
</units>
<units name="J_per_K_per_mol">
<unit units="joule"/>
<unit exponent="-1.0" units="mole"/>
<unit exponent="-1.0" units="kelvin"/>
</units>
<units name="M2_per_second">
<unit exponent="-1.0" units="second"/>
<unit exponent="2.0" units="M"/>
</units>
<units name="M">
<unit exponent="-1.0" units="litre"/>
<unit units="mole"/>
</units>
<units name="uA">
<unit prefix="micro" units="ampere"/>
</units>
<units name="uA_per_A">
<unit exponent="-1.0" units="ampere"/>
<unit units="uA"/>
</units>
<units name="um2">
<unit exponent="2.0" prefix="micro" units="metre"/>
</units>
<units name="C_per_mol">
<unit exponent="-1.0" units="mole"/>
<unit units="coulomb"/>
</units>
<units name="per_M_per_second">
<unit exponent="-1.0" units="M"/>
<unit exponent="-1.0" units="second"/>
</units>
<units name="umol_per_mol">
<unit units="umol"/>
<unit exponent="-1.0" units="mole"/>
</units>
<units name="mM">
<unit prefix="milli" units="mole"/>
<unit exponent="-1.0" units="litre"/>
</units>
<units name="M_per_second">
<unit exponent="-1.0" units="second"/>
<unit units="M"/>
</units>
<component name="environment">
<variable cmeta:id="environment.time" name="time" public_interface="out" units="second"/>
</component>
<component name="parameters">
<variable cmeta:id="parameters.k0_12" initial_value="20000" name="k0_12" public_interface="out" units="per_M_per_second"/>
<variable cmeta:id="parameters.k0_21" initial_value="400" name="k0_21" public_interface="out" units="per_second"/>
<variable cmeta:id="parameters.k0_23" initial_value="1e4" name="k0_23" public_interface="out" units="per_M_per_second"/>
<variable cmeta:id="parameters.k0_32" initial_value="20" name="k0_32" public_interface="out" units="per_second"/>
<variable cmeta:id="parameters.k0_34" initial_value="50" name="k0_34" public_interface="out" units="per_second"/>
<variable cmeta:id="parameters.k0_43" initial_value="50" name="k0_43" public_interface="out" units="per_second"/>
<variable cmeta:id="parameters.k0_45" initial_value="800" name="k0_45" public_interface="out" units="per_second"/>
<variable cmeta:id="parameters.k0_54" initial_value="6.7e6" name="k0_54" public_interface="out" units="per_M_per_second"/>
<variable cmeta:id="parameters.k0_25" initial_value="0.01" name="k0_25" public_interface="out" units="per_second"/>
<variable cmeta:id="parameters.k0_52" initial_value="0.17" name="k0_52" public_interface="out" units="per_second"/>
<variable cmeta:id="parameters.k0_56" initial_value="48" name="k0_56" public_interface="out" units="per_second"/>
<variable cmeta:id="parameters.k0_65" initial_value="50" name="k0_65" public_interface="out" units="per_M_per_second"/>
<variable cmeta:id="parameters.k0_61" initial_value="35" name="k0_61" public_interface="out" units="per_second"/>
<variable cmeta:id="parameters.k0_16" initial_value="100" name="k0_16" public_interface="out" units="per_second"/>
<variable cmeta:id="parameters.delta" initial_value="0.7" name="delta" public_interface="out" units="dimensionless"/>
<variable cmeta:id="parameters.alpha_p" name="alpha_p" public_interface="out" units="dimensionless"/>
<variable cmeta:id="parameters.alpha_pp" initial_value="0" name="alpha_pp" public_interface="out" units="dimensionless"/>
<variable cmeta:id="parameters.N_C" initial_value="8.4e10" name="N_C" units="dimensionless"/>
<variable cmeta:id="parameters.N_Avo" initial_value="6.022e23" name="N_Avo" units="per_mol"/>
<variable cmeta:id="parameters.area" initial_value="1e6" name="area" units="um2"/>
<variable cmeta:id="parameters.C_T" name="C_T" public_interface="out" units="umol"/>
<variable cmeta:id="parameters.n" initial_value="1" name="n" public_interface="out" units="dimensionless"/>
<variable cmeta:id="parameters.z_c" initial_value="-1" name="z_c" public_interface="out" units="dimensionless"/>
<variable cmeta:id="parameters.z_Na" initial_value="1" name="z_Na" public_interface="out" units="dimensionless"/>
<variable cmeta:id="parameters.F" initial_value="96485.34" name="F" public_interface="out" units="C_per_mol"/>
<variable cmeta:id="parameters.R" initial_value="8.314" name="R" public_interface="out" units="J_per_K_per_mol"/>
<variable cmeta:id="parameters.T" initial_value="310" name="T" public_interface="out" units="kelvin"/>
<variable cmeta:id="parameters.V" name="V" public_interface="in" units="volt"/>
<variable cmeta:id="parameters.mu" name="mu" public_interface="out" units="dimensionless"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>alpha_p</ci>
<apply>
<minus/>
<apply>
<minus/>
<cn cellml:units="dimensionless">1</cn>
<ci>delta</ci>
</apply>
<ci>alpha_pp</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>C_T</ci>
<apply>
<divide/>
<apply>
<times/>
<cn cellml:units="umol_per_mol" type="e-notation">1<sep/>6</cn>
<ci>N_C</ci>
</apply>
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</apply>
</apply>
<apply>
<eq/>
<ci>mu</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>F</ci>
<ci>V</ci>
</apply>
<apply>
<times/>
<ci>R</ci>
<ci>T</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="ion_concentrations">
<variable cmeta:id="ion_concentrations.V" initial_value="-150e-3" name="V" public_interface="out" units="volt"/>
<variable cmeta:id="ion_concentrations.Na_i" initial_value="10e-3" name="Na_i" public_interface="out" units="M"/>
<variable cmeta:id="ion_concentrations.Na_o" initial_value="100e-3" name="Na_o" public_interface="out" units="M"/>
<variable cmeta:id="ion_concentrations.glucose_i" initial_value="0e-3" name="glucose_i" public_interface="out" units="M"/>
<variable cmeta:id="ion_concentrations.glucose_o" initial_value="1e-3" name="glucose_o" public_interface="out" units="M"/>
<variable cmeta:id="ion_concentrations.time" name="time" public_interface="in" units="second"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
<bvar>
<ci>time</ci>
</bvar>
<ci>V</ci>
</apply>
<cn cellml:units="volt_per_second" type="e-notation">200<sep/>-6</cn>
</apply>
</math>
</component>
<component name="rate_constants">
<variable cmeta:id="rate_constants.Na_i" name="Na_i" public_interface="in" units="M"/>
<variable cmeta:id="rate_constants.Na_o" name="Na_o" public_interface="in" units="M"/>
<variable cmeta:id="rate_constants.glucose_i" name="glucose_i" public_interface="in" units="M"/>
<variable cmeta:id="rate_constants.glucose_o" name="glucose_o" public_interface="in" units="M"/>
<variable cmeta:id="rate_constants.k_12" name="k_12" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_21" name="k_21" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_23" name="k_23" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_32" name="k_32" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_34" name="k_34" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_43" name="k_43" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_45" name="k_45" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_54" name="k_54" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_25" name="k_25" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_52" name="k_52" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_56" name="k_56" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_65" name="k_65" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_61" name="k_61" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k_16" name="k_16" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k0_12" name="k0_12" public_interface="in" units="per_M_per_second"/>
<variable cmeta:id="rate_constants.ks_12" name="ks_12" public_interface="out" units="per_M_per_second"/>
<variable cmeta:id="rate_constants.k0_21" name="k0_21" public_interface="in" units="per_second"/>
<variable cmeta:id="rate_constants.k0_23" name="k0_23" public_interface="in" units="per_M_per_second"/>
<variable cmeta:id="rate_constants.k0_32" name="k0_32" public_interface="in" units="per_second"/>
<variable cmeta:id="rate_constants.k0_34" name="k0_34" public_interface="in" units="per_second"/>
<variable cmeta:id="rate_constants.k0_43" name="k0_43" public_interface="in" units="per_second"/>
<variable cmeta:id="rate_constants.k0_45" name="k0_45" public_interface="in" units="per_second"/>
<variable cmeta:id="rate_constants.k0_54_temp" name="k0_54_temp" public_interface="out" units="per_M_per_second"/>
<variable cmeta:id="rate_constants.k0_54" name="k0_54" public_interface="in" units="per_M_per_second"/>
<variable cmeta:id="rate_constants.k0_25" name="k0_25" public_interface="in" units="per_second"/>
<variable cmeta:id="rate_constants.k_52_temp" name="k_52_temp" public_interface="out" units="per_second"/>
<variable cmeta:id="rate_constants.k0_52" name="k0_52" public_interface="in" units="per_second"/>
<variable cmeta:id="rate_constants.k0_56" name="k0_56" public_interface="in" units="per_second"/>
<variable cmeta:id="rate_constants.k0_65" name="k0_65" public_interface="in" units="per_M_per_second"/>
<variable cmeta:id="rate_constants.k0_61" name="k0_61" public_interface="in" units="per_second"/>
<variable cmeta:id="rate_constants.k0_16" name="k0_16" public_interface="in" units="per_second"/>
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<variable cmeta:id="rate_constants.n" name="n" public_interface="in" units="dimensionless"/>
<variable cmeta:id="rate_constants.z_c" name="z_c" public_interface="in" units="dimensionless"/>
<variable cmeta:id="rate_constants.z_Na" name="z_Na" public_interface="in" units="dimensionless"/>
<variable cmeta:id="rate_constants.alpha_p" name="alpha_p" public_interface="in" units="dimensionless"/>
<variable cmeta:id="rate_constants.alpha_pp" name="alpha_pp" public_interface="in" units="dimensionless"/>
<variable cmeta:id="rate_constants.C_T" name="C_T" public_interface="in" units="umol"/>
<variable cmeta:id="rate_constants.mu" name="mu" public_interface="in" units="dimensionless"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>ks_12</ci>
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<times/>
<ci>k0_12</ci>
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<exp/>
<apply>
<divide/>
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<times/>
<apply>
<minus/>
<ci>n</ci>
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<ci>mu</ci>
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<cn cellml:units="dimensionless">2</cn>
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</apply>
<apply>
<eq/>
<ci>k_12</ci>
<apply>
<times/>
<ci>ks_12</ci>
<apply>
<power/>
<ci>Na_o</ci>
<ci>n</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>k_21</ci>
<apply>
<times/>
<ci>k0_21</ci>
<apply>
<exp/>
<apply>
<divide/>
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<times/>
<ci>n</ci>
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<ci>alpha_p</ci>
<ci>mu</ci>
</apply>
<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>k_23</ci>
<apply>
<times/>
<ci>k0_23</ci>
<ci>glucose_o</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>k_32</ci>
<ci>k0_32</ci>
</apply>
<apply>
<eq/>
<ci>k_34</ci>
<apply>
<times/>
<ci>k0_34</ci>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<minus/>
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<plus/>
<ci>z_c</ci>
<ci>n</ci>
</apply>
</apply>
<ci>delta</ci>
<ci>mu</ci>
</apply>
<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>k_43</ci>
<apply>
<times/>
<ci>k0_43</ci>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<plus/>
<ci>z_c</ci>
<ci>n</ci>
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<ci>delta</ci>
<ci>mu</ci>
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<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>k_45</ci>
<ci>k0_45</ci>
</apply>
<apply>
<eq/>
<ci>k_54</ci>
<apply>
<times/>
<ci>k0_54_temp</ci>
<ci>glucose_i</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>k_25</ci>
<apply>
<times/>
<ci>k0_25</ci>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<minus/>
<apply>
<plus/>
<ci>z_c</ci>
<ci>n</ci>
</apply>
</apply>
<ci>delta</ci>
<ci>mu</ci>
</apply>
<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>k_52_temp</ci>
<apply>
<times/>
<ci>k0_52</ci>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<plus/>
<ci>z_c</ci>
<ci>n</ci>
</apply>
<ci>delta</ci>
<ci>mu</ci>
</apply>
<cn cellml:units="dimensionless">2</cn>
</apply>
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</apply>
<apply>
<eq/>
<ci>k_56</ci>
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<times/>
<ci>k0_56</ci>
<apply>
<exp/>
<apply>
<divide/>
<apply>
<times/>
<apply>
<minus/>
<ci>n</ci>
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<ci>z_Na</ci>
<ci>alpha_pp</ci>
<ci>mu</ci>
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<cn cellml:units="dimensionless">2</cn>
</apply>
</apply>
</apply>
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<apply>
<eq/>
<ci>k_65</ci>
<apply>
<times/>
<ci>k0_65</ci>
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<power/>
<ci>Na_i</ci>
<ci>n</ci>
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<exp/>
<apply>
<divide/>
<apply>
<times/>
<ci>n</ci>
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<ci>alpha_pp</ci>
<ci>mu</ci>
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<ci>k_61</ci>
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<times/>
<ci>k0_61</ci>
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<divide/>
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<times/>
<ci>z_c</ci>
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<times/>
<ci>k0_16</ci>
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<exp/>
<apply>
<divide/>
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<times/>
<apply>
<minus/>
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<ci>mu</ci>
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<cn cellml:units="dimensionless">2</cn>
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<eq/>
<ci>k_52</ci>
<apply>
<divide/>
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<times/>
<ci>k0_12</ci>
<ci>k_25</ci>
<ci>k0_56</ci>
<ci>k0_61</ci>
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<apply>
<times/>
<ci>k0_21</ci>
<ci>k0_16</ci>
<ci>k0_65</ci>
</apply>
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<apply>
<eq/>
<ci>k0_54_temp</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>k0_23</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_52</ci>
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<apply>
<times/>
<ci>k_43</ci>
<ci>k_32</ci>
<ci>k_25</ci>
</apply>
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</apply>
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<component name="kinetic_equations">
<variable cmeta:id="kinetic_equations.k_12" name="k_12" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_21" name="k_21" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_23" name="k_23" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_32" name="k_32" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_34" name="k_34" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_43" name="k_43" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_45" name="k_45" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_54" name="k_54" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_25" name="k_25" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_52" name="k_52" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_56" name="k_56" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_65" name="k_65" public_interface="in" units="per_second"/>
<variable cmeta:id="kinetic_equations.k_61" name="k_61" public_interface="in" units="per_second"/>
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<variable cmeta:id="kinetic_equations.C_6_temp" initial_value="3.06798e-9" name="C_6_temp" public_interface="out" units="umol"/>
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<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<apply>
<diff/>
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<ci>time</ci>
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<minus/>
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<plus/>
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<times/>
<ci>k_21</ci>
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<times/>
<ci>k_61</ci>
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<apply>
<times/>
<apply>
<plus/>
<ci>k_12</ci>
<ci>k_16</ci>
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<variable cmeta:id="king_altman_states.k_21" name="k_21" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_23" name="k_23" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_32" name="k_32" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_34" name="k_34" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_43" name="k_43" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_45" name="k_45" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_54" name="k_54" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_25" name="k_25" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_52" name="k_52" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_56" name="k_56" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_65" name="k_65" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_61" name="k_61" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.k_16" name="k_16" public_interface="in" units="per_second"/>
<variable cmeta:id="king_altman_states.C1_sum" name="C1_sum" units="per_second5"/>
<variable cmeta:id="king_altman_states.C2_sum" name="C2_sum" units="per_second5"/>
<variable cmeta:id="king_altman_states.C3_sum" name="C3_sum" units="per_second5"/>
<variable cmeta:id="king_altman_states.C4_sum" name="C4_sum" units="per_second5"/>
<variable cmeta:id="king_altman_states.C5_sum" name="C5_sum" units="per_second5"/>
<variable cmeta:id="king_altman_states.C6_sum" name="C6_sum" units="per_second5"/>
<variable cmeta:id="king_altman_states.C_sum" name="C_sum" units="per_second5"/>
<variable cmeta:id="king_altman_states.C_T" name="C_T" public_interface="in" units="umol"/>
<variable cmeta:id="king_altman_states.C1" name="C1" public_interface="out" units="umol"/>
<variable cmeta:id="king_altman_states.C2" name="C2" public_interface="out" units="umol"/>
<variable cmeta:id="king_altman_states.C3" name="C3" public_interface="out" units="umol"/>
<variable cmeta:id="king_altman_states.C4" name="C4" public_interface="out" units="umol"/>
<variable cmeta:id="king_altman_states.C5" name="C5" public_interface="out" units="umol"/>
<variable cmeta:id="king_altman_states.C6" name="C6" public_interface="out" units="umol"/>
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<ci>C5_sum</ci>
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<ci>k_23</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_56</ci>
</apply>
<apply>
<times/>
<ci>k_16</ci>
<ci>k_21</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_56</ci>
</apply>
<apply>
<times/>
<ci>k_16</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_45</ci>
<ci>k_56</ci>
</apply>
<apply>
<times/>
<ci>k_12</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_45</ci>
<ci>k_56</ci>
</apply>
<apply>
<times/>
<ci>k_16</ci>
<ci>k_21</ci>
<ci>k_32</ci>
<ci>k_45</ci>
<ci>k_56</ci>
</apply>
<apply>
<times/>
<ci>k_16</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_56</ci>
</apply>
<apply>
<times/>
<ci>k_12</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_56</ci>
</apply>
<apply>
<times/>
<ci>k_16</ci>
<ci>k_21</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_56</ci>
</apply>
<apply>
<times/>
<ci>k_16</ci>
<ci>k_21</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_54</ci>
</apply>
<apply>
<times/>
<ci>k_16</ci>
<ci>k_21</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_52</ci>
</apply>
<apply>
<times/>
<ci>k_16</ci>
<ci>k_21</ci>
<ci>k_32</ci>
<ci>k_45</ci>
<ci>k_52</ci>
</apply>
<apply>
<times/>
<ci>k_16</ci>
<ci>k_21</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_52</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>C_sum</ci>
<apply>
<plus/>
<ci>C1_sum</ci>
<ci>C2_sum</ci>
<ci>C3_sum</ci>
<ci>C4_sum</ci>
<ci>C5_sum</ci>
<ci>C6_sum</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>C1</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>C_T</ci>
<ci>C1_sum</ci>
</apply>
<ci>C_sum</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>C2</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>C_T</ci>
<ci>C2_sum</ci>
</apply>
<ci>C_sum</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>C3</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>C_T</ci>
<ci>C3_sum</ci>
</apply>
<ci>C_sum</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>C4</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>C_T</ci>
<ci>C4_sum</ci>
</apply>
<ci>C_sum</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>C5</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>C_T</ci>
<ci>C5_sum</ci>
</apply>
<ci>C_sum</ci>
</apply>
</apply>
<apply>
<eq/>
<ci>C6</ci>
<apply>
<divide/>
<apply>
<times/>
<ci>C_T</ci>
<ci>C6_sum</ci>
</apply>
<ci>C_sum</ci>
</apply>
</apply>
</math>
</component>
<component name="NBC_current">
<variable cmeta:id="NBC_current.k_12" name="k_12" public_interface="in" units="per_second"/>
<variable cmeta:id="NBC_current.k_21" name="k_21" public_interface="in" units="per_second"/>
<variable cmeta:id="NBC_current.k_56" name="k_56" public_interface="in" units="per_second"/>
<variable cmeta:id="NBC_current.k_65" name="k_65" public_interface="in" units="per_second"/>
<variable cmeta:id="NBC_current.k_34" name="k_34" public_interface="in" units="per_second"/>
<variable cmeta:id="NBC_current.k_43" name="k_43" public_interface="in" units="per_second"/>
<variable cmeta:id="NBC_current.k_25" name="k_25" public_interface="in" units="per_second"/>
<variable cmeta:id="NBC_current.k_52" name="k_52" public_interface="in" units="per_second"/>
<variable cmeta:id="NBC_current.k_61" name="k_61" public_interface="in" units="per_second"/>
<variable cmeta:id="NBC_current.k_16" name="k_16" public_interface="in" units="per_second"/>
<variable cmeta:id="NBC_current.C_1" name="C_1" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.C_2" name="C_2" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.C_3" name="C_3" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.C_4" name="C_4" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.C_5" name="C_5" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.C_6" name="C_6" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.C1" name="C1" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.C2" name="C2" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.C3" name="C3" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.C4" name="C4" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.C5" name="C5" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.C6" name="C6" public_interface="in" units="umol"/>
<variable cmeta:id="NBC_current.delta" name="delta" public_interface="in" units="dimensionless"/>
<variable cmeta:id="NBC_current.alpha_p" name="alpha_p" public_interface="in" units="dimensionless"/>
<variable cmeta:id="NBC_current.alpha_pp" name="alpha_pp" public_interface="in" units="dimensionless"/>
<variable cmeta:id="NBC_current.n" name="n" public_interface="in" units="dimensionless"/>
<variable cmeta:id="NBC_current.z_c" name="z_c" public_interface="in" units="dimensionless"/>
<variable cmeta:id="NBC_current.z_Na" name="z_Na" public_interface="in" units="dimensionless"/>
<variable cmeta:id="NBC_current.F" name="F" public_interface="in" units="C_per_mol"/>
<variable cmeta:id="NBC_current.I_NaGl_pSS" name="I_NaGl_pSS" public_interface="out" units="uA"/>
<variable cmeta:id="NBC_current.I_NaGl_SS" name="I_NaGl_SS" public_interface="out" units="uA"/>
<variable cmeta:id="NBC_current.J_Na" name="J_Na" public_interface="out" units="umol_per_second"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>I_NaGl_pSS</ci>
<apply>
<times/>
<apply>
<minus/>
<ci>F</ci>
</apply>
<apply>
<plus/>
<apply>
<times/>
<ci>n</ci>
<ci>z_Na</ci>
<ci>alpha_p</ci>
<apply>
<minus/>
<apply>
<times/>
<ci>k_12</ci>
<ci>C_1</ci>
</apply>
<apply>
<times/>
<ci>k_21</ci>
<ci>C_2</ci>
</apply>
</apply>
</apply>
<apply>
<times/>
<ci>z_c</ci>
<ci>delta</ci>
<apply>
<minus/>
<apply>
<times/>
<ci>k_16</ci>
<ci>C_1</ci>
</apply>
<apply>
<times/>
<ci>k_61</ci>
<ci>C_6</ci>
</apply>
</apply>
</apply>
<apply>
<times/>
<ci>n</ci>
<ci>z_Na</ci>
<ci>alpha_pp</ci>
<apply>
<minus/>
<apply>
<times/>
<ci>k_56</ci>
<ci>C_5</ci>
</apply>
<apply>
<times/>
<ci>k_65</ci>
<ci>C_6</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>I_NaGl_SS</ci>
<apply>
<times/>
<apply>
<minus/>
<ci>F</ci>
</apply>
<apply>
<plus/>
<apply>
<times/>
<ci>z_c</ci>
<apply>
<minus/>
<apply>
<times/>
<ci>k_16</ci>
<ci>C1</ci>
</apply>
<apply>
<times/>
<ci>k_61</ci>
<ci>C6</ci>
</apply>
</apply>
</apply>
<apply>
<times/>
<apply>
<plus/>
<ci>z_c</ci>
<apply>
<times/>
<ci>z_Na</ci>
<ci>n</ci>
</apply>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>k_25</ci>
<ci>C2</ci>
</apply>
<apply>
<times/>
<ci>k_52</ci>
<ci>C5</ci>
</apply>
</apply>
</apply>
<apply>
<times/>
<apply>
<plus/>
<ci>z_c</ci>
<apply>
<times/>
<ci>z_Na</ci>
<ci>n</ci>
</apply>
</apply>
<apply>
<minus/>
<apply>
<times/>
<ci>k_34</ci>
<ci>C3</ci>
</apply>
<apply>
<times/>
<ci>k_43</ci>
<ci>C4</ci>
</apply>
</apply>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>J_Na</ci>
<apply>
<minus/>
<apply>
<plus/>
<apply>
<minus/>
<apply>
<times/>
<ci>k_25</ci>
<ci>C2</ci>
</apply>
<apply>
<times/>
<ci>k_52</ci>
<ci>C5</ci>
</apply>
</apply>
<apply>
<times/>
<ci>k_34</ci>
<ci>C3</ci>
</apply>
</apply>
<apply>
<times/>
<ci>k_43</ci>
<ci>C4</ci>
</apply>
</apply>
</apply>
</math>
</component>
<component name="phenomonological_constants">
<variable cmeta:id="phenomonological_constants.epsilon" name="epsilon" public_interface="out" units="per_second"/>
<variable cmeta:id="phenomonological_constants.lambda" name="lambda" public_interface="out" units="per_M2_per_second5"/>
<variable cmeta:id="phenomonological_constants.chi" name="chi" public_interface="out" units="M"/>
<variable cmeta:id="phenomonological_constants.alpha" name="alpha" public_interface="out" units="M2"/>
<variable cmeta:id="phenomonological_constants.beta" name="beta" public_interface="out" units="M"/>
<variable cmeta:id="phenomonological_constants.gamma" name="gamma" public_interface="out" units="M2_per_second"/>
<variable cmeta:id="phenomonological_constants.phi" name="phi" public_interface="out" units="M_per_second"/>
<variable cmeta:id="phenomonological_constants.k_12" name="k_12" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_21" name="k_21" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_23" name="k_23" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_32" name="k_32" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_34" name="k_34" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_43" name="k_43" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_45" name="k_45" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_54" name="k_54" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_25" name="k_25" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_52" name="k_52" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_56" name="k_56" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_65" name="k_65" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_61" name="k_61" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.k_16" name="k_16" public_interface="in" units="per_second"/>
<variable cmeta:id="phenomonological_constants.ks_12" name="ks_12" public_interface="in" units="per_M_per_second"/>
<variable cmeta:id="phenomonological_constants.k0_23" name="k0_23" public_interface="in" units="per_M_per_second"/>
<variable cmeta:id="phenomonological_constants.C_T" name="C_T" public_interface="in" units="umol"/>
<variable cmeta:id="phenomonological_constants.F" name="F" public_interface="in" units="C_per_mol"/>
<variable cmeta:id="phenomonological_constants.Na_o" name="Na_o" public_interface="in" units="M"/>
<variable cmeta:id="phenomonological_constants.glucose_o" name="glucose_o" public_interface="in" units="M"/>
<variable cmeta:id="phenomonological_constants.Imax_Na" name="Imax_Na" public_interface="out" units="uA"/>
<variable cmeta:id="phenomonological_constants.Imax_gluc" name="Imax_gluc" public_interface="out" units="uA"/>
<variable cmeta:id="phenomonological_constants.Khalf_Na" name="Khalf_Na" public_interface="out" units="M"/>
<variable cmeta:id="phenomonological_constants.Khalf_gluc" name="Khalf_gluc" public_interface="out" units="M"/>
<math xmlns="http://www.w3.org/1998/Math/MathML">
<apply>
<eq/>
<ci>lambda</ci>
<apply>
<plus/>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_43</ci>
<ci>k_54</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_34</ci>
<ci>k_54</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_45</ci>
<ci>k_52</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_43</ci>
<ci>k_52</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_34</ci>
<ci>k_52</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_45</ci>
<ci>k_56</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_43</ci>
<ci>k_56</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_34</ci>
<ci>k_56</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_43</ci>
<ci>k_54</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_34</ci>
<ci>k_54</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_45</ci>
<ci>k_52</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_43</ci>
<ci>k_52</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_34</ci>
<ci>k_52</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k0_23</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_56</ci>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>chi</ci>
<apply>
<times/>
<apply>
<divide/>
<cn cellml:units="dimensionless">1</cn>
<ci>lambda</ci>
</apply>
<apply>
<plus/>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_54</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_43</ci>
<ci>k_54</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_34</ci>
<ci>k_54</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_54</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_52</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_32</ci>
<ci>k_45</ci>
<ci>k_52</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_52</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_45</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_65</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_56</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_32</ci>
<ci>k_45</ci>
<ci>k_56</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_56</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_54</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_43</ci>
<ci>k_54</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_34</ci>
<ci>k_54</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_54</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_52</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_32</ci>
<ci>k_45</ci>
<ci>k_52</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_52</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_45</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_61</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_34</ci>
<ci>k_45</ci>
<ci>k_56</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_45</ci>
<ci>k_56</ci>
</apply>
<apply>
<times/>
<ci>ks_12</ci>
<ci>k_25</ci>
<ci>k_32</ci>
<ci>k_43</ci>
<ci>k_56</ci>
</apply>
</apply>
</apply>
</apply>
<apply>
<eq/>
<ci>beta</ci>
<apply>
<times/>
<apply>
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<documentation xmlns="http://cellml.org/tmp-documentation">
<article xmlns="http://cellml.org/tmp-documentation">
<articleinfo xmlns="http://cellml.org/tmp-documentation">
<title xmlns="http://cellml.org/tmp-documentation">Biophysical Characteristics of the Pig Kidney Na/Glucose Cotransporter SGLT2 Reveal a Common Mechanism for SGLT1 and SGLT2</title>
<author xmlns="http://cellml.org/tmp-documentation">
<firstname xmlns="http://cellml.org/tmp-documentation">Jonna</firstname>
<surname xmlns="http://cellml.org/tmp-documentation">Terkildsen</surname>
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<title xmlns="http://cellml.org/tmp-documentation">Model Status</title>
<para xmlns="http://cellml.org/tmp-documentation">This CellML model has been curated, the units are consistent and the model runs in COR and PCEnv. This model recreates the published results.</para>
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<para xmlns="http://cellml.org/tmp-documentation">Abstract: The Na-dependent, low affinity glucose transporter SGLT2 cloned from pig kidney is 76% identical (at the amino acid level) to its high affinity homologue SGLT1. Using two-microelectrode voltage clamp, we have characterized the presteady-state and steady-state kinetics of SGLT2 expressed in Xenopus oocytes. The kinetic properties of the steady-state sugar-evoked currents as a function of external Na and a-methyl-D-glucopyranoside (aMG) concentrations were consistent with an ordered, simultaneous transport model in which Na binds first. Na binding was voltage-dependent and saturated with hyperpolarizing voltages. Phlorizin was a potent inhibitor of the sugar-evoked currents (Ki ~10 mM) and blocked an inward Na current in the absence of sugar. SGLT2 exhibited Na-dependent presteadystate currents with time constants 3-7 ms. Charge movements were described by Boltzmann relations with apparent valence ~1 and maximal charge transfer ~11 nC, and were reduced by the addition of sugar or phlorizin. The differences between SGLT1 and SGLT2 were that (i) the apparent affinity constant (K0.5) for aMG (~3 mM) was an order of magnitude higher for SGLT2; (ii) SGLT2 excluded galactose, suggesting discrete sugar binding; (iii) K0.5 for Na was lower in SGLT2; and (iv) the Hill coefficient for Na was 1 for SGLT2 but 2 for SGLT1. Simulations of the six-state kinetic model previously proposed for SGLT1 indicated that many of the kinetic properties observed in SGLT2 are expected by simply reducing the Na/glucose coupling from 2 to 1.</para>
<para xmlns="http://cellml.org/tmp-documentation">The original paper reference is cited below:</para>
<para xmlns="http://cellml.org/tmp-documentation">
Biophysical Characteristics of the Pig Kidney Na/Glucose Cotransporter SGLT2 Reveal a Common Mechanism for SGLT1 and SGLT2, M. Mackenzie, D.D.F. Loo, M. Panayotova-Heiermann, and E. M. Wright, 1996,
<emphasis xmlns="http://cellml.org/tmp-documentation">Journal of Biological Chemistry</emphasis>
, 271, 32678-32683.
<ulink xmlns="http://cellml.org/tmp-documentation" url="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=pubmed&cmd=Retrieve&dopt=AbstractPlus&list_uids=8955098&query_hl=1&itool=pubmed_docsum">PubMed ID: 8955098</ulink>
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<caption xmlns="http://cellml.org/tmp-documentation">Schematic diagram of the Mackenzie et al 1996 SGLT2 model. C' represents the external-facing carrier. C'' represents the internal-facing carrier.</caption>
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<dcterms:abstract xmlns:dcterms="http://purl.org/dc/terms/">The Na-dependent, low affinity glucose transporter SGLT2 cloned from pig kidney is 76% identical (at the amino acid level) to its high affinity homologue SGLT1. Using two-microelectrode voltage clamp, we have characterized the presteady-state and steady-state kinetics of SGLT2 expressed in Xenopus oocytes. The kinetic properties of the steady-state sugar-evoked currents as a function of external Na and a-methyl-D-glucopyranoside (aMG) concentrations were consistent with an ordered, simultaneous transport model in which Na binds first. Na binding was voltage-dependent and saturated with hyperpolarizing voltages. Phlorizin was a potent inhibitor of the sugar-evoked currents (Ki ~10 mM) and blocked an inward Na current in the absence of sugar. SGLT2 exhibited Na-dependent presteadystate currents with time constants 3-7 ms. Charge movements were described by Boltzmann relations with apparent valence ~1 and maximal charge transfer ~11 nC, and were reduced by the addition of sugar or phlorizin. The differences between SGLT1 and SGLT2 were that (i) the apparent affinity constant (K0.5) for aMG (~3 mM) was an order of magnitude higher for SGLT2; (ii) SGLT2 excluded galactose, suggesting discrete sugar binding; (iii) K0.5 for Na was lower in SGLT2; and (iv) the Hill coefficient for Na was 1 for SGLT2 but 2 for SGLT1. Simulations of the six-state kinetic model previously proposed for SGLT1 indicated that many of the kinetic properties observed in SGLT2 are expected by simply reducing the Na/glucose coupling from 2 to 1.</dcterms:abstract>
<vCard:FN xmlns:vCard="http://www.w3.org/2001/vcard-rdf/3.0#">Mariana Panayotova-Heiermann</vCard:FN>
<semsim:unknown xmlns:semsim="http://www.bhi.washington.edu/SemSim#">The Na-dependent, low affinity glucose transporter SGLT2 cloned from pig kidney is 76% identical (at the amino acid level) to its high affinity homologue SGLT1. Using two-microelectrode voltage clamp, we have characterized the presteady-state and steady-state kinetics of SGLT2 expressed in Xenopus oocytes. The kinetic properties of the steady-state sugar-evoked currents as a function of external Na and a-methyl-D-glucopyranoside (aMG) concentrations were consistent with an ordered, simultaneous transport model in which Na binds first. Na binding was voltage-dependent and saturated with hyperpolarizing voltages. Phlorizin was a potent inhibitor of the sugar-evoked currents (Ki ~10 mM) and blocked an inward Na current in the absence of sugar. SGLT2 exhibited Na-dependent presteadystate currents with time constants 3-7 ms. Charge movements were described by Boltzmann relations with apparent valence ~1 and maximal charge transfer ~11 nC, and were reduced by the addition of sugar or phlorizin. The differences between SGLT1 and SGLT2 were that (i) the apparent affinity constant (K0.5) for aMG (~3 mM) was an order of magnitude higher for SGLT2; (ii) SGLT2 excluded galactose, suggesting discrete sugar binding; (iii) K0.5 for Na was lower in SGLT2; and (iv) the Hill coefficient for Na was 1 for SGLT2 but 2 for SGLT1. Simulations of the six-state kinetic model previously proposed for SGLT1 indicated that many of the kinetic properties observed in SGLT2 are expected by simply reducing the Na/glucose coupling from 2 to 1.</semsim:unknown>
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